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The Freshwater Fishes of Western Borneo 
(Kalimantan Barat, Indonesia) 






The Freshwater Fishes of Western Borneo 
(KaUmantan Barat, Indonesia) 

By 
Tyson R. Roberts 



Published by 
California Academy of Sciences 



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^i^ 




San Francisco 
1989 

Memoirs of the California Academy of Sciences Number 14 




SCIENTIFIC PUBLICATIONS COMMITTEE: 

Daphne Fautin, Scientific Editor 

Frank Almeda 

Luis Baptista 

Wojciech Pulawski 

Frank Talbot 

Sheridan Warrick 

Date submitted for publication: November 29, 1984 
Date accepted for publication: July 23, 1985 

Published by the California Academy of Sciences. 

Copyright ici 1989 by California Academy of Sciences. All rights reserved. No part of this publication 
may be reproduced or transmitted in any form or by any means, electronic or mechanical, included 
photocopying, recording, or any information storage or retrieval system, without permission in writing 
from the publisher. 

Library of Congress Catalog Card Number 88-70981 
ISBN 0-940228-21-1 

Cover Illustration: Acantophthalinus superbus 



The Freshwater Fishes of Western Borneo 
(KaUmantan Barat, Indonesia) 

By 
Tyson R. Roberts 

Department of [chlhyolog): California Academy of Sciences. San Francisco. California 94118 

Table of Contents 

Abstract 1 

Introduction 1 

Acknowledgments 3 

History of East Indian ichthyology, with particular reference to freshwater fishes of Borneo 4 

Reproductive adaptations in freshwater fishes of Borneo 10 

Collecting localities of the 1976 Kapuas ichthyological survey 13 

Systematic Account 20 

Dasyatidae 22 

Himantura 22 

Himantura signifer 23 

Osteoglossidae 23 

Scleropages 23 

Scleropages formosus 23 

Notopteridae 24 

Notopteriis 24 

Notopterus borneensis 24 

Clupeidae 24 

Clupeichthys 24 

Clupeichthys bleekeri 24 

Clupeoides - 25 

Clitpeoides hypselosoma 25 

Engraulidae 25 

Lycolhnssa 26 

Lycothrissa crocodilus 26 

Setipinna 26 

Setipinna melanochu\ 26 

Sundasalangidae 27 

Sitndasalanx 27 

Sundasalanx microps 27 

Sundasalanx sp. undet 27 

Cyprinidae 27 

Albulichlhys 27 

Albulichthys albuloides 28 

Amblyrhyncluchthys 28 

Amblyrltynchichthys Iruncatus 29 

Balantiocheilos 29 

Balantiocheilos melanopterus 29 

Barbichthys 29 

Barbichthys laevis 30 

Barilius 30 

Barilius borneensis new species 30 

Chela 31 

Chela maassi 31 

Cosinochihis 31 

Cosmochiliis falcifer 31 



Crossocheilus 32 

Crossocheilns cobilis - 32 

Crossocheilus oblongus 32 

Crossocheilus sp. undet 32 

Cyclocheilichthys - 33 

Cyclocheilichthys apogon - 35 

Cyclocheilichthys annatus 35 

Cyclocheilichthys enoplos ~ 35 

Cyclocheilichthys heleronema 36 

Cyclocheilichthys janthochir 36 

Cyclocheilichthys niicrolepis 36 

Cyclocheilichthys repasson 36 

Dangila 36 

Dangila cuvieri 38 

Dcingila fasciata 38 

Dangila festiva 38 

Dangila lineata 38 

Dangila ocellala 38 

Eirmotus 38 

Eirmotus octozona - 38 

Epalzeorhynchos - 38 

Epalzeorhynchos kalopterum 39 

Garra 40 

Garra horneensis 40 

Hampala 40 

Hainpala himaculata 40 

Hampala macrolepidota - 40 

Kaliinantania 40 

Kalimantania lawak 40 

Lcpiobarhus 41 

Leptobarbus hoeveuii 41 

Lepioharbus melanopterus - 41 

Lobocheilos 41 

Lobocheilos bo 41 

Lobocheilos hispidiis 41 

Luciosoma 42 

Luciosoma setigeruni - 43 

Luciosoma spilopleiira 43 

Luciosoma trinema 44 

Macrochirichthys 44 

Macrochinchthys inacrochirus 44 

Morulius 45 

Morulius chrysophekadion 45 

Mystacoleucus 45 

Mystacoleiicus margmatus 45 

Osteochilus 45 

Osteochilus borneensis 47 

Osteochilus enneaporus 47 

Osteochilus hasselti 48 

Osteochilus intermedins 48 

Osteochilus kahajanensis 49 

Osteochilus kappeiui - 49 

Osteochilus niclanopleurus 50 

Osteochilus nucrocephalus 52 

Osteochilus pleurotaenia 52 

Osteochilus schlegeli 52 

Osteochilus spilurus 53 

Osteochilus triporos 54 

Osteochilus waandersii 54 

O.xygaster 55 

Oxygaster hypophiiialnnis 55 



Oxygaster oxygaster 56 

Oxygaster oxygastroides 57 

Paracwssochilus 57 

Paracrossochilus acenis 57 

Oaracrossochihts vitiatis 57 

Pectenocypris 57 

Pectenocypris balaena new species 58 

Puntioplites 59 

Pimtioplites bulit 60 

Puntioplites waandersi 60 

Puntius 60 

Puntius anchisponis 60 

Puntius binotatus 60 

Puntius bramoides 61 

Puntius collingwoodii 62 

Puntius endecanalis new species 62 

Puntius eugrammus 64 

Puntius everetti 64 

Puntius lateristriga 65 

Puntius lineatus 65 

Puntius pentazona 65 

Puntius rhotnboocellatus 65 

Puntius schwanenfeldii 66 

Rasbora 66 

Rasbora agilis 66 

Rasbora argywtacnia 67 

Rasbora axebvdi 67 

Rasbora bankancnsis 70 

Rasbora beauforii sp. inq 71 

Rasbora borneensis 71 

Rasbora brittani 72 

Rasbora caudiniaculata 72 

Rasbora cephalotaenia 72 

Rasbora dorsioccllata 73 

Rasbora dusonensis 73 

Rasbora einthovenii 73 

Rasbora elegans 73 

Rasbora ennealepis new species 74 

Rasbora kalochroma 74 

Rasbora myersi 74 

Rasbora paiiciperforata 74 

Rasbora sarawakensis 75 

Rasbora subtilis new species 75 

Rasbora trilineata 76 

Rasbora voizi 77 

Rasbora sp. undet 77 

Rasborichlhys 77 

Rasbortchthys hdfrichii 78 

Rohteicluhys 78 

Rohteichlhys microlepis 78 

Schismatorhynchos 79 

Schismaiorhynchos heterorhynchos 79 

Thryssocypris 79 

Thryssocypris smaragdinus 79 

Thynnichthys 80 

Thynnichtliys polylcpis 80 

Thynnichthys thynnoides 80 

Tor ' __ 80 

TorcC. tanibra 80 

Tor tambroides 81 

Gyrinocheilidae 81 



Gyrinocheilus 81 

Gynnocheilus pustulosus 81 

Homalopteridae 81 

Gastroi>iy:o)i 85 

Gaslroniyzon conlractus 86 

Gastwmyzon fasciatus 86 

Gastromyzon Icpidogaster 87 

Gastromyzon ridens 88 

Hoinaloptera 88 

Homaloptera nebulosa 88 

llomaloptera ogllviei 89 

Homaloptera ophiolepis 89 

Homaloptera orthogoniata 89 

Homaloptera cf stephensoni 90 

Homaloptera tweediei 90 

Homaloptera zollingeri 91 

Hypergaslromyzon new genus 91 

Hypergasiromyzon humilis new species 92 

Neogastromyzon 93 

Neogastromyzon nieuwenhuisii 93 

Neohomaloptera 93 

Neohomalopterajohorensis 94 

Cobitidae 94 

Acanthopsoides 94 

Acanthopsoides gracilis 94 

Acautophlhalmus 95 

Acaiuophthalmus angnillaris 95 

Acautophlhalmus oblongus 97 

Acantophihatmus semiclndiis 98 

Acantophlhatmus shclfordi 98 

Acantophlhalmus sitperhus new species 98 

Acanlopsis 99 

Acantopsis choirorhynchos 99 

Barbucca new genus 100 

Barbucca diabolica new species 100 

Botia 101 

Botia hymenophysa 101 

Bolia macracantha 102 

Bona reversa new species 102 

Elloposloma 103 

Ellopostoma tnegalomycter 103 

Lepidocephalichihys 103 

Lepidocephalichthys hasselti 103 

Lepidocephaluhthys lorcutzi 104 

Lepidocephalichthys pristes new species 105 

Lepidocephaliis 106 

Lcpidocephaliis macrochir 106 

Lepidocephaliis spectrum new species - 106 

Nemacheilus 106 

Nemacheilus kapuasensis 107 

Nemacheilus lactogencus new species 107 

Nemacheilus cf longipectoralis 108 

Nemacheilus maculiceps new species 108 

Nemacheilus saravacensis 108 

Nemacheilus selangoricus 109 

Vaillantella 109 

Vaillantella euepiptera 109 

\ aillantella maassi 109 

Ariidae 110 

Anus 110 



Anus melanochir 1 10 

Arius stormi 110 

Hemipimelodus 1 10 

Hemipimelodus borneensis 1 11 

Bagndae 1 11 

Bagrichthys 11 1 

Bagrichthys hypselopterus 1 12 

Bagrichthys macwptenis 1 14 

Bagrichthys nucranodus new species 115 

Bagroides 115 

Bagroides melapterus 1 15 

Leiocassis 1 16 

Leiocassis arniatus 1 17 

Leiocassis micropogon 117 

Leiocassis myersi new species 1 17 

Leiocassis vaillanli 120 

Leiocassis sp. undet 120 

Mystus 120 

Mystus micracanthus 120 

Mystus nemurus 121 

Mystus nigriceps 122 

Mystus olyroides new species 124 

Mystus wolffii 126 

Mystus wyckii 126 

Pelteobagrus 126 

Peheobagrus ornatus 126 

Clariidae 127 

Clarias 127 

Glorias cf batrachus 127 

Clarias leiacanthus 127 

Clarias meladerma 127 

Clarias meuhofi 128 

Clarias teysmanni 128 

Encheloclarias 128 

Encheloclarias tapeinopterus 128 

Schilbeidae 128 

Pseudeutropius 128 

Pseiideiitropius brachypopterus 129 

Pseudeutropius moolenburghae 129 

Pangasiidae 130 

Laides 131 

Laides hexanema 131 

Pangasius 131 

Pangasiits humeralis new species 131 

Pangasius hthostoma new species 132 

Pangasius micronema 132 

Pangasius nasutus 133 

Pangasius polyuranodon 133 

Sisoridae 133 

Bagarius 133 

Bagarius yarrelli 133 

Glyptothorax 134 

Glyptothorax major 134 

Glyptothorax platypogon 135 

Glyptothorax platypogonoides 136 

Akysidae 136 

Acrochordonichthys 137 

Acrochordonichthys chamaleon 137 

Acrochordonichthys cf. melanogaster 138 

Akysis 138 



Akysis p.sciidobagarius new species 138 

Breitensteinia 140 

Brcitcnstcinia cf insignis 140 

Parakysidae new tamily 141 

Parakysis 142 

Purakysis anomalopteryx new species 142 

Chacidae 143 

Chaca _ 143 

Chaca bankanemis. 143 

Siluridae 143 

Bclodontichihys 144 

BelocJonlichtliys duicina 144 

Ceratoglanis 144 

Ceraioglanis sclcroneiiia 144 

Hemisilurm 145 

Hemisilurus hetewrhynchos 145 

Hemisilums moolenburghi 145 

Kryptoplcnis 145 

Kryptopterus apogon 145 

Kryplopterus bicirrhis 147 

Kryptopterus cryptoplerus 147 

Kryplopterus lais 147 

Kryplopterus limpok _ 147 

Kryplopterus macrocephalus 147 

Kryplopterus nucroneina 149 

Kryplopterus nunor new species 149 

Kryplopterus schilbeules 150 

Kryplopterus sp. undet 150 

Ompok 150 

Onipok eugeneiaius 150 

Ompok hypophthalinus 150 

Ompok subcinus 151 

Ompok weberi 151 

Siluruiilhys 151 

Silunehlhys hasselli 151 

Sihiriehthys phaiosoma 151 

Silwichthys sanguineus new species 151 

W'allago ' 151 

IS'allago leerii ___ _ _ _ 152 

Belonidae 152 

Xeneniodon 152 

Xeneniodon canciloides 153 

Hemiramphidae 153 

Der))utgenys __ _.._ _ _ __ _ 154 

Dermogenys cT piisilhis 154 

Heniirhumpliodon 154 

Ilemirhainphihlon phaiosoma 155 

Uemnhamphodoii pogonognalhus 155 

Uemniiainphodon sp. undet 158 

Syngnaihidae 158 

Doryichlhys 158 

Doryiehthys boaja 159 

Doryichlhys deokhatoides 159 

Doryiehthys heterosoma 159 

Doryiehthys mariensii 159 

Chandidae ' 159 

Gymnoehanda 159 

Gyninochanda hhimenlosa 160 

Paradoxodacna new genus - 1 60 

Farado.xodaena piratiea new species 160 



Parambassis _ 161 

Parambassis apogonoides 161 

Parambassis macwlepis 162 

Parambassis woljfii 162 

Nandidae 164 

Nandus 164 

Nandus nebulosus 164 

Datnioididae 164 

Dalmoides 164 

Datnioides microlepis 164 

Datmoides quadrifasciatus 165 

Pristolepidae 165 

Pristolepis 165 

Pnstolepis fasciata 165 

Toxotidae 165 

Toxotes 165 

Toxotes microlepis 165 

Polynemidae 166 

Polynemus 166 

Polynemm macrophthalmus 166 

Polynemus muliifilis 166 

Eleotridae 166 

Eleolris 166 

Eleotris melanosoma 166 

Oxyeleotris 166 

Oxyeleotris marmorata 166 

Oxyeleotris itrophthalmus 167 

Oxyeleotris urophthalmoides 167 

Gobiidae 167 

Brachygobius 167 

Brachygobius doriae 167 

Brachygobius xanthomelas 167 

Calamiana 168 

Calamiana sp. undet 168 

Mugilogobius 168 

Mugilogobius sp. undet 168 

Mugilogobius sp. undet 168 

Pseudogobiopsis - - 168 

Pseudogobiopsis jurongensis 168 

Pseudogobius 169 

Pseudogobius sp. undet 169 

Stigmatogobius 169 

Sligmatogobius brocki 169 

Stigmatogobius sp. undet 169 

Channidae 169 

Channa 169 

Channa bankanensis 169 

Channa lucius 170 

Channa mandioides 170 

Channa melanoptera 170 

Channa melasoma 170 

Channa micropeltes 170 

Channa orientalis 170 

Channa pleurophthalmus 170 

Channa striata 170 

Anabantidae 171 

Anabas 171 

Anabas testudineus 171 

Belontiidae 171 

Belontia 171 



Beloniia hasselti 171 

Betta 171 

Betta anabatoides 172 

Betta dimidiata new species 172 

Betta piignax 173 

Betta cf taetuata 173 

Parosphwmenus 174 

Pawsphromcnus deissueri 174 

Parosphronwnus parvulus 174 

Sphaerichthys 175 

Sphaerichthys osphromenoides 176 

Sphaerichthys vaillanti 177 

Trichogaster 177 

Tnchogaster leerii 177 

Trichogaster trichopterus 177 

Osphronemidae 177 

Osphronennts 177 

Osphronemiis goramy 177 

Helostomatidae 177 

Helostoina 177 

Helostoma tenvuinckii 178 

Luciocephalidae 178 

Luciocephahis 178 

Liiciocephahis piilcher 178 

Mastacembelidae 178 

Macrogmithus _ 178 

Macrognathus aciileatus 180 

Mastacembehts 180 

Mastacembehis erythrotaenia 181 

Mastacembehts inacidatus 181 

Maslaccnibeliis notophthalmus new species 181 

Mastaccmbeliis itnicolor 182 

Synbranchidae 182 

Monopierus 183 

Monopterus albus 183 

Soleidae 183 

Achiroides 183 

Achiroides leucorhyiichiis 183 

Achiroides melanorhyuchiis 183 

Achiroides sp. undet 183 

Cynoglossidae 183 

Cyiioglossiis 184 

Cynoglossiis kapiiasensis 184 

Cynogh/>ssiis waandersi 185 

Tetraodontidae 185 

Chonerhmos 186 

Choncrhinos amabilis 187 

Chonerhmos modestiis 187 

Chonerhmos ncfastus 187 

Tetraodon 188 

Tetraodon leiurus 188 

Tetraodon nigroviridis 188 

Tetraodon palembangensis 188 

Literature Cited 188 

Index to Genera and Species 195 

General Index 208 



Abstract: This monograph on (he frtshwater fishes of western Borneo or Kalimantan Barat. Indonesia, is based on the extensive collections made 
during the author's ichth>ological surve> of the Kapuas Ri\er basin in 1976. as uell as on material from earlier collections deposited in European 
museums. The ichthyofauna includes about 290 species belonging to 120 genera and 40 families. One new family of catfishes (Parakysidae), three 
ne« genera [Hyper^astwmyion in Homalopleridae. Barbucca in Cobitidae. and Paradoxodacna in Chandidae). and 25 new species (belonging in 
the families Cyprinidae. Ilomalopteridae. Cobitidae. Belontiidae. and Mastacembelidae) are described. Among the new taxa are Pectenocypris 
bataena, a small Rasbora-Wkv cyprinid with over 200 gill rakers on the first gill arch, which feeds on unicellular phytoplankton; Lepidocephatus 
spectrum, the first blind cobitid from a nonsuhterranean habitat; Hypergastromyzon humilis, a highly specialised gastromy/ontin homalopterid; and 
Paradoxodacna pirarica, a scale-eating chandid v\ith remarkable dentition. Original observations on feeding habits and reproduction are reported 
for various species. Scanning electron micrographs reveal the unciliferous quality of the lips of 7 of the 13 species of the herbivorous cyprinid genus 
Osteochilus known from western Borneo. The ant-eating habits of the endemic freshwater halfbeak genus Hemirhamphodon are reported for the 
first time, together with observations on its poorly known internal fertilization and viviparity. Oral-brooding is reported in Parambassis apogonoides, 
the first member of the percoid family Chandidae in which such parental care has been observed. 

The introduction includes general remarks on the freshwater ichthyofauna of western Borneo and its geographical distribution: a brief historical 
sketch of East Indian ichthyology, with particular reference to freshwater fishes of western Borneo: an essay on reproductive biology of freshwater 
fishes in western Borneo; and a detailed description of collecting localities of the 1976 Kapuas survey. The systematic account includes a brief 
introductory statement for each family . formal sy nonymies for almost all included genera and species, identification keys for some groups, comparative 
morphological and meristic data for many groups, detailed information on food habits and reproductive biology for some species, and geographical 
distribution for all species for which reliable information is available. The monograph is extensively illustrated with black-and-white photographs 
and line drawings. The literature cited provides a nearly complete bibliography for systematic study of Bornean freshwater fishes to the end of 1984. 



Introduction 

The ichthyological significance of western Borneo or Kali- 
mantan Barat and its largest river, the Kapuas. lies in the geo- 
graphical and hydrographic history of Sundaiand and the drain- 
age basin of the ancient Central or North Sunda River (hereafter 
referred to simply as the Sunda River). Today, with ocean levels 
only 8 m below the highest ever reached. Sundaiand is sundered 
by the shallow South China Sea into the separate lands of the 
Malay Peninsula. Sumatra. Borneo, and Java. The freshwater 
hydrographic systems of these lands are now entirely separate. 
During much of the Cenozoic and the Pleistocene Ice Age Max- 
ima, however, the oceans were 60-80 m lower than they are 
today. Under such conditions the continental land of the Malay 
Peninsula, Sumatra, Borneo, and Java was connected. During 
Pleistocene times its extensive lowlands presumably were en- 
tirely covered with tropical evergreen rain forest similar to that 
now found in much of the area. Low-lying but nearly continuous 
mountain ranges near the peripheral and southerly limits of 
Sundaiand provided the watershed for the vast Sunda River. 
The Sunda River incorporated all or nearly all of the drainage 
area of eastern and southwestern Malay Peninsula, northern 
Sumatra, western and southwestern Borneo, and northwestern 
Java. It represents the only great equatorial river system of the 
Asian continent, somewhat smaller than, but otherwise com- 
parable to, the Amazon and Congo basins. Like the equatorial 
river systems of South Amenca and Africa, the Sunda River 
basin became populated by an extremely rich and largely en- 
demic ichthyofauna. The Sunda drainage appears to have been 
an important evolutionary center for many fish groups, most 
notably the ostariophysan families Cyprinidae. Gyrinocheilidae, 
Homalopteridae. Cobitidae. Bagridae. Pangasiidae. and Siluri- 
dae, and the non-ostariophysan suborder Anabantoidea. All of 
these groups have endemic genera and more or less numerous 
endemic species in Sundaiand, and the distribution of a large 
proportion of them is today restricted to rivers that formerly 
were part of the unified Sunda drainage. The significance of 
western Borneo's Kapuas river is that it is the largest and prob- 
ably has the richest ichthyofauna of any of the modem rivers 
derived from the Sunda drainage. The Sunda drainage was last 
unified during the period 26,000 to 6,000 ybp. During this pe- 
riod a preponderance of the Sunda drainage fishes probably were 
widely or generally distributed in what is now the Malay Pen- 



insula, Sumatra, western Borneo, Java, and the shallow floor of 
the intervening South China Sea. The present high levels of 
identical freshwater fish species now inhabiting these lands pre- 
sumably is primarily due to dispersal events initiated 26,000 
ybp and interrupted 6,000 ybp. 

The richness of the Kapuas ichthyofauna was not fully ap- 
preciated until recently and even now many species probably 
remain to be discovered. Prior to 1 976 there had been only four 
major ichthyological surveys of the Kapuas. these and other 
efforts resulting in a known freshwater ichthyofauna of approx- 
imately 1 50 species (many incorrectly identified). In 1 976, spon- 
sored by the Smithsonian Tropical Research Institute. Indo- 
nesian National Research Council, and Museum Zoologicum 
Bogorense, I undertook a comprehensive ichthyological survey 
of the Kapuas resulting in collection of some 260 freshwater 
fish species. 

Subsequently I visited museums in Europe, examining and 
reidentifying significant portions of all but one of the important 
Kapuas collections previously made. These studies, reported 
herein, have resulted in a presently known freshwater ichthyo- 
fauna of western Borneo of about 290 species, a large proportion 
known only from within the hydrographic limits of the Sunda 
River drainage. 

Excluded from the present account are exotic fish species 
introduced into western Borneo for aquaculture or other rea- 
sons. In Kalimantan Barat. in 1976. I heard that a number of 
exotic species had been introduced as pondfish. but have no 
documentation of this. Iwamatsu et al. ( 1 982) reported the rice- 
fish Oryziasjavaniciis (Bleeker. 1 854) from Pontianak. The hab- 
itat is described simply as a "stream," width 1 m, depth 20-30 
cm at Pontianak. and the specimens were collected in 1978. 
Oryzias has not been reported previously from the Kapuas or 
anywhere else in Borneo. Its occurrence in Pontianak presum- 
ably is due to introduction for mosquito control. 

Some species found in western Borneo do extend into adjacent 
areas, especially southern and eastern Borneo and Thailand, but 
the Chao Phrya and Mekong ichthyofaunas are largely distinct 
from the Sunda ichthyofauna. Very few of the Sunda species 
occur in India, almost none in China. Of the approximately 40 
or so species reportedly shared by Borneo and India (see Ja- 
yaram 1 982). only about five or six appear to be correctly iden- 
tified. It IS doubtful that more than one or two species are shared 



CALIFORNIA ACADEMY OF SCIENCES 



by Borneo and China. At taxonomic levels above the species 
level, however, biogeographic relationships are much more 
complex; the fishes of Borneo show numerous phylogenetic re- 
lationships with those of the Mekong and India, and some with 
those of China. 

Kapuas fishes exhibit many of the kinds of adaptations as- 
sociated with the nchest tropical freshwater ichthyofaunas. There 
are numerous small or minute species, including the tiny sal- 
moniform Siindasalanx minor and sexually dimorphic Rasbora 
axclrodi. males of which rival the Amazonian neon tetras in 
their striking red and blue coloration. Many species are secretive 
and apparently rare, especially among the catfishes, some of 
which are camouflaged like dead leaves. Some have highly spe- 
cialized feeding habits, including ant-eating halfbeaks, and a 
scale-eating chandid. and. perhaps most unexpectedly, a small 
Rashoru-Uke phytoplanktonophagous cyprinid with over 200 
elongate gill rakers on the first gill arch {Pcclcnocypris balaena). 
There are also two large midwater piscivores highly specialized 
for lunging upwards at prey, the cyprinid Macmchinclilhys and 
silurid Belodoiitichthys. Reproductive behavior of Kapuas fish- 
es, although many have not been studied in this respect, is 
extraordinarily diverse. At least 10 species in six families are 
oral-brooders, and many others have specialized modes of re- 
production. The large lowland rivers, such as the mainstream 
of the Kapuas and its major tributaries, are inhabited by more 
than 100 fish species; small to moderately large lowland forest 
tributaries have up to 40 species, and highland streams of com- 
parable size probably somewhat fewer species. As in the richest 
freshwater ichthyofaunas in other areas, there is little overlap 
in the kinds offish species inhabiting the large lowland riverine 
habitats, smaller lowland tributaries, and montane tributaries. 
An idea of species composition of fish communities in these 
habitats can be gained by consulting the species lists accom- 
panying collecting localities of the 1976 Kapuas ichthyological 
survey (pp. 13-20). 

Some of the fishes discovered in 1976 have already been 
reported upon. Thus. Leonard J. V. Compagno kindly joined 
me in revising the veiT poorly known freshwater stingrays or 
Dasyatidae of southeast Asia and in describing Himantuni sig- 
nifcr. the first freshwater species of stingray known from Borneo 
(Compagno and Roberts 1982). Another 1976 Kapuas discovery 
is Siindasalanx minor, representing a new family as well as a 
new genus and species (Roberts 1981). Snndasalanx are the 
tmiest and most tropical members of that large order of lower 
teleosts the Salmoniformes, which includes our northern salmon 
and trout. The existence of a salmoniform fish in Borneo was 
totally unexpected. The skeleton oi Snndasalanx is largely car- 
tilaginous and such bones as are present do not stain well or at 
all with alizarin. In order to study its osteology, specimens were 
prepared using the new alcian-alizarin technique for counter- 
staining bone and cartilage. In reality, alcian stains some kinds 
of bone that do not stain with alizarin, as well as cartilage. These 
osteological studies confirmed the suspected relationship of 5h/;- 
dasalanx to their nearest relatives, the Salangidae of Korea, 
.lapan, and China, and also revealed unique characteristics jus- 
tifying their recognition as a distinct family. Sundasalangidae. 
A monograph describing and comparing the skeletal anatomy 
of all of the salangoid fishes and revising their classification has 
been published (Roberts 1984). 

Other papers published in preparation for the present mono- 
graph include the description of Thrrssocypris smaragdmus. a 



new genus and species of compressed, emerald-green Cyprini- 
dae, superficially resembling an anchovy (Roberts and Kottclat 
1984) and several generic revisions. The revisions concern the 
mountain stream loach genus Gastromyzon (with descriptions 
of three new species from the Kapuas) (Roberts I9&2d)\ the 
catfish genera &/^?cy/7;/.v (Roberts 1983). Chaca {Rohens 1982/)), 
and Wallago {Rohens 1 982t'); the spiny eel genus Macrognalhus 
(Roberts 1980); and the previously monotypic pufferfish genus 
Chonerhinos (with descriptions of two new species from the 
Kapuas) (Roberts 1982c). The results of these studies are in- 
corporated into the present monograph, but the individual pa- 
pers should be consulted for additional information. 

Attention should also be called to a paper on unculi (Roberts 
1 982a), unicellular horny projections on the lips, fins, and other 
epidermal features of ostariophysan fishes. Unculi represent one 
of the key adaptive features of the Ostariophysi (including carps, 
loaches, and catfishes) and are ideally suited for observation 
with scanning electron microscopy. Unculiferous structures in 
Kapuas species of Cyclochcilichlhys. Osleochilus. Paracrosso- 
clnlns. (iasiromyzon. and Baganus are illustrated and dis- 
cussed in the paper cited; additional observations on the di- 
versity of the highly specialized unculiferous lips of Osteochilus 
are presented herein. My most recent observations of unculi, 
done with the scanning electron microscopy facility of the Cal- 
ifornia Academy of Sciences, but as yet unpublished, reveal the 
unculiferous nature of the extraordinarily specialized ostracoph- 
ilous larvae in members of the Eurasian cyprinid subfamily 
Rhodeinae. and the presence of filamentous algae and other food 
Items on unculiferous portions of the lips of the Chinese moun- 
tain stream loach Psciidogaslromyion niyersi Herre, 1932. For 
a full discussion of the adaptive and evolutionary significance 
of unculi in adult ostariophysans see Roberts (1982a). 

The approximately 7.000 specimens of 263 fish species ob- 
tained b\ the Kapuas surxey of 1976 have been deposited in 18 
museums and research institutions in Europe. North America, 
Indonesia, and Thailand. In many instances, specimens of species 
that are rare, poorly known, or diflncult to identify have been 
placed in museums that have primary type specimens of the 
same or closely related species. 

The main series of specimens, including holotypes of all new 
species and the 1st, 9th, and 10th specimen in every series of 
10 specimens, is deposited in the Museum Zoologicum Bogo- 
rense (Bogor. Indonesia) and bears catalog numbers MZB 3000- 
4005. The next most important series, including paratypes of 
nearly all new species, are deposited in the California Academy 
of Sciences and the Smithsonian Institution (catalog numbers 
CAS 49 1 8 1-5 1 3, 53049, USNM 2 1 8069-74, 2 1 8603, 229492, 
2301 52-362). The other institutions which have received mate- 
rial are the Academy of Natural Sciences of Philadelphia (.ANSP 
152021-2), American Museum of Natural History (AMNH 
41578-9, 48878-9. 48918-42), British Museum (Natural His- 
tory) (BMNH 1978.1.10:1-4. 1979.3.21:145-152. 1981.4.13: 
20-26. 1981.11.16:1, 1982.3.29:1-255), Field Museum of Nat- 
ural History (94193-255), Institut Royal des Sciences Natu- 
relles, Brussels (IRSNB 631-2, 1 9726-5 1 ), Kasetsart University 
Museum of Fisheries, Bangkok (KUMF 2848-60). Museum 
d'Hisloire NaturcUc de Geneve (MHNG 2087.3-5). Museum 
National d'Histoire Naturelle. Paris (MNHN 1982-650-724), 
Museum of Comparative Zoology (MCZ 57598-9, 58339-53), 
Museum of Zoology, University of Michigan (UMMZ 209852- 
930), National Inland Fisheries Institute, Bangkok (NIFI, col- 



ROBERTS- WESTERN BORNEO FISHES 



lection uncatalogued). Rijksmuseum van Natuurlijke Hislorie, 
Leiden (RMNH 28822-924), Royal Ontario Museum (ROM 
37406, 38599-625), Zoological Museum. Amsterdam (ZMA 
116.516-549), and Zoologisches Museum und Institut, Ham- 
burg (ZMH 6475-76). 

One of my aims in writing this monograph has been to point 
out numerous areas for further investigation. More species will 
surely be found in the Kapuas. and much systematic work re- 
mains to be done on the species already found there. Enough is 
known about the biology of the fishes to indicate that their food 
habits and modes of reproduction are extremely diverse, but 
for most of the species basic biological information is still lack- 
ing. For example, very little is known about fruit- and seed- 
eating fishes in the Kapuas. Whether the Kapuas fishes and 
flooded forest plain species have coevolved and interact to the 
extent found by Michael Goulding in the Amazon basin is un- 
known and perhaps doubtful. The Kapuas clearly does not have 
such a diverse array of large fruit- and seed-eating fishes as the 
Amazon, but those that are present may be important in dis- 
persing the seeds of ecologically dominant plant species which 
play an important role in fish ecology. Readers interested in this 
topic should consult Goulding ( 1 980). The cyprinid genera Lep- 
toharhus and Punttoplitcs. the catfish genus Pangasius. and the 
giant goramy (Osphroncmus goramy) are probably the most 
important fruit- and seed-eaters in the Kapuas. 

Another phenomenon of potential ecological significance about 
which little is known in the Kapuas is fish migration. Migrations 
evidently do occur, but their direction, extent, biological sig- 
nificance, and even the species involved are virtually unknown. 
According to fishermen interviewed in 1976, large ikan tapah 
(Wallago leehi) formerly migrated in large numbers in the Ka- 
puas mainstream, but their numbers have declined due to in- 
tensive gill-netting. There may have been a primary migration 
related to reproduction or some other seasonal activity of ikan 
tapah, or perhaps it was a secondary or adventitious migration 
enabling them to feed upon migratory prey species. Another 
large fish species suspected of migratory activity in the Kapuas 
mainstream is ikan pachil, Cosmochiliis falcifcr. 



Acknowledgments 

The decision to undertake an ichthyological survey of the 
Kapuas originated in a discussion with Ira RubinoflT. Director 
of the Smithsonian Tropical Research Institute, at the Museum 
of Comparative Zoology toward the end of 1974. Support for 
travel to Indonesia and for the survey was provided by the Office 
of the Assistant Secretary for Science of the Smithsonian Insti- 
tution. This was facilitated and coordinated by David Challinor, 
Ross Simons, and Rita Jordan. The work in Indonesia was done 
in cooperation with the Museum Zoologicum Bogorense and 
with the permission of the Indonesian Science Research Council 
(Lembaga Ilmu Pengatahuan Indonesia). I am greatly indebted 
to Didin Sastrapradja, Sjamsiah Achmad, Sampumo Kadarsan, 
and Adisoemarto Soenartono for their sympathetic support. 
Soetikno Woerjoatmodjo of the Museum Zoologicum Bogo- 
rense accompanied and assisted me during the first half of the 
Kapuas survey. 

The research was done mainly at the Tiburon Center for 
Environmental Studies of San Francisco State University and 



in the Department of Ichthyology of the California Academy of 
Sciences. I especially wish to thank Dean .lames Kelley, Ken 
Carpenter, Erwin Siebel, and Curtis O. David for facilitating 
my work at Tiburon, and George C. Lindsay, Frank Talbot, 
William N. Eschmeyer, and Tomio Iwamoto for their under- 
standing and assistance at the California Academy of Sciences. 
Additional research was done at the Rijksmuseum van Na- 
tuurlijke Historic, Zoologisch Museum Amsterdam, Zoolo- 
gisches Institut und Museum Hamburg, Museum National 
d'Histoire Naturelle, Naturhistorisches Museum Wien, and 
British Museum (Natural History). Specimens were borrowed 
for study from these as well as a number of other institutions, 
including the Field Museum of Natural History, Museum of 
Zoology of the University of Michigan, and the Smithsonian 
Institution. I am grateful to the following colleagues for their 
assistance: Marinus Boeseman, Lipke Holthuis, Martien van 
Oijen, Peter van Helsdingen (Leiden); Han Nijssen (Amster- 
dam); Alfred Post, Matthias Stehman (Hamburg); Marie-Louise 
Bauchot (Paris); Harald Anhelt. Rainer Hacker (Vienna); Oliver 
Crimmen. Gordon Howes, Alwyne Wheeler. Peter Whitehead 
(London); Donald J. Stewart (Chicago); Ree\e M. Bailey. Robert 
R. Miller, Walter Rainboth (Ann Arbor); and Susan Jewett 
(Washington, D.C.). 

Many persons responded to my requests for information, re- 
prints, bibliographic assistance, and so forth. I am especially 
grateful to colleagues who assisted in identification of species 
belonging to groups with which they were working: Thosapom 
Wongratana (Clupeidae and Engraulidae). Jaranthada Kama- 
sula (Ostcochilus), Walter Rainboth (Puninis. Tor). Maurice 
Kottelat {Nemacheilus). C. E. Dawson (Syngnathidae), Douglas 
Hoese (Gobiidae), and Donn E. Rosen (Synbranchidae). Generic 
citations and method of typification were reviewed with William 
N. Eschmeyer using his "Genera of Fishes" database at the 
California Academy of Sciences. Technical assistance at the 
California Academy of Sciences (including radiography and cat- 
aloging) was provided by Charles Ruark, Michael Heam, and 
David Catania. Bibliographic assistance was provided by Ray 
Brian, Doris Cantou, Lillian Dempster, James Jackson. Ruth 
Opper, Lesley Segedy (California Academy of Sciences); Ingrid 
Radkey (University of California at Berkeley); Jack Marquardt 
(Smithsonian); and Verlejean Parker and Robert R. Rofen (Kor- 
don Corporation. Hayward. California). Most of the photo- 
graphs were prepared by Alphonse Coleman in the Photography 
Lab of the Museum of Comparative Zoology. Other photo- 
graphs were kindly provided by Martin R. Brittan, W. I. Follett, 
Orrin Moon, Robert R. Rofen, Edward S. Ross, and Lloyd 
Lllberg. Scanning electron micrographs were prepared in the 
SEM Lab of the National Museum of Natural History by Susann 
Braden and Mary-Jaques Mann. The manuscript was typed by 
Francis Bertetta and reviewed by Reeve M. Bailey, William N. 
Eschmeyer, John Lundberg, and Lynne Parenti. 

Support for study of the material obtained by the 1976 Kapuas 
survey and of material from earlier collections was provided by 
the National Science Foundation under Grant DEB77-24759. 
This publication is funded by National Science Foundation Grant 
BFR86 12812. The Foundation provides awards for research in 
the sciences and engineering. The awardee is wholly responsible 
for the conduct of such research and preparation of the results 
for publication. The Foundation, therefore, does not assume 
responsibility for such findings or their interpretation. Any opin- 
ions, findings, conclusions, or recommendations expressed in 



CALIFORNIA ACADEMY OF SCIENCES 



this publication are those of the author and do not necessarily 
reflect the views of the National Science Foundation. 

History of East Indian Ichthyology, with Particular 
Reference to Freshwater Fishes of Borneo 

The present narrative on ichthyological collecting and sys- 
tematic research in the East Indies has been written because 1 ) 
such background is essential to systematists who have to study 
historical collections, manuscripts, and publications: 2) the sub- 
ject is inherently interesting and worthwhile, particularly in that 
it reveals the importance of individual contributions; and 3) in 
the course of my museum visits and systematic investigations 
significant new information came to light concerning some of 
the events in this history. As in other branches of systematic 
biology, knowledge of East Indian freshwater fishes accumu- 
lated not steadily, evenly, and as a matter of course, but slowly 
and unevenly, and only with great individual efforts and sac- 
rifices. The persons in this history should be memorialized, not 
least for the inspiration this may provide to future generations 
of workers. 

Before proceeding to the narrative, I wish to point out that 
some of the important contributions could have been described 
at greater length, and that some minor episodes have been in- 
cluded but others omitted, so that this is by no means a complete 
history of our topic. Only a relatively small amount of research 
time has been devoted to this essay; no doubt it could have 
been greatly enriched by reference to archival materials espe- 
cially in Leiden, Paris, and Vienna. 

The history of East Indian or Indonesian ichthyology begins 
with Johan or Johannes Nieuhof(c. 1610-1672), who in 1653- 
1662 represented the Nederlandsche Oost-Indische Compagnie 
on an exploratory and diplomatic mission to the East Indies 
and China. The extensively illustrated folio report of this mis- 
sion was first published in Dutch and French in 1665, and 
shortly thereafter in German, Latin, and English. It includes 
relatively detailed accounts of the people, towns, industries, 
agriculture, and natural productions of China and Indonesia. 
Some 43 East Indian species of fishes are named and briefly 
described, and nearly all of them arc illustrated. This fish ma- 
terial was incorporated as an Appendix to Willughby's De His- 
foria Piscluiii { 1 686). NieuhoPs work stands not only as the first 
but almost as the sole contribution to knowledge of freshwater 
East Indian fishes for more than a century and a half ( 1 58 years). 
It is therefore of interest to inquire more closely as to the actual 
species he discovered. To my knowledge there is no review of 
the subject. Nieuhof apparently did not bring any fish specimens 
back to Europe, or if so they probably were lost long ago, but 
all or almost all of the families, most of the genera, and many 
of the actual species can be recognized from his figures, some 
of which were identified by Cuvier and Valenciennes and many 
others by Bleeker. The great majority of the species are marine 
and will not be discussed further here. Five species, however, 
are freshwater (or estuarine and freshwater in one instance) and 
arc of particular interest. These arc Nieuhofs "Penne Visch'," 
"Wit Visch," "Bont-ael" or "Negen-oogen." "Kabos" or "Ael- 
quabben"" and "Vijfoogen," or "Vyf Oogh." 

The "Penne Visch"" was identified by Bleeker (1862-63;60) 
as Aspiclohagnis i=MystHs) giilio. a species described from the 
Ganges by Hamilton-Buchanan (1822) and generally considered 



to be widely distributed along the coasts, in estuaries, and in 
river mouths, from India to southeast Asia including the Greater 
Sunda Islands. My observations, however, indicate that at least 
two species are involved; I therefore agree with Bleeker's generic 
identification but consider the species tentative. The "Wit Visch" 
was identified by Bleeker (1862-63:84) as Callichrous (=0m- 
pok)bii}iaciilanis and I concur. The excellent figure of the "Bont- 
ael" was identified by Valenciennes in Cuvier and Valenciennes 
( 1 839:387) as his new species Clarias nieiihofi. The generic iden- 
tification is unquestionably correct but the species probably can- 
not be confirmed. The "Kabos" was identified by Bleeker ( 1 862: 
103) as Clanas melanoderma {=Clarias meladerma). but this 
IS cleariy in error. Although deficient in details, the figure strong- 
ly resembles a Channa or snakehead. This identification is sup- 
ported by Nieuhofs vernacular name, a spelling variant of ikan 
gabus. the Indonesian name usually applied to Channa striata. 
Although crude, the figure of "Vyf Oogh" is based unmistakably 
on Maaognatlnis acideatiis. This highly specialized genus and 
species is indicated by the very elongate rostrum, anterior por- 
tion of dorsal fin consisting of about 13 separate sharp spines, 
and base of soft dorsal fin with a row of large, well-defined, and 
evenly spaced "eye-spots" or ocelli. 

In summary, the identities of the five freshwater fishes re- 
ported by Johan Nieuhof insofar as they can be reasonably 
inferred are Mystus cf gulio. Ompok bimaculalus. Clanas sp., 
Channa cf. striata, and Macrognathiis acideatus. These are al- 
most certainly the earliest published illustrations of these species. 
That oi Clarias probably is the earliest illustration of the family 
Clariidae (see Valenciennes //; Cuvier and Valenciennes 1840: 
359). That of Macrognathus aciileatiis considerably antedates 
the earliest published (1718) and unpublished ( 1 7 1 5) accounts 
and figures of Mastacembelidae cited by Sufi (1956). Most of 
Nieuhofs fishes were collected in or near Batavia (M. Boese- 
man. L. Holthuis. pers. comm.. July 1984). 

The seventeenth and eighteenth ccntunes saw a great flow- 
ering of private natural history cabinets in Europe. This led to 
a flourishing market for natural curiosities in centers such as 
London and Amsterdam. Many cabinets obtained specimens or 
illustrations of East Indian zoological material, most of it never 
recorded and subsequently lost. There were exceptions, how- 
ever, some of them notable. In 1704 Michael Valentini of 
Frankfort published some illustrations of East Indian fishes ob- 
tained for his Naturkammer; an enlarged second edition ap- 
peared in 1714 (neither edition seen by me). In 1715 Cornelis 
de Vlaming, an official of the Dutch East Indies Company, and 
Samuel Fallours, a physician stationed in Amboina, returned 
to Holland with a very large number of illustrations of East 
Indian fishes and other marine life. .A large, two volume work 
based on their materials was published in Amsterdam in 1718 
by Louis Renard. This contains 459 "crudely drawn and bar- 
barously colored figures" of East Indian fishes, crabs, etc. Several 
editions and reissues of this work with amendments and ad- 
ditions appeared. Francis Valentijn. a Dutch Protestant mis- 
sionary stationed in the East Indies in 1685-94 and 1706-14, 
published a five- volume description of the Orient in 1724-26, 
The third volume contains descriptions and figures of 528 fishes 
of Amboina. The authors of these works and their collaborators 
were not ichthyologists or even biologists, and the artists who 
prepared the illustrations had not been suitably trained. Never- 
theless, the large number of species and the variety of their color 



ROBERTS- WESTERN BORNEO FISHES 



and form, even crudely portrayed, served to proclaim the ex- 
traordinary ichthyological richness of the East Indies. 

The beginning of modem systematic knowledge of nearly all 
groups of organisms in the East Indies is intimately related to 
the founding of the Rijksmuseum van Natuurlijke Historic in 
Leiden and of the Natuurkundige Commissie voor Neder- 
landsche Indie in 1820. The guiding light of both organizations 
was Coenraad Jacob Temminck, the Rijksmuseum's director 
from 1820 to 1858. A considerable number of young and ca- 
pable naturalists, mainly from Holland and Germany and most 
with university education, were recruited by the Natuurkundige 
Commissie to obtain biological materials in the East Indies for 
systematic studies. They were usually instructed to make general 
collections, but many also pursued their individual interests. 
Unfortunately, during this early period medical knowledge and 
facilities were inadequate to cope with diseases in the East In- 
dies, and most of the Natuurkundig Commissie's recruits died 
within a short time (usuallly one to four years) after undertaking 
fieldwork. Despite individual tragedies, the collections they made 
almost always arrived intact at the Rijksmuseum. and Tem- 
minck saw to it that they were promptly reported on by that 
museum's systematists or by systematists at other institutions. 

Modern knowledge of East Indian fishes begins with Hemrich 
Kuhl and Johan Coenraad van Hasselt. After completing studies 
at the University of Groningen and receiving appointments in 
the Natuurkundige Commissie for the Dutch East Indies, they 
arrived in Batavia (Jakarta) in December 1820. They traveled 
widely in western Java, collecting plants as well as animals. 
Manuscript descriptions and illustrations were made for many 
species, which they presumably intended to publish upon re- 
turning to Holland, but such was not to be. Kuhl died after only 
nine months in Java, van Hasselt after two years and nine months. 
The sickness which caused their deaths has not been identified, 
but malaria, amoebic dysentery, or yellow fever, or perhaps a 
combination of these diseases, could have been responsible. 
Kuhl was only 25 when he died, van Hasselt 26. They are buried 
in the cemetery of the Botanic Gardens at Bogor (Buitenzorg). 

Kuhl and van Hasselt's collections and manuscripts arrived 
safely at the Rijksmuseum van Natuurlijke Historic and were 
soon reported upon by various systematists. (Several East Indian 
plants, especially palms, and many fishes have been named after 
them.) The job of reporting on the fishes fell to the great French 
ichthyologists George Cuvier and Achille Valenciennes at the 
Museum National d'Histoire Naturelle. Before he died, van 
Hasselt wrote two long letters to Temminck, providing an over- 
all impression of the ichthyological discoveries. The letters con- 
tained manuscript latinized names for many new genera and 
species, some with no descriptions at all. others accompanied 
by very brief remarks on morphology, but none with what van 
Hasselt himself would have regarded as a publishable scientific 
description. Temminck published the two letters in the Alge- 
meene Konst- en Lei!er-Bode{\S22). Shortly thereafter a French 
translation appeared in Ferussac's Bulletin des Sciences Naiu- 
relles et de Geologie. Although many or perhaps all of the fish 
names in the letters are based on specimens that arrived in 
Leiden, and some of those specimens apparently were accom- 
panied with labels bearing the manuscript names, presumably 
none of the specimens were labeled specifically as types by van 
Hasselt. Many of the fish names were written on the fish illus- 
trations, which had been prepared in Java, but the illustrations 



were not published in the Algemeene Konst- en Letter-Bode or 
in Ferussac's journal; so far as I know none of the illustrations 
has ever been published. An English translation of the letters, 
prepared with M. Boeseman, was published by Alfred (1961). 
The full extent of the ichthyological discoveries made by Kuhl 
and van Hasselt cannot be judged from van Hasselt's ichthy- 
ological letters because many species they obtained are not men- 
tioned. This became evident to me while going through the 
volumes of Cuvier and Valenciennes's Histoire Naturelle des 
Poissons. in which Kuhl and van Hasselt's material is more fully 
reported upon. Cuvier and Valenciennes retained many manu- 
script names coined by Kuhl and van Hasselt (occasionally mod- 
ifying the original spelling) but rejected many others, including 
some of those published in the Algemeene Konst- en Letter- 
Bode. They invariably cited Kuhl and van Hasselt as the authors 
of these names, even though it is clear in nearly every instance 
that Cuvier and Valenciennes drafted the descriptions them- 
selves. Kuhl and van Hasselt manuscript names published for 
the first time by Cuvier and Valenciennes are attributable either 
to Cuvier, or to Valenciennes, in Cuvier and Valenciennes, not 
to Kuhl and van Hasselt. For the names published in the Al- 
gemeene Konst- en Letter-Bode ( 1 823), the situation is compli- 
cated. A fair number of them are outnght nomina nuda; the 
original descriptions of some of these date from the volume of 
Cuvier and Valenciennes in which they first appeared, while 
others remain unpublished and it is unknown which species 
they represent. A few species described in the van Hasselt letters 
(1823) are so distinct that even the brief original description is 
perhaps adequate for their recognition, for example. Dermog- 
enys pusillus and Acantopsis diahtzona. For most of the names 
published in 1823, however, a strong case can be made for 
rejecting them as nomina nuda or dubia, and in most instances 
rejection is in the best interest of nomenclatural stability. Rec- 
ognition of any of the 1823 names poses serious problems either 
of identification or typification at generic or species levels. Ho- 
lotypes cannot be identified for any of the species. It seems to 
me that Cuvier and Valenciennes, Bleeker. and other ichthy- 
ologists of the time did all that they could to give credit to Kuhl 
and van Hasselt and to preser\e their memory, by recognizing 
their manuscript names and attributing original authorship to 
them. This unfortunately led to considerable nomenclatural in- 
stability, because some authors uncritically accepted names re- 
jected by others. Chief offender among recent authors is H. M. 
Smith (ill Smith 1945 and in earlier papers), who recognized a 
number of Kuhl and van Hasselt names which earlier workers, 
equally or more familiar with their work and the historical cir- 
cumstances, had chosen not to recognize. A case in point is 
provided by the genus Labiobarbus and the species L. Itpocheilus 
and L. leptocheilus of van Hasselt in Kuhl and van Hasselt 
(1823). These are unquestionably nomina nuda. as Smith him- 
self presumably would have recognized had he consulted the 
Algemeene Konst- en Letter-Bode. Of the relatively few names 
that can or should be accepted, it should be noted that the 
original spellings of the genera currently known as Acanthoph- 
thalmus and .Acanthopsis are .Acantophthahmis and .-icantopsis. 
and that these spellings undoubtedly were intentional and there- 
fore must be used according to ICZN. Although Noemachcilus 
Kuhl and van Hasselt, 1823, has been accepted by some authors, 
it has been rejected by others. It can be subjectively defined as 
a nomen nudum, and. because of the difficulty in determining 



CALIFORNIA ACADEM'>' OF SCIENCES 



ils generic type species, I have rejected it in favor oWenmcheilus 
Bleeker, 1863. with the originally designated type species Cobitis 
fasciatus Valenciennes in Cuvier and Valenciennes, 1846. This 
seems to be clearly in the best interest of nomenclatural stability 
as well as in accord with ICZN. A more perplexing problem 
arises from Acantophthalmus van Hasselt. 1823. This name is 
also subjectively definable as a nomen nudum, but its rejection 
would lead to generic name changes for numerous Asian loaches. 
The resulting confusion and opposing viewpoints probably could 
not be resolved except by petition to the International Com- 
mission of Zoological Nomenclature. These and other nomen- 
clatural problems posed by van Hasselt (1823) are taken up 
more fully in the systematic accounts of the taxa involved. 

The specimens obtained by Ruhl and van Hasselt, including 
the type specimens of many new species described by Cuvier 
and Valenciennes, are deposited in the Rijksmuseum van Na- 
tuurlijke Historic and in the Museum National d'Histoire Na- 
turelle. Their original illustrations of .lavanese fishes were con- 
sulted in preparing descriptions for the Histoire Nalurelle des 
Poissons but were never published. At least 32 species reported 
herein from western Borneo were first discovered by Heinrich 
K.uhl and Johan Coenraad van Hasselt in .lava. 

In 1826 the Natuurkundige Commissie sent Heinrich Boie, 
Heinrich Cristian Macklot, and Salomon Miiller to the East 
Indies. Boie, a herpetologist, died in Buitenzorg not long after 
his arrival. His collections and manuscripts were reported upon 
by his older brother Friedrich Boie, who remained in Holland. 
Macklot collected in Java, New Guinea, and Timor; his collec- 
tions and manuscripts were lost in a fire during the Chinese 
revolt in Poerwakarta, Java, in May 1 834, and he was murdered 
there a few days later. Salomon Miiller was perhaps the most 
fortunate and therefore the most successful of the Commissie's 
zoological recruits. It is said that Miiller was a waiter in a bar 
in Heidelberg in 1823 when he overheard Boie and Macklot 
discussing their forthcoming trip to the Indies. The three young 
men became close friends, and the Natuurkundige Commissie, 
presumably persuaded by Boie and Macklot, gave Miiller an 
appointment. Mullercollectedm Java (1826-28, 1829-32, 1836), 
Celebes ( 1 828), Amboina (1828), southwest New Guinea ( 1 828- 
29), and Samauw ( 1 828). The holotype of the earliest freshwater 
fish discovered in New Guinea, Ariiis Icpia.yns Bleeker. 1863, 
was collected by Miiller, but his most important zoological dis- 
coveries were made in Borneo in 1836, when he ascended the 
Doeson or Barito River as far as Moara Teweh. There he found 
the long-snouted piscivorous crocodilian Tomislonm sch/cgeli 
(van der Hoeven and de Vries, 1838) and the bony-tongue fish 
Sclcropages fonnosus (Muller and Schlegel, 1 844). According to 
Popta (1906:270), Muller collected nearly 100 species of fishes 
in Borneo; these are deposited in the Rijksmuseum but their 
identifications have not been reported. 

In 1843 Johann Jacob Heckel. the first member of the nine- 
teenth century ichthyological triumvirate of the Naturhisto- 
risches Museum Wien (Heckel, Kner, Steindachner), published 
his "Fische Syriens," one of the most remarkable papers in the 
history of ichthyology. The full title of this paper is "Abbildun- 
gen und Beschreibungen der Fische Syriens, nebst einer neuen 
Classification und Characteristik sammtlicher Gattungen der 
Cyprinen." Into its 109 closely written pages Heckel packed an 
original classification and synopsis of all known cyprinoid fishes, 
the first comprehensive descriptive account of cyprinoid pha- 



ryngeal teeth, and a detailed systematic account of all fishes 
known from Syria. The account of cyprinoid fishes includes 
descriptions of the first freshwater fishes known from Borneo: 
Cyreiic fesliva. Cyreue ocellata. and Barbus carassioides. The 
first two of these were well described and are valid species of 
the genus Dangila. The third, for which Heckel gave only a brief 
diagnosis, has not been identified previously but is here tenta- 
tively referred to the genus Puntioplites. The only collection of 
Bornean freshwater fishes of record at this early date is that of 
Salomon Muller from the Barito or Doeson. In going through 
accession records of the Naturhistorisches Museum Wien, I found 
early records for three additional freshwater species from Borneo 
(no additional locality information); Helostoma temminckii 
( 1 839 and 1 842), Scatophagus argiis ( 1839), and Tetraodon jlu- 
vialilis ( 1 840). Either these specimens and those of the cyprinids 
described by Heckel were part of Miiller's Barito collection or 
they represent some other early Bornean collection of which 
there is no published record. 

The second earliest known collection of Bornean freshwater 
fishes was made by geologist Carl Schwaner, another member 
of the Natuurkundig Commissie who died young. Schwaner was 
charged with investigating the coal beds of Borneo. In 1843-47 
he explored the Barito, Mahakam. Kahajan, and Kapuas basins. 
He was perhaps the first explorer to traverse the Barito-Ma- 
hakam and Kahajan-Kapuas watersheds. The latter traverse was 
done in the vicinity of Mount Raja, at 2,279 m, the second 
highest peak in Borneo (the highest, at 4,102 m, is Mount Kin- 
abalu in the northeastern corner of the island). Schwaner then 
descended the Melawi River and the Kapuas mainstream to 
Pontianak. A collection of some 40 species of Bornean fishes 
(all from the Barito'!') made by him is deposited in the Rijks- 
museum (Popta |Q06:271-272). but again their identifications 
have not been reported. The mountain ranges forming the wa- 
tershed between the Kapuas and the rivers of southern Borneo 
are named after him. The fishes of the Schwaner Mountains, 
including Mount Raja, are still almost entirely unknown. 

In 1851 Ida PfcilTer left Vienna on her second voyage around 
the world. This time she traxeled to London, the East Indies, 
and the United States. From Kuching. against the advice of 
Rajah James Brooke, she ascended the Batang Lupar, trans- 
versed the Lupar-Kapuas watershed and discovered the Kapuas 
Lake District. Pfeifltr then descended the Kapuas River to Pon- 
tianak, where a collection of freshwater fishes was obtained. 
This collection, or at least part of it. was presented to Bleeker, 
whom she met in Batavia. Bleeker (1852/)) identified 16 species 
from this collection (half marine, half freshwater). In the Natur- 
historisches Museum Wien I found accession records of two 
additional Bornean freshwater fishes collected by her, Anibly- 
rhynehichlhys tnincatus and Leptobarbus hoevemi. both de- 
scribed by Bleeker one year prior to her visit. For additional 
information concerning Ida Pfeiffer and her exploration of Bor- 
neo see Pfeiffer (1856) and Lebzeller (1910). 

By far the most important contributions to East Indian ich- 
thyology are those of Pieter Bleeker (1819-1878). Bleeker's bib- 
liography comprises some 730 items. 520 on fishes, including 
numerous monographs of substantial length. At a time when it 
was difficult to stay alive in the East Indies, he remained for 18 
years, amassing a personal collection of some 2,500 fish species, 
over 1,100 of which he described as new, and published some 
270 papers on systematic ichthyology. This work was accom- 



ROBERTS-WESTERN BORNEO FISHES 



plished in addition to official duties as surgeon and medical 
officer in the Dutch East Indian army and other substantial non- 
ichthyological activities. 

During his lengthy sojourn in the East Indies (1842-1860), 
Bleeker organized or assisted societies and initiated and edited 
journals which served as the main stimuli for scientific activities 
in the East Indies, notably the Koninklijke Natuurkundige Ve- 
reenigung in Nederlandsch Indie. Military and civil associates 
sent a steady stream of fish collections from throughout the 
Dutch East Indian islands to Bleeker; reports on such collections 
were promptly presented at meetings of the scientific societies 
in Batavia and printed in their journals. In 1860 Bleeker re- 
turned to Holland (retiring from military service in 1863) to 
continue his ichthyological work and care for his ailing wife, 
whom he survived by one year. From 1860 until the end of his 
life in 1878 Bleeker published continuously. The te.\t and il- 
lustrations of his monumental opus, the Atlas Ichthyologiqiie 
des Indes Orientales Neeiiandaises. were completed during this 
period. The first fascicles of the Atlas were published in 1862. 
By 1 878 all but the last fascicle of text for the first nine volumes 
and all of the plates for the first 10 of the projected 14 or 15 
volumes had been published (the uncompleted text of volume 
9 ends in the middle of the description of Pristolepis fasciala). 
The remaining plates for volumes 11-14 were finally published 
(without text) m 1984 (Bleeker 1984). 

It is largely due to the efforts of Marinus Boeseman of the 
Rijksmuseum van Natuurlijke Historic that the plates for vol- 
umes 1 l-Uofthe.-t/tearenow published, that many of Bleek- 
er"s works have been reprinted (including the first three volumes 
of the Atlas), and that the basic facts about Bleeker's life and 
the fate of his ichthyological collections (including the type spec- 
imens of his new species) are readily available. Readers may 
wish to consult Boeseman's English translation of Bleeker's fas- 
cinating autobiography (Bleeker 1973), and his Introduction to 
the plates of volumes 11-14 of the Atlas (in Bleeker 1984). I 
have also consulted the manuscript of an unpublished paper on 
Bleeker's life and collections prepared by Robert R. Rofen with 
the help of Boeseman and Lipke Holthuis (Rofen, ms). 

Despite repeated efforts, Bleeker never received a salaried 
appointment from any museum. A great deal of his time and 
energy was devoted to raising funds for his ichthyological col- 
lections and publications. Bleeker's fish specimens were his per- 
sonal property until he disposed of them, and specimens re- 
mained in his possession in Batavia and in Holland for varying 
periods of time before they were deposited in museums. The 
overall history of Bleeker's collections is now fairly well known. 
Bleeker material is scattered in at least a dozen museums, but 
with very few exceptions his type specimens are deposited either 
in the British Museum (Natural History), or, as a result of public 
auction of Bleeker's effects after his death, in the Rijksmuseum 
van Natuurlijke Historic. Unfortunately, some of the types 
probably have been lost or were not identified as types, and 
some non-type specimens have been incorrectly labeled as types. 
It seems that at least some of the material Bleeker assembled 
in Batavia did not reach Europe. 

The main facts of Bleeker's life are recounted in his auto- 
biography. In 1840 and again in 1841, before departing for 
Batavia, Bleeker applied to the Rijksmuseum van Natuurlijke 
Historic for an appointment. The reason for his rejection has 
been unclear and might be revealed by consulting archival ma- 



terial in Leiden. Perhaps it was due to his youth and humble 
origins. Certainly this very brief summary does not do justice 
to Pieter Bleeker, an extraordinary man who surmounted great 
obstacles to achieve what he did, and whose greatness and hu- 
manity found expression in a wide range of non-ichthyological 
endeavors. 

Of the 290 species of freshwater fishes now known from west- 
em Borneo, no fewer than 123 were originally described by 
Bleeker, mainly from specimens collected by his associates in 
Sumatra or southeastern Borneo (particularly Banjarmasin). 
Bleeker named many of these species after the persons who 
collected them; their names and position are recorded in Bleek- 
er's Atlas and other works. Bleeker received specimens of only 
about 60 species of freshwater fishes from western Borneo (mainly 
Sambas and Pontianak), somewhat of a colonial backwater dur- 
ing his time. From 1850 to I860 Bleeker published 13 papers 
on freshwater fishes of Borneo. His principal collaborators in 
obtaining specimens of freshwater fishes from that island, all 
doctors or medical officers, were C. Helfrich (Barito and Ka- 
hajan), .1. Wolffand M. A. Feldmann (Barito), .1. Einthoven and 
A. J. Andresen (Sambas), and M. J. N. Stevens, E. F. J. van 
Kappen, A. H. Thepass, and D. Sigal (Kapuas). 

Researchers on systematics of Indo-Pacific fishes are indebted 
to Max Weber and Licven F. de Beaufort for bringing so much 
information together in their great faunal work. The Fishes of 
the Indo-Australian Archipelago (which unfortunately ceased 
publication, although nearly completed, with volume 12). The 
first volume of this work comprises a nearly complete bibli- 
ography of the ichthyological publications of P. Bleeker, num- 
bered from I to 500, together with indices to the principal 
geographical localities and to the genera and species. Without 
these tools it would be very difficult to work with the Bleeker 
literature. A number of libraries, including that of the Depart- 
ment of Ichthyology of the California Academy of Sciences, 
have utilized Weber and de Beaufort's numbering system for 
filing their Bleeker papers; the numbers are given (in brackets) 
after each Bleeker item in the literature cited herein. 

In working with systematics of East Indian fishes one is fre- 
quently confronted by problems requiring examination of Bleek- 
er type specimens. In numerous instances, including rare species 
of which Bleeker obtained only a single specimen or a single lot 
of specimens, identification of the holotype or of the syntypes 
presents no problems. In many instances, however, due to the 
exigencies of circumstance, Bleeker's type specimens were placed 
in the same jar with more or less numerous (and likewise un- 
labeled) non-type specimens, often from widespread East Indian 
localities. There is also a distinct possibility that some of his 
types were destroyed or lost in the East Indies. Sometimes Bleek- 
er types can be recognized by pencil marks on the scales, a 
method used by Bleeker to facilitate scale counts. (Bleeker pre- 
sumably had a proper dissecting scope for his ichthyological 
researches in Holland after I860, but it may be that he lacked 
such a facility for all or most of the time during his stay in the 
East Indies.) Some ichthyologists have avoided the issue of deal- 
ing with Bleeker type specimens by regarding the plates in the 
Atlas as equivalent to types or at least based on types. This 
practice is unsound, since it is known that in many instances 
non-type specimens in better condition were utilized in the prep- 
aration of the plates. Due to the great amount of painstaking 
work involved and limited resources available for such a task, 



CALIFORNIA ACADEMY OF SCIENCES 



the Bleeker material deposited in the Rijksmuseum has never 
been thoroughly revised. In some instances lectotypes or neo- 
types may have to be designated for Bieei<er species, but this 
should only be done after careful study by persons thoroughly 
familiar with the history of Bleeker's material. In revising the 
supposedly monotypic freshwater pufferfish CluuicrhinosiRoh- 
erts 1982(') I found that Borneo was inhabited by five species 
which had been lumped under Chonerhinos modest us (Bleeker, 
1852). As Bleeker's holotype could not be positively identified, 
a neotype was designated for it. This was only done after the 
revision of the genus had been completed, all Bleeker material 
of Choucrhnws had been examined, and I had consulted with 
M. Boeseman. The only other Bleeker species for which I have 
designated a neotype is Myslus micracanlhus (herein). Addi- 
tional examples of Bornean freshwater fishes for which diffi- 
culties have been encountered in locating or identifying Bleeker 
type specimens include Misgwnus barbatuloidcs (see comments 
under Lcpidocephalichthys hasselti) and Glyptothorax platypo- 
gonoides. 

Eduard von Martens ( 1 83 1-1 904), one of the most productive 
malacologists in the second half of the nineteenth century, rep- 
resented the Berlin Museum on the Preussische Ost-Asien Ex- 
pedition in I860. After one year he left the expedition vessel 
Thetis and remained in Asia for three years on his own. He 
visited Japan, China, the Philippines, Thailand, Singapore, Su- 
matra, Java, Celebes, Molukkas, Timor, and western Borneo 
before returning to Berlin in 1 864. Although primarily interested 
in terrestrial and freshwater mollusks, von Martens made ex- 
tensive collections of other groups including fishes. His out- 
standing zoological discovery is the Yangtze paddlefish Psephu- 
riis gladius {Manens, 1 86 1). In March-June 1863 he collected 
some 90 species of freshwater fishes along the west coast of 
Borneo at Sinkawang, Seminis. Lumar. and Benkayang and in 
the interior of the Kapuas basin at Pontianak. Mandhor. Pulo 
Matjan (=Pulau Matjang), Danau Sriang, Selimbouw (=Selim- 
bau), and Sintang. These collections were identified in Berlin 
by Wilhelm Carl Peters. Only two new species were described, 
the pipefish Syngnaihus (=Daryichthys) nuiriemti and the 
snakehead Ophiocephalus vagus (a junior synonym of Chaiina 
striata). All of the Asian fishes collected by von Martens and 
identified by Peters are listed with their localities in the official 
account of the Preussische Ost-Asien Expedition (Martens 1 876). 
Peters was a very competent ichthyologist but his research was 
mainly on freshwater fishes of east Africa and evidently he was 
not very familiar with species from southeast Asia. The collec- 
tions, apparently intact, are now kept in the Zoologische Mu- 
seum of the Universiteit Alexander von Humboldt in East Berlin 
(H. J. Paepke, pers. comm., June 1981). It seems that none of 
the Bornean specimens of Eduard von Martens have been crit- 
ically reexamined since their original identification. 

A little known episode in the ichthyological history of Borneo 
concerns collections made in the Barito basin by Dr. H. Brei- 
lenstein. A German, Breitenstein served as regimental physician 
in the Dutch East Indian army in the Netherlands East Indies 
for 21 years. In 1880 he was stationed for some 18 months at 
Moara Teweh, at the junction of the Barito and Teweh rivers, 
then perhaps the most remote administrative post in the interior 
of Borneo. He sent anthropological and zoological collections 
to Europe. His fishes, all freshwater species obtained in the 
vicinity of Moara Teweh. were purchased for the Naturhisto- 
risches Museum Wien by Franz Steindachner. .'\fter retiring 



Breitenstein published a fascinating memoir on his experiences 
in the Dutch East Indies, filled with anthropological and medical 
observations, but with only a passing mention of fishes and fish 
collecting (Breitenstein 1899). Steindachner (1881^, /') de- 
scribed two new genera and species collected by Breitenstein, 
the cyprinid Parachela hrcitensteini and the very strange and 
highly specialized akysid Breitensteinta instgnts. Many other 
species were identified by Steindachner but never reported upon. 
During a visit to the NHW in September 1982, with the assis- 
tance of Harald Ahnelt and the late Rainer Hacker, I was able 
to locate accession records for the following 38 species collected 
by Breitenstein (as identified by Steindachner, with modern 
equivalents in some instances): Cyprinidae: Balantioeheilos 
nu'lanopteiiis. Barbichthys laevis. Barbus (Capoeta) suiiiatranus 
(=Puntius sunuitranus). Barynotus {=Cyelocheilichthys} micro- 
lepis. Chela (=Macrochiriehthys) macrochir. Dangila faseiata. 
Epaheorhynchos kal(>pterum. Luciosoma trinenia. Osteocheilus 
vittatus. Parachela breitenstetiu. Rasbora argyrotaenia, Thyn- 
nichthys thynnoides. Cobitidae: Acanthopsis eholrorhynehus. 
Botia hyiiu'iiophysa. B. macracantha. Siluridae: Calhchrous 
macronema (^Outpok hypophthalnius), IVallago leeni. Pan- 
gasiidae: Lais he.xanenia. Pangasiiis micronema. Bagridae: Bag- 
richthys hypselopterus. Bagroides melapterus. Maerones(=Mys- 
tus) neniurus. Maerones ( = Mystus) nigneeps. Akysidae: 
Breitensteinta insignis. Polynemidae: Polynennis midtifilis. Pris- 
tolepidae: Catopra faseiata (=Pristolepis Jasciatus), C. grootii 
{=P. faseiatus?). Toxotidae: Toxotes microlepis. Syngnathidae: 
Doryiehthys boaja. Channidae: Ophiocephahis (=Channa) lu- 
cnis. Ophiocephahis (=Channa) maruhoides. Ophiocephalus 
( = Channa) niicropeltes. Ophiocephalus striatus (=Channa stria- 
ta). Anabantidae: .Anabas oligolepis (=testudineusl). Betta pug- 
nax. Gobiidae: Boleophthalimis boddaerti. Tetraodontidae: Te- 
traodoii paleiiibungensis. \enopterus(=Chonerhinos) niodestus. 
It IS possible that this is only a partial list of the species obtained 
by Breitenstein. During my visit I examined four of the actual 
lots of specimens: Botia hyinenophysa. Breitcnsteinia insignis 
(holotype). P<^lyiiei}ius niultihhs. and .Xenoptenis (or Chone- 
rhinos) niodestus. My observations indicate that Botia hyineno- 
physa (otherwise unknown from the Barito) is correctly iden- 
tified; that the species reported from western Borneo and Sumatra 
as Breitensteinta insignis may represent one or more unde- 
scribed species; and that the Barito is inhabited by two species 
of Chonerhinos. C. iiitiJestus and C nefastus. It seems likely 
that knowledge of other Bornean fish species and their distri- 
bution could be similarly enhanced by examining other speci- 
mens collected by Breitenstein. Ichthyologists look forward to 
the day when the great collections of the Naturhistorisches Mu- 
seum Wien will again be available for examination. Our es- 
teemed friend and colleague Rainer Hacker was working to 
make this possible, and we can only hope that his untimely 
death will not signify an end to the project he began. 

In 1884. according to collection records in Vienna. Franz 
Steindachner purchased a small collection of Kapuas fishes from 
one Fritz Grabowski. At least four species were included: Chela 
hypophthalnius. Cyclocheilichthys apogon. Leptobarbus hoe- 
renii. and Chaca bankanensis. These specimens apparently were 
never reported on; the collector presumably was zoologist Frie- 
derich Grabowsky. who traveled extensively in Borneo in 1881- 
84 including over four months in the Kapuas basin. He sub- 
sequently held professional positions in Breslau and Brunswick. 

The last decade of the nineteenth century witnessed a re- 



ROBERTS-WESTERN BORNEO FISHES 



markable burst of ichthyological exploration of the Kapuas ba- 
sin, greatly increasing knowledge of its fishes. Civil mining en- 
gineer, biologist, and explorer Maurice Chaper traveled to western 
Borneo in November 1890-January 1891, making a valuable 
fish collection in the Upper Kapuas and two of its tributaries, 
the Knapei and Sebruang. This collection, deposited in the Mu- 
seum National d'Histoire Naturelle, comprised some 500 spec- 
imens of about 100 species and was reported upon by Leon 
Vaillant. From examining selected portions I learned that Vail- 
lant underestimated the number of species and overlooked or 
misidentified several that were undescribed. Although Vaillant 
described three species from Chaper's material as new, two of 
these are junior synonyms of species described by Bleeker. Vail- 
lant believed that Chaper's material did not contain many new 
species (Vaillant 1893a:55), but in fairness it should be noted 
that prior to Chaper's collections the Paris museum did not 
have any material of freshwater fishes from Borneo. Among the 
undescribed fishes collected by Chaper are the minute noodlefish 
Sundasalanx (mistaken for young needlefish), the remarkably 
specialized scale-eating chandid Paradoxodacna piralica (de- 
scribed herein), the belontiid Sphacrichthys vaillanti. and the 
pufferfish Choncrhinos amahilis. One specimen of the cyprinid 
genus Rashora and one of the bagrid genus Leiocassis collected 
by Chaper have not been identified to species and may represent 
species still undescribed. If one of these does prove to be new, 
it would be fitting to name it after Maurice Chaper, for the only 
Bornean fish species named after him, Diaslolomycter chapen 
Vaillant, 1 893, is a junior synonym of Hemisilums heterorhyn- 
chus Bleeker, 1853. Chaper has left two accounts of his travels 
in western Borneo (Chaper 1894a. b). 

Within three years of Chaper's Kapuas exploration, the So- 
ciety for the Advancement of Natural History Exploration in 
the Dutch Colonies (Maatschappij ter Bevorderung van het Na- 
tuurkundig Onderzoek der Nederlandsche Kolonien) sponsored 
an important series of scientific expeditions to central Borneo. 
These expeditions were in the field in 1893-94, 1896-97, and 
1898-1900. General accounts were published by Molengraaff 
(1895), Buttikofer (1897), and Nieuwenhuis (1900, 1904-07); 
zoological results appeared mainly in the Notes from the Leyden 
Museum (1897-1906). Zoologist J. Buttikofer collected fishes 
as well as birds, mammals, and other animals in the Kapuas 
basin. Anton Willem Nieuwenhuis, medical doctor, undertook 
ethnographic and anthropological studies and collected fishes 
in the Kapuas and Mahakam basins. Geologist Gustaaf Molen- 
graaff was the first scientist to describe the history of the river 
systems submerged by the South China Sea and to point out 
the significance of this history for distribution of freshwater 
fishes in the area (Molengraaff 1921; Molengraaff and Weber 
1921). The important fish collections, deposited in the Rijks- 
museum van Natuurlijke Historic in Leiden, were reported on 
by Vaillant ( 1 902) and Canna M. L. Popta ( 1 903, 1 905, 1 906). 
In her final compilation of the freshwater fishes of central Bor- 
neo, Popta indicated some 173 species from the Kapuas and 97 
from the Mahakam. .About 35 of her Kapuas species are re- 
dundant, so that the true number of species known from there 
at that time is closer to 140. The names in the Mahakam list 
are not redundant; little further ichthyological exploration of 
the Mahakam, the second largest river basin in Borneo, has 
been undertaken, and the total number of species now known 
from there is only a little more than 100. Hardenberg (1935/)) 
listed 39 species of freshwater fishes from the lower Mahakam, 



but many of his identifications are doubtful and the disposition 
of the specimens is unknown. The headwaters of the Mahakam 
and Kapuas evidently share a number of fish species, but Ma- 
hakam ichthyofauna is too poorly known to permit a worthwhile 
comparison of species composition in the two basins. 

A digression is needed here concerning ichthyological explo- 
ration in northern Borneo (including Sarawak) and the Malay 
Peninsula. Despite Rajah James Brookes's keen interest in nat- 
ural history, ichthyological knowledge in the British possessions 
of Borneo lagged far behind that in the Dutch territories, a 
condition which has not been rectified. One of the earliest fresh- 
water fishes described from northern Borneo is the remarkably 
specialized mountain stream loach Gastwmyzon borneensis 
Gunther, 1874. The first five specimens of Gastromyzon. col- 
lected in Labuan (now part of Sabah). were purchased from their 
collector by Albert Gunther of the British Museum (Natural 
History) for £1.00 (according to a pencil note apparently in 
Giinther's hand in the register of vertebrate accessions). Not 
until a recent revision of the genus (Roberts \9B2d) was it re- 
alized that the type-series of G. borneensis includes two very 
distinct species, and that northern Borneo is inhabited by no 
fewer than eight species of Gastromyzon. As late as 1 896 only 
about 60 species of freshwater fishes were known from all of 
northern Borneo (Bartlett 1896). This number was moderately 
increased by small collections of fishes made by C. Hose, A. 
Everett, R. Hanitsch, and others and reported on by G. A. 
Boulenger(1894, 1899) and C. T. Regan (1906, 1913/)) of the 
British Museum (Natural History), and by Willy Kiikenthafs 
collection of 32 species from the Baram reported upon by F. 
Steindachner(1901). In recent years Robert F. Inger of the Field 
Museum of Natural History, Eugene K. Balon of the LIniversity 
of Guelph, and Antony Lelek of the Senckenberg Museum and 
their associates have made extensive collections in the Rejang 
and Baram but little of this material has been reported upon. 
Extensive collections of fresh- and brackish water fishes from 
northeastern Borneo (including the Kinabatangan) have been 
reported upon (Inger and Chin 1962) so that this somewhat 
impoverished area is relatively well known. The freshwater fish- 
es of the Malay Peninsula, which one might suppose would be 
well known systematically, are poorly known. The first sub- 
stantial contribution to knowledge of freshwater fishes of the 
Peninsula is by Georg Duncker (1904), with about 100 species, 
including 1 4 described as new. In this paper Duncker announced 
the discovery of some remarkable cyprinodont-like fishes, but 
it fell to Regan (1913/). 1 9 1 6) to give the first formal descriptions 
of the family Phallostethidae. These small or tiny fishes, in which 
males have a highly specialized subcephalic copulatory organ, 
or priapium, are now known from the Philippines, Thailand, 
Sumatra, and northern Borneo as well as the Malay Peninsula, 
but have not been found in the rest of Borneo or in Java. Despite 
papers by Alfred, Herre, Herre and Myers, Hora, Hora and 
Gupta, and Tweedie much more ichthyological collecting and 
museum study will be needed for the freshwater fishes of the 
Malay Peninsula to become well known. A checklist of marine 
and freshwater fishes of the Peninsula by Fowler (1938) is useful 
for its bibliography. The recent account of the freshwater fishes 
of the Malay Peninsula by Mohsin and Ambak (1983) is con- 
cerned primarily with ecology but includes some distribution 
records of systematic interest. 

Returning to events in western Borneo, H. A. Loreniz visited 
the Kapuas in 1911, obtaining a small fish collection which was 



10 



CALIFORNIA ACADEMY' OF SCIENCES 



deposited in the Zoological Museum of Amsterdam. This ma- 
teria! is reported on in Weber and de Beaufort's Fishes of the 
Indo-Australian Archipelago. Fisheries biologist J. F. D. Har- 
denberg published a series of papers on marine, estuarine, and 
freshwater fishes of the Dutch East Indies from 1930 to 1948, 
including several on freshwater fishes from western Borneo. 
Unfortunately Hardenberg's identifications are not very reliable, 
and the disposition of most of the specimens reported on in his 
papers is unknown. A few Hardenberg specimens are deposited 
in Amsterdam. The validity or identification of three of the new 
species he described from western Borneo {Rashora beauforti. 
Paiigasius hoeksi. and Onipok wcben) is problematical. The type 
specimens of these nominal species are not deposited in Am- 
sterdam, Leiden, or Bogor; my letters concerning them ad- 
dressed to the Director of the Marine Fisheries Institute at Den 
Pasar in Jakarta have not been answered. A group of Japanese 
fisheries biologists reported on some 109 species obtained dur- 
ing a survey of the middle Kapuas in 1978 (Imaki et al. 1978); 
these specimens are deposited in the Tokyo University of Ag- 
riculture. In 1983 Pier Georgio Bianco of the Universita degli 
Studi delTAquila collected about 1,000 specimens from a trib- 
utary of the upper Kapuas; the Cyprinidae he obtained have 
been reported upon (Banarescu and Bianco 1984). 

Reproductive Adaptations in Freshwater Fishes 
OF Borneo 
Virtually every reproductive adaptation known in vertebrates 
has been found in fishes. An ecological classification of the modes 
of reproduction in tropical freshwater fishes (with Bomean ex- 
amples) is as follows: 

I. Reproduction continuous 

A. viviparity (Hemirhamphodon) 

B. oviparity 

1. spawning partial 

a. fertilization external (Chacal) 

b. fertilization internal (Phallostethidae) 

2. spawning total (Cahimiana sp.) 
II. Reproduction periodic 

A. viviparity (Himaniura) 

B. oviparity 

1. spawning partial (some Osteochihis'!) 

2. spawning total [Cosniochihisl] 

This simple classification does not account for all physiological 
and behavioral adaptations involved in fish reproduction. The 
viviparous or live-bearing categories could be further divided 
into so-called "ovoviviparity" (yolk nutntion) and "true vivi- 
parity" (maternal nutrition); for a critical discussion of this dis- 
tinction in teleosts see Wourms (1981). While of considerable 
interest from the standpoints of developmental and evolution- 
ary biology. It is relatively unimportant ecologically whether 
young internally developed receive nutrition from the yolk or 
more directly from the mother (in fact both sources of nutrition 
are maternal); the essential point is that free-living young emerge 
from the mother. A more important consideration ecologically 
is the matter of parental care, and this is also of much interest 
to behaviorists and evolutionary biologists. Parental care is a 
fundamental aspect of the reproductive biology of many ovipa- 
rous fishes, and possibly of many viviparous forms as well. This 
IS particularly true of tropical freshwater fishes. 
The main kinds of behavioral parental care found in fishes 



are as follows (adapted with substantial modification from Blu- 
mer 1982): 

1 . Nesting (including nest building, nest maintenance, egg bur- 
ying). 

2. Guarding (agonistic and other protective behavior directed 
mainly against potential predators on eggs and fry, including 
coiling body around eggs or fry). 

3. Cleaning (removal of foreign matter from eggs, fry, or spawn- 
ing site; removal of dead eggs or fry). 

4. Aerating (fanning eggs or fry with fins or forcing water cur- 
rents over them through mouth or gills; similar actions may 
be involved separately or simultaneously in cleaning). 

5. Oral transport (carrying one or a few individual eggs or fry 
for a short period, possibly a fixed action response, in which 
the primary object is to transport eggs; examples include 
placing spawn in nest, returning lost eggs or fry to nest). 

6. Oral brooding (carrying an entire clutch of eggs or fry in 
mouth for more or less prolonged periods). 

7. External brooding (carrying eggs or fry attached to body or 
in special brood pouches). 

8. Provision of food (see discussion below). 

Eggs and larvae of fishes are currently receiving long-overdue 
attention. Substantial surveys of marine fish larvae have been 
published (e.g., Leisand Rennis 1984, on Indo-Pacific coral reef 
fishes) and others are in preparation. A detailed classification 
of the "reproductive guilds" of fishes, based mainly on larval 
adaptations, was proposed by Kryzhanovsky (1949). This clas- 
sification has been translated into English, with extensive emen- 
dations and commentary, by Balon (1975, 1981). With notable 
exceptions, the information on freshwater fishes available to 
these authors came mainly from the temperate regions of North 
America and Eurasia. Svensson (1933) made numerous obser- 
vations on larval fishes in the Gambia River of west Africa, but 
comparable observations have not been made for other tropical 
rivers, and no truly broad survey of tropical freshwater fish 
larvae exists. The eggs and larvae of the great majority of south- 
east Asian and particularly Bomean freshwater fishes are entirely 
unknown. Highly specialized mountain stream fishes such as 
Gastmmyion and peculiar catfishes including Chacidae, Sisor- 
idae, Akysidae, and Parakysidae are likely candidates for dis- 
tinctive larval adaptations, but their larvae have never been 
observed. 

Observations of any kind on reproductive biology are avail- 
able only for a few of the freshwater fishes of western Borneo. 
This is especially true of the two largest groups, carps and cat- 
fishes. More is known about those with "advanced" parental 
care, especially the anabantoids. many of which are oral-brood- 
ers or bubble-nestersand have been studied in aquaria (the single 
most important reference to the reproductive and other behav- 
ior of these fishes is Forselius 1957). Western Borneo is peculiar 
in that it has the most diverse assemblage of oral-brooding fishes 
found anywhere on earth, involving at least 1 1 species in six 
families. This is truly remarkable when it is recalled that the 
total number of fish families known to practice oral-brooding 
IS only 1 1 . Anabantoid oral-brooders of western Borneo include 
Channidae (Channa oricntalis). Belontiidae {Betta pugnax. 
Sphaeriihthys). and Luciocephalidae; non-anabantoid oral- 
brooders are Osteoglossidae, Ariidae, and Chandidac. The last 
family was not previously known to have any species with oral- 
brooding, but this behavior occurs in the Bomean species Par- 



ROBERTS-WESTERN BORNEO FISHES 



anibassis apogonoides. in which a male has been observed with 
nearly 100 young of two age classes in its mouth (reported 
herein). In addition to oral -brooders, western Borneo has four 
species of the freshwater syngnathid (pipefish) genus Doryich- 
thys. in which the males (as in all members of the family) carry 
the developmg young in an abdominal brood pouch, a vivip- 
arous freshwater elasmobranch (Himantura signifer), and four 
species of viviparous freshwater halfbeaks (Hemiramphidae). 

While it has not been possible for me to make original ob- 
servations on reproductive biology for all families of freshwater 
fishes inhabiting western Borneo. 1 have tried to identify and 
document significant or unusual phenomena and to point out 
worthwhile areas of investigation. Some interesting generaliza- 
tions or hypotheses have emerged fom this process. A number 
of notions currently prevalent concerning reproductive biology 
of freshwater fishes will need to be reconsidered as more groups 
and species are investigated from tropical regions, especially 
where the ichthyofauna is exceptionally rich and diverse. There 
has been almost no study of the very numerous species that are 
small or very small, or of those that are rare. Small fishes do 
not have the same reproductive modes as the few relatively 
large species that have been most studied. The phenomenon of 
"rarity" or very low population density is poorly studied, but 
is prevalent in the richest tropical ichthyofaunas, such as that of 
western Borneo, and here again the modes of reproduction are 
not or are not likely to be the same as those of better known 
species. Consideration of reproductive modes in secondary as 
opposed to primary freshwater fishes should yield further in- 
sights. 

Freshwater fishes of at least some secondary freshwater fam- 
ilies tend to have more "advanced" forms of reproduction and 
parental behavior than closely related marine species. This is 
particularly true in rich tropical ichthyofaunas dominated by 
primary freshwater fishes. The outstanding southeast Asian ex- 
ample of this tendency is provided by the family Hemiram- 
phidae. in which the numerous marine species are all pelago- 
philes. spawning large numbers of eggs which develop offshore, 
while the somewhat less numerous freshwater species all have 
internal fertilization and are mostly viviparous. In Hemirham- 
phodon pogonognathits sexually mature females are almost in- 
variably pregnant, apparently producing one or a few relatively 
large young every few days until the end of their lives. Further 
examples are provided by Chandidae. Gobiidae. and Tetra- 
odontidae. One of the five endemic freshwater chandids in west- 
em Borneo is an oral-brooder, the only known member of the 
family with such behavior. At least some and perhaps all of the 
marine species are pelagophiles with relatively small eggs. Pa- 
rental behavior in other freshwater chandids is unknown; as of 
now there is no indication that they are also oral-brooders. 
Western Borneo has three species of the southeast Asian fresh- 
water tetraodontid genus Choncrlunos. all of which produce 
ovarian eggs 1.5-2.0 mm in diameter (Roberts 1982c). This is 
far larger than the eggs observed in Indo-Pacific coral reef tetra- 
odontids, which are demersal and under 1 .0 mm (Leis and Ren- 
nis 1984:244), No other information on reproduction in Cho- 
nerhinos is available; egg size and other aspects of reproductive 
biology are unknown for the freshwater species of Teiraodon 
inhabiting western Borneo, 

So far as I have been able to determine, there are no diad- 
romous fishes inhabiting the Kapuas River. This is in marked 
contrast to the situation in the Fly River, the largest river in the 



Australian region, where endemic freshwater species of numer- 
ous secondary freshwater families live in the total absence of 
primary freshwater fishes. In the Fly large species belonging to 
no fewer than eight secondary freshwater families have been 
positively or probably identified as diadromous. migrating to 
brackish or marine environments for spawning (Roberts 1978). 
Of these eight families, only two— Engraulidae and Datnioidi- 
dae — occur in the Kapuas. and they are represented by non- 
diadromous species. Comparisons of the taxonomic composi- 
tion of secondary freshwater fishes in the Fly, Kapuas, and other 
large tropical rivers with different ichthyofaunas and consid- 
eration of the meager information available on their reproduc- 
tive biology leads to the hypothesis that fishes with diadromous 
life histones generally do not survive where primary freshwater 
fishes are dominant (Roberts 1978:15-21). 

The widespread notion that all fishes must have a reproduc- 
tive "season" is contradicted repeatedly by species in many 
groups inhabiting large rivers in the "evergreen" equatorial rain 
forests. In the largest tropical rivers with the richest ichthyo- 
faunas, including the Amazon, Congo, and Kapuas, fishes re- 
producing throughout the year appear to be particularly nu- 
merous. Most information available on reproductive periodicity 
in such rivers has come from relatively few commercially im- 
portant species, mainly large forms, reproductively atypical of 
the ichthyofaunas as a whole. Even some large tropical species 
of commercial importance may breed continuously, e,g., pira- 
nhas {Serrasalnms), loricariid catfishes (Hypostomus sp,), air- 
breathing catfishes (Callichthyidae, Clariidae). and many Cich- 
lidae. As more groups with small and rare or otherwise unusual 
species are examined, it seems likely that specialized reproduc- 
tive adaptations will be found, and that many will have non- 
seasonal or continuous reproduction. Examples or probable ex- 
amples from Borneo include species of such diverse families as 
Cyprinidae, Chacidae, Hemiramphidae, Syngnathidae, Masta- 
cembelidae, Channidae, Belontiidae, Luciocephalidae, and Tet- 
raodontidae. In all of these families one or more species have 
been observed in which mature females tend to be reproduc- 
tively active regardless of the time of year. In most instances 
very small juveniles have also been found whenever the species 
was moderately abundant and extensive collections of small 
fishes were made. Some observations are presented here, and 
more information can be found scattered in the systematic ac- 
count of this monograph (e.g., under Rasbora axelivdi. R. ban- 
kanensis, Bella dimidiata. Calamiana sp.). Some species of 
Mastacembelus. especially the large M. erythrotaenia. evidently 
have highly restricted reproductive periods, since the young are 
unknown or very rarely collected, and even large females are 
rarely gravid. A notable exception, however, is provided by M. 
macidatiis. In this relatively small species, large samples usually 
include gravid females. Very small Mastacewbehts collected in 
the Malay Peninsula and western Borneo usually belong to M. 
maculatus. Females have a darkly pigmented, elongate, tubular 
genital papilla which I tentatively identify as an ovipositor, but 
oviposition and other aspects of reproductive behavior have not 
been observed. A similar genital papilla occurs in the closely 
related species M. gucntheri of India but is otherwise unknown 
m Mastacembelidae. More observations are needed, but the 
freshwater pipefishes of the genus Doryichlhys are probably re- 
productively active at all times. Syngnathidae as a group evi- 
dently have continuous ovulation, each ovan,' producing a uni- 
cellular layer of follicles of gradually increasing size (see Wallace 



12 



CALIFORNIA ACADEMY OF SCIENCES 



and Selman 1981:334-335, fig. 14c). While ovulation is inter- 
rupted for prolonged periods in pipefishes in temperate regions, 
tropical species might continuously produce synchronous 
clutches of eggs without interruption for long periods. It seems 
fairly clear from numerous accounts that various oral-brooding 
and bubble-nesting anabantoids breed throughout the year. Ex- 
tensive observations are reported herein indicating that the vi- 
viparous halfbeak Heniiramphodon pogonognalhus bears live 
young throughout the annual cycle and that its reproduction 
continues uninterruptedly for long periods (presumably until 
the end of life). For the great majority of fishes in western Bor- 
neo, however, many of which are smaller than any of those 
discussed here, no information is available concerning frequency 
or timing of reproduction. Based on the information available 
on some of the small and very small species," reproduction prob- 
ably occurs throughout the year in many of them, unless ter- 
minated by food scarcity or other unfavorable conditions. In 
my opinion, considering tropical freshwater fishes as a whole, 
it is generally inappropriate to speak of "reproductive seasons." 
Many of the large, highly migratory species with marked repro- 
ductive periodicity are total spawners. undergoing massive 
spawning in response to environmental cues or triggers within 
a very short period of time which hardly constitutes a "season." 
At the other extreme, numerous statements like "reproductive 
season March-October" are suspect; they are often based on 
inadequate information, and often do not correspond to any 
recognized seasonal phenomena of an ecological or climatolog- 
ical nature. In many instances it might be more appropriate to 
think rather in terms of "interrupted reproduction" or a "non- 
reproductive season." Probably it would be best to dispense 
with the notion of seasonality altogether and instead look for 
factors such as resource availability (mates, food, spawning sites) 
and other parameters which affect reproductive activities. For 
a critical review and new approaches to the study of reproductive 
periodicity in tropical freshwater fishes see Kirschbaum (1984). 

Extensive comparative studies have not been done on the 
gonads of tropical freshwater fishes, and surprisingly little in- 
formation is available about their gametes. Students of fecundity 
have assumed that ecological and life history parameters are 
directly related to fecundity. It is commonly taken for granted 
that in oviparous species all eggs produced arc spawned, and 
that all eggs spawned are fertilized, but neither assumption is 
justified. I suspect that in some highly fecund fishes, only a 
portion of the eggs produced may be spawned, and that the 
proportion fertilized varies greatly depending on many factors. 
Internal fertilization occurs in all viviparous (including ovovi- 
viparous) fishes, but the extent of internal or "oviducal" fertil- 
ization among oviparous fishes is unknown; it may be more 
widespread than realized. An advantage of oviducal fertiliza- 
tion, apart from insuring fertilization, is that it may permit 
temporal separation of mating and spawning. Thus mating can 
occur whenever sexually active males and females come to- 
gether, and spawning can occur intermittently over a long period 
after they have separated. Oviducal fertilization, documented 
in the oviparous Neotropical catfish family Auchenipteridae 
(von Ihering 1937), is possibly widespread among catfishes. 

Many catfishes have males with an elongate genital papilla, 
the significance of which is unknown. An even more widely 
distributed characteristic of male catfishes is possession of lo- 
bate testes. These first came to my attention while studying the 
west African amphiliid Aniphihus utcsiicnsis Boulenger, 1904. 



mature males of which have an elongate genital papilla. Since 
then I have found lobate testes in one or more species repre- 
senting 18 of the 32 families of catfishes currently recognized. 
These families are Amblycipitidae. Bagridae {Bagnchihys. Leio- 
cassis, Mysiiis). Pimelodidae. Ictaluridae. Cranoglanidae, 
Olyridae, Schilbeidae (Pseiideutropiiis hrachypopleriis). Panga- 
siidac, Mochokidae. Auchenipteridae. Ageneiosidae, Cetopsi- 
dae. Amphiliidae. Sisoridae {Bagarius. Glyptofhorax). Akysidae 
(all genera). Parakysidac, Clariidae, Heteropncustidae, and Plo- 
tosidae. Lobate testes have not been observed in Ariidae or any 
of the South American loricarioid families (many of which have 
been examined), or in any other ostariophysans. The morpho- 
logical peculiarity of catfish testes has received little attention. 
Lobate testes were reported, apparently for the first time, in the 
South American auchenipterid Trachycorystes galealus (Lin- 
naeus, 1 758), by von Ihering (1937); in this species males have 
an enlarged genital papilla attached to the modified anterior 
portion of the anal fin, and fertilization is internal ("oviducal"). 
Whether internal fertilization occurs in some or many other 
catfishes is unknown. It may be significant that males of catfishes 
living in mountain streams often have a particularly long genital 
papilla (e.g., Astroblepidae), but this does not necessarily in- 
dicate it IS employed as an intromittent organ. A few other 
reports of catfish lobate testes have come to my attention; Sun- 
daraj (1958) for Hetewpneiistes fossilis (Bloch, 1797); Nawar 
(1959) for Clarias lazcra Valenciennes m Cuvier and Valen- 
ciennes, 1840ri; Sneed and Clemens ( 1964) for Ictaluridae; and 
Tilak and Hussain (1973) for Glyptothorax brevipinnis Hora, 
1923. In catfishes both testes are usually well developed. The 
number of lobes is typically 20-30 for each testis, but in many 
instances they number in the hundreds. The lobes generally are 
elongate and tapered, but may be threadlike, globose, or oth- 
erwise modified; Pangasius pangasiiis (Hamilton-Buchanan, 
1822) has multilocular lobes. The lobes may become so hyper- 
trophied that the abdomen of a ripe male is nearly as distended 
as that of a gravid female. Lobe formation may begin long before 
sexual maturity. The disposition of the lobes, which may be 
obvious or obscured in adults, is usually (invariably?) lateral. 
In many instances size and superficial morphology of the lobes 
is identical for the entire length of the testes, but in others the 
anterior and posterior lobes may be strongly difltrentiated (e.g., 
in Trachyanystc's. Clariidae. Heteropncustidae, and Ictaluri- 
dae). In such instances the antenor lobe are reportedly semi- 
niferous and the posterior lobes non-seminiferous. In Trachy- 
corystes the posterior lobes secrete a gelatinous substance 
interpreted as an "oviducal plug" (von Ihering 1937), but which 
is probably a spermozeugma. In a few catfishes I have observed 
amber or orangish concretions at the posterior end of the testes 
(e.g.. in Psciulciitropiiis brachypoptcrus), but thus far I have not 
found similar concretions in the oviduct of females. For a review 
of spermozeugma and spermatophores in teleosts see Nielsen 
et al. (1968). 

One area in which field observations of parental behavior is 
likely to provide new information is parental food supply for 
the young. Observers should watch for the following possible 
kinds of behavior: 

a) ectodermal feeding (production of ectodermal secretions fed 
upon by fry); 

b) egg feeding (mother provides eggs fed upon by fry); 

c) regurgitation (parents regurgitate food for fry to feed upon); 



ROBERTS-WESTERN BORNEO FISHES 



13 



d) fecal feeding (parents facilitate feeding of fry on fecal matter); 

e) other forms of food provisioning (including killing or dis- 
abling prey fed upon by fry). 

Ectodermal feeding is well documented in a few tropical fresh- 
water fishes (e.g., the South American cichlids of the genus 
Symphysodon) but is as yet unknown in those of Borneo. Most 
catfishes (including many in Borneo) have pectoral glands which 
discharge a copious milk- or mucus-like substance through an 
axillary pore, but there is no evidence as yet that this substance 
is fed upon by the young (see observations on pectoral glands 
of Pangasiiis herein). The literature on reproductive and feeding 
behavior of fishes is replete with references to oophagy or egg- 
eating: fish eggs are preyed upon by conspecifics as well as a 
vast array of vertebrate and invertebrate heterospecifics. Many 
fishes guard the eggs to protect them from predation. In many 
instances, however, one or both parents will feed upon their 
own eggs. Intrauterine oophagy is now well documented in vi- 
viparous sharks of the family Isuridae. Egg feeding behavior 
should be watched for in species with free-living fry. If the egg- 
eating opportunity arises as a result of maternal behavior, it 
would constitute a form of parental care. So far as I am aware, 
there is no direct evidence of parental feeding by regurgitation 
or fecal supplying in fishes. Fecal feeding is very important in 
fishes generally, and conceivably may be involved in parental 
care. Observations of gut contents in young of the ant-eating 
hemiramphid Hcmirhaniphodon pogonognathus (q.v.) indicate 
that parental regurgitation or defecation may be involved. Other 
forms of parental food provisioning probably do occur. I suspect 
that killing of "excess" prey by large individuals of the snake- 
head Channa micropelies (Smith 1945: herein) represents pa- 
rental food supplying, and that similar behavior occurs in other 
large predatory fishes with protracted parental care. Finally, it 
should be noted that parental food supplying involving regur- 
gitation, feces, or killed or disabled prey might provide impor- 
tant leammg experiences for the young. 

To summarize, no observations on reproduction are available 
for the majority of taxa of freshwater fishes in Borneo. Enough 
is known to indicate that their reproductive biology is diverse 
and that excitmg discoveries await investigators with access to 
the fauna. Reproduction of the relatively few large species is 
generally difl^erent from that of the numerous small and very 
small species and should not be regarded as typical. Continuous 
or nearly continuous reproduction appears to be widespread. 
Periodic reproduction is poorly documented and not well under- 
stood. 

Collecting Localities of the Kapuas Ichthyological 
Survey of 1976 

Fieldwork in western Borneo (Kalimantan Barat) from 1 July 
through 23 August 1976 resulted in 55 collections from diverse 
freshwater habitats almost exclusively within the Kapuas River 
basin. These collections are designated Kapuas 1976-1 through 
55 (Fig. 1). It should be noted that Kapuas 1976-19, 52. and 
53 were obtained from fish markets at Sintang, Pontianak, and 
Kampong Djunkat, respectively, and that Kapuas 1976-4 through 
7 were made in small coastal drainages (including the Mem- 
pawah) immediately N of the Kapuas basin. The rest are from 
habitats within the Kapuas basin. 

Localities of the Kapuas survey of 1 976 (Fig. 2.3) were plotted 
in the field on series TPC topographic maps (scale 1:500,000, 



edition 2-GSGS). Distances and coordinates were subsequently 
calculated and Figure 1 prepared from the same maps. All place 
names employed in the locality descriptions are indicated on 
these maps. 

Kapuas 1976-1. Small forest stream and man-made ditch, 2- 
3 m wide and 1 m deep, flowing into Sungai Sepatah, a tributary 
to Sungai Mandor; 24 km NE of Pontianak. Lat. 0°07.5'N, long. 
109°30'E. Current moderately swift. Rotenone. 10 July. 

Piintius pentazona. Rashora cephalotaenia. R. dusonensis. R. 
cinihovcni. R. pauciperforala. Stigmatogobiiis sp., Betta ana- 
batoides. B. taeniata, Channa hicius. 

Kapuas 1976-2. Man-made canal. 1-2 m wide and 20 cm-1 
m deep, flowing into Sungai Landok: 5 km E of Pontianak. Lat. 
0°0I'S. long. 109°23'E. Current sluggish to nil. Push net, dip 
nets. 10 July. 

Rasbora dusonensis. Brachygobius doriae. Miigilogobius spp., 
Channa striata. 

Kapuas 1976-3. Sungai Durian. tidal creek 2-6 m wide and 
lOcm-1 m deep (low tide), flowing directly into Kapuas Ketchil; 
7-8 km E of Pontianak. Lat. 0°OrN, long, 109°16'E. Current 
moderate, pH 6. Push net. dip nets. 13 July. 

Rasbora dusonensis. Dermogenys cf orienlalis. Doryichthys 
doekhaloides. Butts gyninoponius. Brachygobius doriae, Miigi- 
logobius sp., Pseudogobiopsis sp., Stigmatogobiiis brocki. 

Kapuas 1976-4, Small stream tributary to Sungai Penju, a 
small coastal drainage; about 43 km NW of Pontianak and 8 
km SW of Andjongan. Lat. 0°18.5'N, long. 109°07'E. Current 
moderate, water brownish tinted, pH 4,5-5. Push net, dip nets. 
13 July. 

Puntius rhontboocellatus. Rasbora pauciperforala. Neohom- 
aloptera johorcnsts. Belontia hasseltii. Betta taeniata. Paros- 
phromenus deissneri. P. parvulus. Sphaerichthys osphrome- 
noides. 

Kapuas 1976-5 (Fig. 2a). Sungai Kepayang, 3-8 m wide and 
maximum 2 m deep, flowing into Sungai Penju; about 44 km 
NW of Pontianak and 7 km SW of Andjongan. Lat. 0°19'N, 
long. I09°08'E. Current moderate, water dark brown tinted, 
blackish when viewed from above, 27°C, pH 4.5-5. Push net; 
rotenone and dip nets. 13 July. 

Osteochilus spihirus. Puntius rhontboocellatus. Rasbora agilis, 
R. axelrodi. R. paucipcrforata. Kiyptopterus macroccphalus, 
Hemirhamphodon pogonognathus. Stigmatogobiiis brocki. Bet- 
ta taeniata. Parosphromenus deissneri, P. parvulus. Sphae- 
richthys osphromenoides. 

Kapuas 1976-6. Sungai Paklehung, a low-lying forested hill- 
stream tributary to Sungai Mempawah, 5 m wide and 1 m deep; 
48 km NNW from Pontianak, 9 km NE of Andjongan, and 1- 
3 km upstream from Toho, Elevation less than 100 m. Lat. 
0°24.5'N. long. 109°13.5'E. Current moderately swift; water clear 
to slightly turbid, 25''C, pH 7; bottom with boulders, cobbles, 
and coarse gravel grading to fine sand. Rotenone. 13 July. 

Crossocheilus oblongus, Cyclocheilichthys armatus, Hatnpala 
macrolepidota, Lobocheilus hispidus. Osteochilus enneaporos, 
Osteochilus hasseiti, O. waandersii, Paracrossochilus vittaius, 
Puntius bmotatus. P. lateristnga. Rasbora caudimaculata. R. 
myersi. R. sarawakensis, Homaloptera nebulosa, Acantophthal- 
mus shelfordi, Mystus nemurus, Silurichthys hasseiti. Clarias 
leiacanthus. Hemirhamphodon phaiosoma. Betta piignax. Mas- 
tacembelus maculatiis. 

Kapuas 1976-7. Sungai Sekilap. tributary to Sungai Mem- 



CALIFORNIA ACADEMY OF SCIENCES 




FioLiRE I. Drainage nclwork of Kapuas and Mempawah rivers, with cnllecling localities of Kapuas survey of \^lb 



pawah; about 64 km N of Pontianak and 38 km by road NE of 
Andjongan. Lai. 0°33.5'N. long. 109022.5'E. Water temp. 24°C. 
pH 7. Rotenone. 14 July. 

Chela Dhuissi. Cyclocheilichthys apogon. Osicochilus hasselli. 
Piiniius tincliisporus. P. binotatus. Rasbora agilis. R. bankaiien- 
sis. R. hnihiiu. R. pauciperforata, AcaiUophthalmus shclfonii. 
Neniachciliis sclangoricus. Hcmirhainphoddn pogonognatlnis. 
Doryichthys marlensii. Pristolepis fasciata. Brachygobnis xan- 
thomelas. Bettataeuiala. Pawsphromenus dcissnen . Mastacem- 
hi'his inaculatus. 

Kapuas 1976-8, Sungai Kenyatan, shaded forest tributary to 
Sungai Landak, 3-7 m wide and 1.5 m deep; about 65 km NE 
of Pontianak and 27 km by road W of Ngabang. Lat. 0°24'N. 
long. 109°44.5'E. Current slow, water turbid, brownish, 27°C, 
pH 6-6.5. Rotenone. 14 July. 

Cychxiu'ilichthys apogon. Ostcochilits hassclti. O. spiliinis. 
Oxygastcr hypophihalmits. Rasbora agilis. R. bankanensis. R. 
ccphalolacnia. Honialoptera nebulosa. .-Icaniflphthaliuus shel- 
fonli. Neinacheilus saravacensis. N. selangoriats. Mystiis mi- 
cracanthus. M. nigriceps. Kryptoplenis macwcephalus. Silii- 
richthys phaiosotna. Henurhamphodon pogonognathiis. 
Xcucuhtdon canciloidcs. Gymnochanda filaiticntosa. Nandiis 
iwhulosus. Pristolepis fasaata. Bella tacn/ata. Parosphromcmis 
deissneri. Lucioccphalus pulcher. Chaiiiia lucius. .Macroguatbus 
aculeatus. Maslacembehis maculatm. 

Kapuas 1976-9. Sungai Landak at Ngabang, large river about 
50 m wide; 83 km ENE of Pontianak, Lat. 0°23.5'N, long. 
1 09°53'E. Current moderate, water turbid, 26°C,pH 7.5; bottom 
coarse sand to gravel. Seine at night. 15 July. 



Clupeichlhys bleekeri. Cyclocheilichlhys heteronema. Oxy- 
gasicr hypopththalmus. Puntiiis bramoides. Rasbora bankanen- 
sis. R. trilmcata. Acanlopsischoirorhynchos. Krypiopterus minor 
(type locality), Laidcs hexauema. Xenenlodon canciloides. Cho- 
ncrhlnos amabilis (type locality). 

Kapuas 1976-10. Sungai Belantian, 20-30 m wide and 1-2 
m deep; 87 km ENE of Pontianak and 6 km by road E of 
Ngabang, Lat, 0°22'N, long, 109°59,5'E, Current swift; water 
clear with brownish tint, 25°C, pH 7.5; bottom sand or gravel 
with some cobblestones, branches and logs. Push net and cast 
net, 15 July. 

Cyclocheilichthys annalus. Rasbora agilis. R. bankanensis. R. 
bntlani, R. pauciperforata. Honialoptera ogilviei. H. tweediei. 
.icanthopsoides gracilis. Neniacheiliis selangoricus. Acrochor- 
donichthys cf melanogasler. Hemirhamphodon pogonognathiis. 
Doryichthys inartcnsii. 

Kapuas 1976-1 1. Swampy, densely overgrown ditch, 1-2 m 
wide and 1 m deep. 87 km ENE of Pontianak, 7 km by road E 
of Ngabang, Lat, 0°22'N, long, 110°00E. Current sluggish or 
ml; water 26°C, pH 6. Push net. 15 July. 

Puntiiis pentazona. Rasbora einthovcnii. Rasbora pauciper- 
forata. Heniirhaniphodon pogonognathiis. Luciocephalus pul- 
cher. 

Kapuas 1976-12. Small forested stream, 2-4 m wide and 20- 
50 cm deep, 91 km ENE of Pontianak and 17 km by road E of 
Ngabang, Lat. 0°19.5'N, long. 1 10°05.5'E. Current moderately 
swift; water clear, slightly brownish, 25°C, pH 6.5; bottom gravel 
or sand, with rocks, logs, and leaf litter. Push net, rotenone, 15 
July. 



ROBERTS-WESTERN BORNEO FISHES 



15 



Rasbora sarawakensis. R. suwalrana, Acantophthahnus shel- 
fordi, Nemacheilus selangoricus. Glyplolhorax major. Hemi- 
rhampbodon phaiosoma. 

Kapuas 1976-13. Sungai Engkonis, small forest stream 2-4 
m wide and 1 m deep. 13 km WNW of Sanggau. Lat. 0°10'N, 
long. 110°28'E. Current moderate; water turbid, 25°C pH 6. 
Push net. 15 July. 

Osteochilus hasseltii, Rasbora bankanensis. R. sarawakensis, 
Homaloptera tweediel. Acantophlhalmus shelfordi. Hemirham- 
phodon sp., Doryichthys maricnsii. Nandus ucbidosus. Mugi- 
logobius sp., Betta piignax. Luciocephalus pidcher, Channa In- 
dus, Mastacembelus notophthalmus (type locality). 

Kapuas 1976-14. Kapuas mainstream and mouth of Sungai 
Sekayam at Sanggau. Lat. 0°07'N, long. 1 10°35'E. Current mod- 
erate; water turbid. Kapuas mainstream 26°C, pH 6.5; Sekayam 
mouth 25°C, pH 7.5. Cast net, floating gill nets, seine. 17 July. 

Clupeichthys bleekeri, Chipeoides hypselosoma, Lycothrissa 
crocoddus, Crossocheihts cobilis. C. oblongiis, Cyclochedichthys 
armatus. Dangila fasciata. D. lineata, Epaheorhynchos kalop- 
terum. Lobochedus sp., Luciosoma spdopleiira, Oxygaster hy- 
pophthalmus, Puntiophtes bulu, P. waandersi, Punlius bra- 
moides, Rasbora borneensis. R. diisonensis. R. myersi. 
Homaloptera tweediei. Ellopostoma megalomycter, Kryptopte- 
nts hicirrhis. K. cryptopterus. K. schdbeides. Laides hexanema, 
Chanda apogonoides. C. wolffii, Paradoxodacna piratica (type 
locality), Toxotes microlepis, Brachygobius xanthomelas, Os- 
phromemis goramy. Cynoglossiis kapuasensis. Achiroides mela- 
norhynchiis. Tetraodou leturus. 

Kapuas 1976-15. Sungai Engkayas where it flows into right 
side of Kapuas mainstream about 2 km upstream from Sanggau. 
Lat. 0°07.5'N, long. 1 10°36'E. Water clear, brown tinted, 27°C, 
pH 7.5; bottom leaf litter and logs. Rotenone. 16 July. 

Clupeichthys bleekeri, Lycothrissa crocoddus, Balantiocheilos 
melanoplenis. Cyclochedichthys heteronema, Dangda lineata, 
Epaheorhynchos kalopterum, Leptobarbus hoevenii, Lobochei- 
lus sp., Osteochilus microcephalus, Oxygaster hypophthalmus, 
Puntioplites bulu, Puntius bramoides. Rasbora dusonensis, R. 
myersi, Rasbora sp., Thynnichthys polylepis, Ellopostoma meg- 
alomycter, Mystus nemurus, Chanda apogonoides, Toxotes mi- 
crolepis, Osphromenus goramy. 

Kapuas 1976-16. Sungai Tekam, small forest stream, where 
it enters right side of Kapuas mainstream about 5-6 km up- 
stream from Sanggau. Lat. 0°06.5'N, long. 1 10°37'N. Water 25°C, 
pH 6.5. Rotenone. 16 July. 

Cyclochedichthys apogon, Dangila fasciata, Eirmoius octo- 
zona, Epaheorhynchos kalopterum, Luciosoma trinema, Osteo- 
chilus borneensis, O. microcephalus. O. spilurus. O. waandersii. 
Oxygaster anomalura. Puntius anchisporus. P. eugrammis. P. 
pentazona, Rasbora agilis, R. argyrotaenia, R. borneensis, R. 
dorsiocellata, R. trilineata, Homaloptera tweediei, Acan- 
tophthalmussemicinctus, .4. shelfordi, A. superbus(type locality), 
Lepidocephalichthys pristes (type locality), I 'aillantella euepip- 
tera, Mystus micracanthus, Mystus nemurus, Silurichthys san- 
guineus (type locality), Wallago leerii, Chaca bankanensis, 
Hemirhamphodon pogonognathus. Doryichthys boaja, Dory- 
ichthys deokhatoides, Monopterus albus, Nandus nebulosus: Oxy- 
eleotris marmorata, Brachygobius xanthomelas. Calanuana sp.. 
Belontia hasseltii. Betta dimidiata (type locality), Betta pugnax, 
Sphaerichthys vaillanti. Trichogaster leerii, T. trichopterus, He- 



lostoma temminckii, Macrognathus aculeatus, Mastacembelus 
erythrotaenia, Achiroides sp. 

Kapuas 1976- 1 7. Several small forest streams where they flow 
into Kapuas mainstream within 10 km upstream from Sanggau. 
Lat. 0°06-07'N, long. I 10°35-38'E. Water 24°C, pH 6-6.5. Ro- 
tenone. 16-17 July. 

Cyclochedichthys apogon. Eirmoius octozona, Luciosoma se- 
tigerum, Osteochilus borneensis, O. intermedins, O. kahajanen- 
sis. O. microcephalus, O. spilurus. Oxygaster anomalura. Pec- 
tenocypris balaena (type locality), Puntius rhomboocellatus, 
Rasbora agilis, R. argyrotaenia, R. bankanensis, Acantophlhal- 
mus semicinctus. A. shelfordi, A. superbus (type locality). Lep- 
idocephalichthys pristes (type locality), I 'aillantella euepiptera. 
Mystus micracanthus, M. nemurus, Glyptothorax major, Clarias 
leiacanthus, Hemirhamphodon pogonognathus. Doryichthys 
deokhatoides. Monopterus albus. Eleotris melanosoma. Oxy- 
eleotris marmorata. Brachygobius xanthomelas. Betta dimidiata 
(type locality), B. pugnax, Trichogaster leerii. Luciocephalus pul- 
cher. Macrognathus aculeatus, Mastacembelus maculatus. 

Kapuas 1976-18. Small swampy stream about 30 km W of 
Sintang on road from Sanggau to Sintang. Approx. lat. 0°00', 
long. 1 1 1°14'E. Push net. 18 July. 

Eirmoius octozona. Puntius eugrammus. Lepidocephalichthys 
pristes (type locality). 

Kapuas 1976-19. Fish market at Sintang. 19 Juiy-1 August. 

Himantura signifer (type locality), Nolopterus borneensis. 
Scleropages formosus. Lycothrissa crocoddus. Amblyrhynch- 
ichthys truncatus, Barbichthys laevis, Cosomochilus falcifer. 
Dangila [estiva, Morulius chrysophekadion, Osteochilus micro- 
cephalus, O. schlegeli, O. triporos. Oxygaster hypophthalmus. 
Botia macracantha, Bagrichthys hypselopterus. B. macropterus. 
Bagroides melapterus, Mystus micracanthus, M. nigriceps, M. 
olyroides (type locality), Belodontichthys dinema. Ceraloglanis 
scleronema. Hemisilurus heterorhynchos, H. moolenburghi. 
Kryptopterus apogon, K. hicirrhis, K. cryptopterus. K. limpok. 
Pangasius humeralis (type locality). P. lithostoina (type locality), 
P. nasuliis. P. polyuranodon, Bagarius yarrelli. Chaca banka- 
nensis, Arius melanochir, A. stormii, Chanda macrolepis, C. 
wolffii, Datnioides microlepis, Pristolepis fasciata. Polynemus 
macrophthahnus. P. midtifilis. Oxyeleotris marmorata. Tricho- 
gaster leerii. Osphromenus goramy, Channa lucius, Cynoglossus 
kapuasensis, Tetraodon palembangensis. 

Kapuas 1976-20. Lower part of Sungai Kebian, a large for- 
ested stream flowing into Kapuas mainstream 5 km upstream 
from confluence of Sungai Melawi. Lat. 0°07'N, long. 1 1 l°32'E. 
Current swift; water clear, tinted dark brown, pH 5.5 Local 
fishermen using "jermal." 19 July-1 August. 

Notopterus borneensis, Balantiocheilos melanopterus, Bar- 
bichthys laevis, Cyclochedichthys janthochir, Dangila ocellata. 
Leptobarbus hoevenii. Luciosoma setigerum, L. trinema. Macro- 
chirichthys macrochir. Osteochilus melanopleura, O. triporos, 
Oxygaster oxygastroides. Puntioplites bulu, Puntius eugrammus, 
Thynnichthys polylepis, Leiocassis micropogon, Mystus nigri- 
ceps. Ompok hypophthalmus. Pseudeutropius brachypopterus. 
Chanda macrolepis. Datnioides microlepis. Trichogaster leerii. 
Channa niarulioides, C pleurophthalmus. 

Kapuas 1976-21. Sungai Belimbing. large forested stream 30 
m wide and 2 m deep, 46 km SSE of Sintang and 15 km by 
road WNW of Nangapinoh. Lat. 0°I8.5'S, long. 1 1 1°38.5'E. 



16 



CALIFORNIA ACADEMY OF SCIENCES 






)>.. 




mf^vsmmmmm^^ 





-A^ 




Figure 2, 
32). 



Kapuas survey of 1976. Habitals. a. Sungai Kcpayang (Kapuas 1976-5); b. lowland forest stream (K.apuas 1976-371; c-d. Danau Piani (Kapuas 1976- 



Water clear, little or no brown tint. Push net in finely divided 
roots of large tree underneath overhanging bank and in riffles. 
20 July. 

Cyclochcilichthys armatus. Osicochilus wicroccphalus. Ras- 
hora agilis, Acantophthalmus anguUlaris, A. shelfonii. Acan- 
thopsoides gracilis. \ 'aillantella euepiptcra. Leiocassis armatus. 
Mystiis nciinirus. Doryichthys martcnsii. Mastacembclus no- 
lopluhalimis (type locality). M. unicolor. 

Rapuas 1976-22. Sungai Melavvi and mouth of Sungai Pinoh 
at Nangapinoh. Lat. 0°19.5'S, long. 1 1 l°44'E. Current moderate 
to strong; water clear to slightly turbid; Melawi mainstream 
28°C. pH 7.5. Pinoh near mouth 28°C, pH 7-7.5. Seine daytime 
and night. 21 and 26 July. 

Chipeichihys bleekeri. Cyclochcilichthys armatus. Luciosoma 
spilopleura. Rashora bankanensis. Homaloptera zollingen, 
Acantopsis choirorhynchos. Acanthopsoides gracilis. Botia mac- 
racantha. Elloposloma megalomyctcr. Kryptopterus nu nor Hype 
locality), A', cryplopterus, Xenentodoii canciloides. Doryichthys 
martensii. Gymnochanda filamentosa. 

Kapuas 1976-23. Sungai Serundung. small forested stream 
flowing into Sungai Melawi mainstream 1-2 km upstream from 
Nangapinoh. Lat. 0°20'S, long. 1 1 1°45.5'E. Local fishermen. 21 
July. 

Rasbora kalochmma. R. paucipcrforata. Bella anabatoides. 

Kapuas 1976-24. Mainstream of Sungai Pinoh 20-60 km up- 
stream from Nangapinoh. Lat. 0°27.5-4l.5'S, long. 1 1 1°39- 



45.5'E. Current swift; water clear. 24-25°C. pH 7.5; bottom with 
gravel, cobbles, or large rocks. Cast nets. 22-26 July. 

Crossocheilus cobitis. C. oblongus. Cyclochcilichthys repasson. 
Epalzeorhynchos kaloptenim. Garra borneensis. Hampala bi- 
maculata. Luciosoma setigcrum. Mystacoleucus marginatus, 
Osicochilus cnneaporos. Paracrossochilus accrus. P. viitatus. 
Puntiiiscollingwoodi. Rasbora baukaiiciisis. R. cicgans. R. myersi. 
R. sarawakcnsis, R. \olzi. Schismatorhynchos heterorhynchos. 
Tor tambroidcs. Gastromyzoii contractus (type locality) G. fas- 
ciatus. G. ridens (type locality), Nemachcilus kapuasensis (type 
locality), Mystus nemurus, Glyptothorax platypogon. G. platy- 
pogonoidcs. Doryichthys martcnsii. Prislolcpis fasciata. Osphro- 
mcniis goramy. Mastacembclus maculatus. M. unicolor. Cho- 
ncrhinos amabilis (type locality). 

Kapuas 1976-25. Sungai Tebelian, small forest stream 3^ m 
wide and 50 cm deep, where it flows into Sungai Pinoh, 19 km 
of S Nangapinoh. Lat. 0°30'S, long. 1 1 1°45'E. Current moder- 
ately swift; water clear, not tinted, 24°C, pH 7.5; bottom sand 
or gravel with cobblestones, logs. Rotenone. 22 July. 

Cyclochcilichthys armatus. Garra borneensis. Hampala bi- 
maculata. Osleochilus cnneaporos. Paracrossochilus vittatus. 
Puntius binotatus. P. laleristnga. Rasbora bankanensis. R. ele- 
gans. R. enncalepis (type locality). R. saranakensis. R. volzi. 
Tor tambroidcs. Gastromyzon contractus (type locality), G. fas- 
ciatus. Homaloptera ncbulosa. Acanthopsoides gracilis. Ne- 
machcilus selangoricus. Ncmacheilus kapuasensis (type locali- 



ROBERTS- WESTERN BORNEO FISHES 



17 



ty), Mystus neiuiinis. Hemirhampbodon phaiosoma. Doiyichthys 
inartensii, Maslacembelus maciilatm. M. unicolor. 

Kapuas 1976-26. Sungai Sekumpai, a small forest stream 3- 
4 m wide and 50 cm deep, where it flows into Sungai Pinoh, 
19 km S of Nangapmoh. Lat. 0°32'S, long. 111°39.5'E. Water 
clear. 25°C, pH 7.5. Rotenone. 22 July. 

Paracrossochi Ills villains. Rasboraelegans. R. ennealepis {type 
locality) R. sarawakensis. Gasiromy:on fascialiis. Homalopiera 
iidnilosa, Nemacheihis kapiiasensis (type locality), Silunchihys 
hasselli, Hemirhampbodon phaiosoma. 

Kapuas 1976-27 (Fig. 3c). Rocky channel in mainstream of 
Sungai Pinoh, 37 km S of Nangapinoh. Lat. 0°39.5'S, long. 
1 1 1°40'E. Current swift; water clear, 25°C, pH 7.5. Rotenone. 
cast net. 24 July. 

Baribiis borneensis (type locality), Cwssocheilus cobilis. C. 
oblongiis. Epalzeorhynchos kaloplenim. Hampala macrolepi- 
dola, Myslacoleucus marginatiis. Osleochihis eniieaporos. O. 
nncrocephabis. Paracrossochilus villatiis. Pimlius cotbngwoodi. 
Rasbora bankanensis. Rasbora cf ennealepis. Schismatorhyn- 
chos heterorhynchos. Gyhnocheilus ptisliilosus. Gastwmyzoii 
contractus (type locality), G. fascialiis. Homalopiera nebidosa. 
H. ophiolepis, H. sicphensoni. H. zolbngeri. Botia reversa (type 
locality), Nemacheilus maculiceps (type locality), Nemacheihis 
kapiiasensis. Leiocassis micropogon. Mystus nemurus, Glypto- 
ihoraxplatypogon. Pnstolepisfasciata. Maslacembelus unicolor. 

Kapuas 1976-28. Small forested streams flowing into Sungai 
Pinoh near village of Ribang-Rabing, about 55 km SSE of Nan- 
gapinoh and 2 km NE of Kotabahru. Lat. 0°47'S, long. 1 1 l°33'E. 
Push net; most fishes caught by villagers using herbigenous 
ichthyocides. 24 July. 

Cyclocheilichthys apogon. Osteochilus kappenii, Rasbora ban- 
kanensis, R. sarawakensis. Leiocassis micropogon. Mystus ne- 
murus, Kryptoptcriis sp., Doryichthys martensii, Pristolepis fas- 
ciala, Maslacembelus maculalus, M. nolophthalmus (type 
locality). 

Kapuas 1976-29 (Fig. 3a). Rocky channel in mainstream of 
Sungai Pinoh at Nanga Saian, 45 km S of Nangapinoh. Lat. 
0°43'S, long. 1 1 1°38.5'E. Current moderate; water tinted dark 
brown, 25°C, pH 7. Rotenone. 26 July. 

Crossocheiliis cobilis. Epalzeorhynchos kalopierum. Osteo- 
chilus enneaporos. O. microcephaliis. Paracrossochilus acerus, 
P. vitlatus, Rasbora bankanensis, Schismatorhynchos hetero- 
rhynchos. Tor tambroides, Homalopiera nebidosa. H. slephen- 
soni, H. zollingcri. Acanlophthalmiis oblongiis. Botia macra- 
canlha, Nemacheilus maculiceps. Nemacheilus kapiiasensis, 
I 'aillantella maassi, Mystus nemurus, M. wyckii, Kryptopterus 
minor (type locality), Doryichthys martensii, Osphromeniis go- 
ramy. Maslacembelus unicolor. 

Kapuas 1976-30. Sungai Tamang. small forested stream with 
rocky bottom flowing into Sungai Pinoh opposite mouth of 
Sungai Kelawai. Lat. and long, uncertain due to irreconcilable 
map sources, approx. 0°35'S, 1 1 1°44'E. 26 July. 

Cyclocheilichthys armalus. Luciosoma setigerum. Paracros- 
sochilus vitlatus. Puniiiis latensiriga. Rasbora volzi. Rasbora 
sp.. Tor tambra. Gastromyzon fascial us. Homalopiera nebidosa, 
H. orthogoniala. Hypergastromyzon humilis (type locality), 
Mystus nemurus. Glyptothora.x platypogon. Clarias leiacanthus. 
Hcniirhamphodon phaiosoma. Bella anabatoides. Channa ori- 
cntalis. Maslacembelus maculalus. 

Kapuas 1976-31 (Fig. 3d). Bar in mouth of Sungai Mclawi at 



Sintang. Lat. 0°35'S, long. 1 1 1°29'E. Bottom gently sloping, fine 
sand to coarse gravel. Seine at night. 29 July. 

Crossocheiliis cobilis, C. oblongiis. Dangila ocellala. Lucio- 
soma spiloplcura. Osteochilus microcephaliis. Puntius hra- 
moides. Rasbora bankanensis, R. borneensis, R. myersi, R. tri- 
lineata, Bagariiis yarrelli, Akysis pseiidobagariiis (type locality), 
Clarias leiacanthus. Paradoxodacna piratica (type locality). 

Kapuas 1976-32 (Fig. 2c, d). Danau Piam near Ketungau, 38 
km NNE of Sintang. Lat. 0''23.5'N, long. 1 1 1°37.5'E. Water 
26°C, pH 5.5-6. Seine, cast net, rotenone. 5-6 August. 

Cyclocheilichthys armalus. C. janthochir. Leptobarbiis mela- 
noplerus. Osteochilus spilurits, Oxygastcr hypophthalmiis, O. oxy- 
gastroides, Peclenocypris balacna (type locality), Puntius lin- 
eatiis, P. rhomboocellatus. Rasbora dorsiocellata. R. myersi, R. 
pauciperforata. R. subtilis (type locality), Homalopiera stephen- 
soni. Lepidocephalichthys pristes. Afystus micracanlhus. Ompok 
eugeneialiis. Pseiideiitropius brachypopteriis. P. moolenburghae. 
Xenenlodon canciloides, Bella dimidiata (type locality), Sphae- 
richlhys vaillanii, Trichogaster leerii, Channa pleurophthalma, 
Cynoglossus waandersi. 

Kapuas 1976-33. Kapuas mainstream, Danau Mawan, and 
"Danau Bahru" near Kampong Nibung, about 100 km NE of 
Sintang and 7 km NE of Selimbau. Lat. 0''39'N, long. 1 12°I0.5'E. 
Kapuas mainstream 26°C, pH 5.5-6, Danau Mawan 33-34°C, 
pH 8-9.5. Many specimens caught by fishermen of Nibung using 
cast nets and life nets; seine at night; gill nets. 5-7 August. 

Clupeichthys hleekeri, Lycolhrissa crocodilus, Sctipinna mela- 
nochir. Siindasalanx microps (type locality), Amblyrhynchich- 
thys iruncatus. Barbichthys laevis. Cyclocheilichthys helerone- 
ma. C. repasson. Cyclocheilichthys sp., Dangila ocellala. 
Epalzeorhynchos kalopierum. Hampala macrolepidoia. Lucio- 
soma trinema, Osteochilus borneensis. O. microcephaliis. O.xy- 
gaster hypophthalmiis. Puiitioplites bulu. P. waandersi, Puntius 
bramoidcs, Rasbora bankanensis. R. borneensis. R. myersi, Ras- 
bora sp., Rasborichthys helfrichii. Rohteichlhys microlepis. 
Thynnichlhys polylepis. T. thynnoides. Acanthopsoides gracilis. 
Botia macracantha, Mystus micracanlhus. M. nigriceps. Kryp- 
topterus bicirrhis, K. cryptopterus. K. lais. K. micronema. K. 
schilbeides. Ompok hypophthalmiis. Pseiideulropius brachypop- 
teriis. P. moolenburghae. Xenenlodon canciloides. Channa apo- 
gonoides. C. macrolepis, Paradoxodacna piratica (type locality), 
Toxoles microlepis, Polynemus nndtifilis, Trichogaster leerii. 
Helosioma temminckii. Osphronemus goramy. Macrognathus 
acidealus. M. erythrotaenia. Achiroides melanorhynchus. Cho- 
nerhinos amabilis (type locality), C. nefastiis (type locality). 

Kapuas 1976-34. Kapuas mainstream 53 km W of Putussi- 
bau. Lat. 0°46.5'N, long. 1 1 2°27.5'E. Water 26°C, pH 6.5. Seine, 
gill nets. 6-7 August. 

Cyclocheilichthys heleronema. C. microlepis. Dangila lineata, 
D. ocellala. Kalimanlania lawak. Oxygasler hypophthalmiis. O. 
oxygastroides. Puntioplites bulu, Puntius bramoides, Rasbora 
myersi. Afystus nigriceps. Kryptopterus bicirrhis. Paradoxodacna 
piratica (type locality), Cynoglossus waandersi. Chonerhinos 
amabilis (type locality), C nefastiis (type locality). 

Kapuas 1976-35. Kapuas mainstream about 23 km WSW of 
Putussibau. Lat. 0°48'N, long. 1 12''45'E. Water 25°C, pH 7-7.5. 
Seine, gill nets. 8-9 August. 

Noiopieriis borneensis. Clupeichthys bleekeri. Lycolhrissa 
crocodilus. Crossocheilus oblongiis. Cyclocheilichthys helerone- 
ma. Dangila lineata. Osteochilus microcephaliis, Oxygasler hy- 



CALIFORNIA ACADEMY OF SCIENCES 




hiuLiKt i. kapuas survey ol IVTb. Habitats, a, Sungai Pinoh al Nariga Saian (Kapuas 1*^76-29), b, Sungai Pinoh (Kapuas 1976-27); c, Sungai Tawang (Kapuas 
1976-44); d, Kapuas mainstream at Sintang, gravel bar in mouth of Sungai Melawi at upper left (Kapuas 1976-31). 



pophthalmus. Rasbora bankanensis. Acantopsis choirohynchos. 
Chaniia apogonoides. Chonerhinos amabilis (type locality), C. 
nefastus (type locality). 

Kapuas 1976-36. Kapuas mainstream, 6 km WofPutussibau. 
Lat. 0°50.5'N, long. 1 12°52'E. Seine. 9 August, 

Cyclochi'ilichthys repasson. Epaheorhynchos kaloptenim. Os- 
teochilus waandcrsi. Rasbora bankanensis. R. myersi. Thrys- 
socypris sniuragdinus (type locality), Acanlopsis choirorhynchos. 
Botia hyiucnophysa. Mystus micnicanlhus. Clianda apogo- 
noides, Chonerhinos amabilis (type locality). 

Kapuas 1976-37 (Fig. 2b), Small forested stream, 3-5 m wide 
and 1 m deep, where it flows into Sungai Mandai 2-3 upstream 
from its confluence with Kapuas mainstream, 17 km WSW of 
Putussibau. Lat. 0°47'N, long, 112°48'E. Current weak, water 
dark brown, turbid, 30°C, pH 6; bottom with logs and leaf litter. 
Rotenone. 10 August. 

Rurbichthys laevis. Chela niaassi. Cycloehcilichthys apogon. 
C. helcroneiua, Dangila fasciaia. D. lineala. Oskvchiliis kap- 
peni. O. mierocephaliis. O. spihirus, Oxygasrer anomaliira. O. 
hypophthalmus. Puntius anclusponis. P. bramoides. P. endeca- 
nalis (\ype\oca\\Xy). Rasbora bankanensis. R. brittani. R. myersi. 
R trilincata. Homaloplera tweediei. Acantophlhahmis shelfordi. 
Acanihopsoides gracilis. Barbucca diabolica (type locality), El- 
loposloDia megalomyeier. Nemacheilus selangoriciis. I 'aillan- 
lella euepiplera. Myslns micracamhiis. Hemirhamphodon po- 
gonognarhiis. Xenentodon canciloides. Pristolepis fasciaia, Bella 
piigna.x. Osphonennis goramy. Achiroides mehinorhynchus. 
Chonerhinos amabilis (type locality). 



Kapuas 1976-38. Mouth of Sungai Mandai, 17 km WSW of 
Putussibau. Lat. 0°48'N, long. 112°47'E. Current swift, water 
dark brown, slightly turbid. Hook and line. 10 August. 

Kryplopicrus limpok. 

Kapuas 1976-39. Sungai Mandai Ketchil near its confluence 
with Kapuas mainstream, 18 km WSW of Putussibau. Lat. 
0°48'N, long. 1 12°47'E. Rotenone. 1 1 August. 

Cyclocheilichthys repasson. Dangila fasciaia. D. lineala, D. 
ocellata. Osleochilus kahajanensis. O. mierocephaliis. O. waan- 
dcrsi. Oxygaslcr anomaliira. Puntius anchisporiis. P. endecanalis 
(type locality). Rasbora bankanensis. R. brlliani. R. myersi, R. 
sumatrana, R. irilineala, Homaloplera tweediei. Acanlophlhal- 
miis semlcincliis. A. shelfordi. Acanihopsoides gracilis. Barbucca 
diabolica (type locality), Nemacheilus selangoricus, I 'alllaniella 
euepiplera. Leiocassis armalus, Afyslus micracanlhiis, Ompok 
sabaniis. Parakysis anomalopteryx (type locality), Hemirham- 
phodon pogonognalhus. Xenenlodon canciloides. Nandus neb- 
ulosus. Calamiana sp., Bella dimidiala (type locality), Bella 
piignax. Channa luciiis. Macrognalhus aculealus, Achiroides 
Icucorhynchus, Chonerhinos nefaslus (type locality). 

Kapuas 1976-40, Kapuas mainstream near Telokabik, 29 km 
WofPutussibau. Lat. 0°50.5'N, long. 112''40'E. Rotenone. 11 
August. 

Oxygasler hypophihalmus. Punliiis bramoides. Rasbora ban- 
kanensis, R. borneensis, R. sublilis (type locality), Ariiis melano- 
chir. Doryichlhys boaja, Achiroides leucorhynchus. A. melano- 
rhynchus. Chonerhinos nefastus (type locality). 

Kapuas 1976-41. Oxbow Lake at Nangaembaluh with direct 



ROBERTS-WESTERN BORNEO FISHES 



19 



connection to Kapuas mainstream; 31 km W of Putussibau. 
Lat. 0°50'N, long. 1 12°39'E. Local fishermen. 12 August. 

Lycothrissa crocodllus. Cyclocheilichthys heteronema, Oxy- 
gaster hypophthalmus, Rasborichlhys helfrichii. Thynnichthys 
polylepis. 

Kapuas 1976-42. Sungai Seriang, forest tributai^ of Sungai 
Palin, 5-10 m wide and 2 m deep, 37 km W of Putussibau and 
3-5 km up Sungai Palin from Kapuas mainstream. Approx. lat. 
0°51.5'N, long. 112°36'E. Current swift; water clear, reddish- 
brown tinted. 30°C, pH 6.5; many trees fallen across and into 
stream, bottom sand. Rotenone. 12 August. 

Chela maassi. Crossocheilus sp., Cyclocheilichthys apogon. 
Osteochilus intermedius, O. spilums. O. trtporos. Rasbora ar- 
gyrolaenia. R. bankanensis. R. bhttanl. R. dorsiocellata. R. tri- 
lineata. Barhucca diabolica (type locality), Lepidocephahchthys 
phsles (type locality). Nemacheilus selangoricus. Vaillantella 
euepipiera, Leiocassis myersi (type locality), Mystus micracan- 
thus, M. nemurus, Ompok hypophthalmus, Silurichthys phaio- 
soma. Parakysis anomaloptery.x (type locality), Chaca banka- 
nensis. Xenentodon canciloides. Calamiana sp., Betta dimidiata 
(type locality), Bella piignax. Trichogaster leerii. Macrognathus 
aciileatus. Mastacembelus macidatus. .Achiroides leucorhynchus. 

Kapuas 1976-43. Small oxbow lake completely cut off from 
Kapuas mainstream opposite Empangau, 124 km NE of Sin- 
tang. Lat. 0°44'N, long. 1 1 2°23'E. Water clear, dark brown, 30°C, 
pH 6-6.5; bottom mud, with many tree branches. Rotenone. 
13 August. 

Sclewpages formosus. Chela maassi. Cyclocheilichthys repas- 
son. Dangila ocellata. Epalzeorhynchos kaloptcnim. Osteochilus 
intermedius. O. keppeni, O. micwcephalus. Punlius anchisponis. 
P. endecanalis (type locality), Rasbora agilis, R. argyrotaenia, 
R. brittani. R. dorsiocellata, Acanthopsoides gracilis, Mystus mi- 
cracanthus, M. nigriceps, Nandtis nebulosiis. Pristolepis fasciata, 
Betta dimidiata (type locality), Sphaerichthys vaillanti. Tricho- 
gaster leerii. Helostoma temminckii. Channa striata. 

Kapuas 1976-44. Sungai Tawang near Danau Pengembung. 
Lat. 0°48'N, long. 1 12°03'E. Current slow; water turbid, cafe- 
au-lait, 26°C, pH 6 (early morning). Local fishermen, cast nets 
and "jermal." 14-15 August. 

Lycothrissa crocodllus, Amblyrhynchichthys truncatus. Bal- 
antiocheilos melanopterus, Barbichthys laevis, Cyclocheilichthys 
apogon. C. armatus. C. heteronema. C. microlepis. Dangila ocel- 
lata. Leptobarbiis hoevenii, Lobocheilus sp., Macrochirichthys 
macrochir, Osteochilus schlegeli, Oxygaster hypophthalmus, O. 
oxygastroides, Puntioplites bulu, Pimtius anchisporus, Rohteich- 
thys microlepis. Bagrichthys hypselopterus. Mystus micracan- 
thus. M. nigriceps. Kryptopterus bicirrhis. K. micronema. Ompok 
eugeneiatiis. O. hypophthalmus. Pseudeutropius brachypoptenis. 
P. moolenburghae. Xenentodon canciloides. Chanda woljfii, 
Paradoxodacna piratica (type locality). Polynemus multifilis, 
Oxyeleotris marmorata, Trichogaster leerii, Achiroides melano- 
rhynchus, Chonerhinos nefastus (type locality), Tetraodon pa- 
lembangensis. 

Kapuas 1976-45. Kapuas mainstream 58 km NE of Sintang 
and 1 km downstream from Sebruang. Lat. 0°25.5'N, long. 
1 1 1°52.5'E. Current moderate; water very turbid, cafe-au-lait, 
28°C, pH 6.5; bottom probably muddy, but with numerous large 
rocks or boulders. Rotenone. 16 August. 

Notopterus borneensis, Osteochilus microcephalus, Puntio- 
plites bulu. Rasbora borneensis. Botia macracantha, Lepido- 
cephalus spectrum (type locality), Nemacheilus lactogeneus (type 



locality), Nemacheilus cf longipectoralis. Vaillantella maassi. 
Bagrichthys macropterus. B. micranodus (type locality), Mystus 
wyckii, Kryptopterus bicirrhis. Breitensteinia cf insignis. Chanda 
apogonoides. Toxotes microlepis. 

Kapuas 1976-46. Sungai Gentu, small forest stream, near 
where it flows into Kapuas mainstream, 55 km NE of Sintang. 
Lat. 0°27'N, long, i 1 1°49'E. Current swift; water clear, brown 
tinted, 25°C, pH 6-6.5; bottom sand, many trees fallen across 
stream. Rotenone. 16 August. 

Osteochilus microcephalus. O. spilwus. Rasbora argyrotaenia. 
R. bankanensis. R. dorsiocellata. R. trillneata. Barbucca dia- 
bolica (type locality), Vaillantella euepipiera. Leiocassis myersi 
(type locality), Mystus nemurus, Wallago leerii. Pseudeutropius 
moolenburghae. Hemirhamphodon pogonognathus. Xenento- 
don canciloides. Nandus nebidosus. Calamiana sp., Betta dimid- 
iata (type locality), Mastacembelus macidatus. 

Kapuas 1976-47. Small forest stream flowing into Kapuas 
mainstream NE of Gunung Setunggul, 53 km SW of Sintang 
and 10 km NW of Silat. Lat. 0°24'N, long. 111°51'E. Current 
moderately swift; water clear, brown tinted, 26°C, pH 6.5. Ro- 
tenone. 16 August. 

Osteochilus Intermedins. O. microcephalus. O. spilurus. O. 
waandersi. Rasbora agilis. R. argyrotaenia, R. bankanensis, R. 
dorsiocellata, R. trillneata. Homaloptera tweediel. .icantoph- 
thalmus oblongus, A. semicinctus. Barbucca diabolica (type lo- 
cality), Lepldocephalichlhys pristes (type locality), Nemacheilus 
selangoricus. Vaillantella euepipiera, Leiocassis myersi (type lo- 
cality), Glyptothorax major. Parakysis anomalopteryx (type lo- 
cality), Chaca bankanensis. Xenentodon canciloides. Gymno- 
chanda lilamentosa. Nandus nebulosus. Pristolepis fasciata. 
Oxyeleotris martnorata, Brachygobius xanthomelas, Calamania 
sp., Betta dimidiata (type locality), Macrognathus aculeatus, 
Mastacembelus maculatus, Achiroides melanorhynchus. 

Kapuas 1976-48. Kapuas mainstream opposite Silat, rocky 
area on left bank. Lat. 0''20.5'N, long. 1 1 1°47'E. Current swift; 
water ver>' turbid, cafe-au-lait, 30°C, pH 6.5. Rotenone. 1 7 Au- 
gust. 

Lepidoccphalus spectrum (type locality), Bagroides melapte- 
riis. Mystus wyckfii. Hemisilurus heterorhynchos, Arius melano- 
chir, Polynemus multifilis. .Achiroides melanorhynchus. Cyno- 
glossus waandersi. Chonerhinos modestus. 

Kapuas 1976-49. Sungai Djentawang, forest stream 5-10 m 
wide and 3-4 m deep near Ketungau, 37-38 km NNE of Sintang. 
Lat. 0°23'N, long. 111°37.5'E. Current swift; water clear, dark 
brown, 30°C, pH 6. Rotenone. 17 August. 

Balantiocheilos melanopterus. Cyclocheilichthys armatus. C. 
heteronema. C. janthochir. Dangila ocellata. Leptobarbiis hoev- 
enii. L. melanopterus. Luciosoma trinema. Macrochirichthys 
macrochir. Osteochilus melanopleura. O. triporos. Oxygaster hy- 
pophthalmus. O. oxygastroides. Puntioplites bulu. Thynnichthys 
polylepis. Bagrichthys micranodus (type locality), Chanda mac- 
rolepis. 

Kapuas 1976-50. Rocky ledge in middle of Sungai Melawi 
near confluence with Kapuas mainstream, about 0.5 km up- 
stream from Sintang. Lat. 0°04.5'N, long. 1 1 1°28'E. Current 
swift; water clear or slightly turbid, dark brown. Rotenone. 18 
.'August. 

Luciosoma spilopleura. Rasbora borneensis. Homaloptera 
ophlolepis. Lepldocephalus spectrum (type locality), Nema- 
cheilus macullceps (type locality), Vaillantella maassi. Mystus 
wyckii, Chanda apogonoides, Achiroides melanorhynchus. 



20 



CALIFORNIA ACADEMY OF SCIENCES 



Kapuas 1976-51. Small forest stream. 3-5 m wide and 50 cm 
deep, about 1 km up Sungai Tajan from Tajan, 87 km E of 
Pontianak. Lat. 0°02'S, long. 1 10°07'E. Current sluggish; water 
moderately turbid, dark brown. 28°C, pH 6.5; bottom mud, 
with leaf litter and logs. Rotenone. 22 August. 

Eirmotus octozona. Osteochilm spilurus. Rasbora agllls, R. 
argyrotaenia. R. bankanensis. R. dorsiocellata. R. diisonensis. 
Acantophthalmus semianctiis. A. siiperbiis {type locality), Acan- 
thopsoides gracilis. Nemacheihis sc/angoriciis. Mystus micra- 
canlluis. Clanas inclanodenua. Hcmirhamphodon pogono- 
gnathus. Xeiwntodon canciloides. Doryichthys deokhatoides. 
Nandus nebulosus, Oxyeleotris urophthahuoides. Pseudogobiits 
sp., Betta dimidiata (type locality), Bella piignax, Parosphw- 
mcnus deissncri. Trichigaster Iccrii. Channa Indus. Macrogna- 
ilnis aadeatiis. Maslacembclus niaculatus. Tctraodon paleni- 
bangensis. 

Kapuas 1976-52. Pontianak fish market. Specimens from a 
single catch, said to be from Sungai Kapuas Ketchil. 23 August. 

Sclipinna luclanochir. Cosmochdus falcifcr. Chanda wolffii. 

Kapuas 1976-53. Market at Kampong Djungkat (near mouth 
of Sungai Kapuas Ketchil, 1 7 km WNW of Pontianak). 1 3 July. 

Codia coomami. Amblyrhynchichthys truncatus. 

Kapuas 1976-54. Kapuas mainstream and Sungai Ketungau 
near Ketungau. Obtained from local fishermen by fisheries of- 
ficer Ponejo. July-August. 

Himantura signifer Hype locality), Andi/yrhyiichichiliys trun- 
catus. Cyclocheilichlhys microlepis. Acluroides inelauorhyuchus. 
Cynoglossus kapuascnsis. 

Kapuas 1976-55. Kapuas mainstream at Selimbau, 92 km 
NE of Smtang. Lat. 0°36.5'N, long. 112°07'E. Water turbid. 
26°C, pH 6.5. Dip netting at night with flashlight from floating 
house. 4 August. 

Crflssocheihts i^blmigus. Rasbora bankanensis. R. borneensis. 
R. sp., Krypiopterus minor (type locality). A', lais. .Achiroides 
niekmorhynclius. 

Systematic Account 

The present account includes all species of freshwater fishes 
known from western Borneo or Kalimantan Barat. The only 
other region of Borneo for which a monographic study of the 
freshwater ichthyofauna is available is northeastern Borneo (In- 
gcr and Chin 1962). In recent years material has been collected 
from the Rejang and Baram basins which should provide the 
basis for a reasonably comprehensive report on the freshwater 
fishes of northern Borneo or Sarawak. The fishes of eastern and 
southern Borneo (Kalimantan Timor and Selatan) have not been 
sufficiently collected to make a modern synthesis worthwhile; 
they are known chiefly through the researches of Bleeker, Vail- 
lant (1902), and Popta (1906). Thus the present work requires 
two sequels to provide a reasonably thorough systematic ac- 
count of the freshwater fishes of the entire island of Borneo; one 
on the fishes of northern Borneo, for which a great deal of 
material has been collected and is deposited in North American 
museums; and one on southern and eastern Borneo, for which 
extensive field surveys need to be conducted in areas still poorly 
known or completely unexplored ichthyologically. 

While it has not been possible for me to provide an account 
of the freshwater fishes of all of Borneo, western Borneo is 
ichthyologically the richest part of the island and probably in- 



cludes at least half of all Bomean freshwater fish species. I have 
examined western Bomean material of almost every species 
reported or known from there, and the basis for inclusion of 
each species is indicated clearly. Some species have been in- 
correctly reported from western Borneo by previous authors; I 
have attempted to correct all such errors by examining and re- 
identifying the actual specimens involved. Type specimens of 
many species have been examined in order to verify identifi- 
cations, and a great deal of extralimital material has been ex- 
amined to differentiate species, investigate systematic problems 
including geographical variation, and verify distributions. 

Before proceeding to the factual systematic accounts of taxa, 
a number of features should be drawn to the reader's attention. 

Classification.— The formal taxonomic categories utilized 
herein are mainly family, genus, and species. Subfamilies or 
other categories are occasionally discussed, especially when I 
have new information concerning their classification or when 
they are relevant to biogeography. Categories above family level 
are rarely discussed or employed herein, but there is one im- 
portant exception I wish to emphasize. This is especially rele- 
vant to students offish evolution and distribution in southeast 
Asia, and has eflfected the sequential listing of families employed 
herein. Sundaland has apparently been the main evolutionary 
center of the labyrinth fishes or anabantoids, or at least a very 
important late evolutionary center for the group. Contrary to 
Berg (1940) and the long-held opinions of Liem (1963 et seq.), 
I belie\e that the snakehead family Channidae (formerly Ophi- 
cephalidae or Ophioccphalidae) and the pike-like family Lucio- 
cephalidae are integral parts of the anabantoid radiation. I have 
long held this view concerning Luciocephalidae and discussed 
it with Liem on several occasions while we were colleagues at 
Harvard; he recently placed Luciocephalidae within Anaban- 
toidea (Lauder and Liem 1983). The full extension of the ex- 
traordinarily protrusible jaws of Luciocephalidae, observed by 
me on living specimens in Malaysia in 1 970 and communicated 
to Liem, has been reported by Lauder and Liem ( 1 98 1 ). On the 
other hand, Liem has maintained steadfastly that snakeheads 
are not anabantoids without proposing a well defined alternative 
hypothesis of their relationships. At the time of our discussions 
of Luciocephalidae I had not given serious consideration to the 
problem of snakehead relationships. Having now reviewed the 
biology and examined radiographs and ostcological prepara- 
tions of many anabantoids and Channidae, I feel the best avail- 
able hypothesis is that snakeheads are anabantoids. The func- 
tional significance of the posterior extension of the swim bladder 
present in all anabantoids may be to provide an adjustable 
counterpoise to the airbubble in the suprapharyngeal respiratory 
organ (W. C. Freihofer. pers. comm.). I predict that evidence 
supporting the broad phyletic relationships among anabantoids 
outlined here will accumulate in proportion to comparative ob- 
servations on their morphology, development, physiology, and 
behavior. The recent observation that Channa striata is an oral- 
brooder (Ettrich 1982) is relevant. I suggest that this is further 
evidence of the anabantoid relationships of Channidae. and that 
detailed observations of its oral-brooding and comparisons with 
oral-brooding in other fishes will bear this out. The distinctive 
spawning embrace characteristic of numerous anabantoids ap- 
parently also occurs in Channa. 

A few additional comments concerning recognition of families 



ROBERTS-WESTERN BORNEO FISHES 



and their sequential arrangement should be noted. The entire 
external body surface of Sisoridae and Akysidae is covered by 
specialized unculiferous plaques which appear to be basically 
identical and have not been observed in any other catfishes 
(Roberts 1982«). indicating that the two families are closely 
related. Perhaps Akysidae (comprising three genera which un- 
doubtedly form a monophyletic unit) should be placed as a 
subfamily of Sisoridae, but for now it is retained as a separate 
family. Parakysis, a Sundaic genus hitherto placed in Akysidae, 
seems not closely related to Akysidae or any other family; due 
to an extraordinary number of unique specializations and no 
known uniquely shared specializations with any other family, 
it is placed in a new family of its own, Parakysidae. Elloposloma. 
a Sundaic genus of previously doubtful family relationships 
(Roberts 1972; Nelson 1984) is referred to Cobitidae. This pe- 
culiar and highly specialized genus cannot be placed in any of 
the four currently recognized cobitid subfamilies, and further 
observations on its osteology and biology should be made. The 
rather generalized percoid Datmoidcs. traditionally placed in 
the tripletail family Lobotidae, is recognized as the sole genus 
in the family Datnioididae. Datnioididae superficially resembles 
the leaf-fishes or Nandidae and may be related to them but is 
much more generalized. There seems to be no valid evidence 
in support of its relationship to Lobotidae. 

Workers familiar with systematics of Asian freshwater fishes 
will find classification at the genus level presented here is gen- 
erally conservative. In particular several large "traditional" or 
"catch-all" genera are employed which almost certainly are 
polyphyletic. Some of them may in fact include several genera; 
such are Barilius. Piintius. Rashora. Nemacheilus. Mystiis. and 
Arius. For all of these, additional genera have been proposed or 
recognized by various authors, but most are poorly defined or 
based on such superficial characters that it does not seem worth- 
while to recognize them at present. In some instances what may 
be distinct genera have been recognized and reasonably well 
defined, but many species are so poorly known that they cannot 
be assigned to these genera. Thus, Howes (1980) recently split 
Rasbora into Rashora and Parluciosoma but was able to assign 
only a few of the species involved to each genus. The situation 
in Rasbora is further complicated because it may include ad- 
ditional groups which should be recognized as separate genera 
(see discussion of Rasbora axehvdi herein). Thus it seems the 
most reasonable course is to leave all of the species in Rasbora 
until a more definitive study has been completed. 

A similar situation exists in Myslus. which Bleeker (1862) 
split into four genera we probably should recognize. But mor- 
phological study of many poorly known species will be needed 
before these genera can be defined adequately and most species 
assigned to them, and the problem of generic typification of 
Mystiis itself has to be resolved. Arius probably represents the 
most challenging problem of generic classification posed by any 
large group of catfishes. Some specialized or very distinctive 
ariids have been placed in separate genera, but a large number 
of species are morphologically generalized or poorly known and 
attempts to distinguish them generically from Arius have very 
little scientific basis. Thus the genera Ccphalocassis and Hemi- 
arius proposed by Bleeker for the species Arius melanoehir and 
Arius stormii are not recognized because adequate generic def- 
initions cannot be provided for them at this time. I have tried 
to list all of the subjective generic synonyms or at least all of 



those relevant to southeast Asia for each traditional or catch- 
all genus employed herein. 

Generic Typification. — Virtually any systematic decision 
involving a genus requires a knowledge of its type species. In 
studying Asian freshwater fishes I have repeatedly found that 
secondary statements in the literature regarding type species 
are untrustworthy. Statements about type species that do not 
include the nature of type species determination and the relevant 
page references simply cannot be relied upon, and in any case 
cannot be verified readily. 

In order to clarify the systematic status of genera, and to 
minimize confusion and needless duplication of effort by other 
workers, I have documented the type species of every genus 
insofar as practical. The following methods of type species de- 
termination or fixation are recognized: original designation, 
monotypy, absolute tautonymy, and subsequent designation (in- 
cluding subsequent monotypy and type species selection by the 
ICZN). Workers unfamiliar with these procedures should con- 
sult the latest edition of the Code. I am happy to report that 
type species have been determined for nearly all included genera. 
The most notable exception is that of Afystus Scopoli, 1777, 
typification of which probably will require formal action by the 
ICZN. In documenting generic typification, especially of older 
names and junior synonyms, it is dilTicult not to overlook key 
references. This is especially so for fixation by subsequent des- 
gination, the references to which are often obscure. In such 
instances it is particularly important to cite the relevant page 
references. I hope that I have made few errors of omission of 
this sort, but in any event the statements herein are documented 
so that they can be readily checked. 

Failure to take type species into account can lead to unfor- 
tunate lapses. Thus numerous Asian loaches referred to Lcpi- 
doccphalus are not congeneric with its type species. The loaches 
in question should be referred to Lcpidocephalicluhys. 

The utility of systematically oriented anatomical studies (such 
as that on cobitid genera by Sawada 1982) could be significantly 
enhanced by including observations on type species of as 
many genera as possible. Such observations immediately place 
the anatomical observations on a sounder taxonomic basis and 
are helpful to other workers concerned with defining the genera. 

Synonymies. — Generic and species synonymies presented in- 
clude primary nomenclatural references (i.e., original publica- 
tions of senior and junior objective and subjective synonyms), 
especially when directly relevant to the systematics of fishes in 
Borneo. In addition important re-identifications or locality rec- 
ords have been included in many synonymies, especially when 
necessary to document inclusion or exclusion of a particular 
species in the western Borneo ichthyofauna. This is particularly 
true for poorly known species or for the few species of which 
western Bomean specimens have not been examined by me. 

Keys. — Keys have been provided for many groups, including 
the important food fishes Cyclochcilichthys and Osteochilus. re- 
cently revised by my Thai colleagues Suebsin Sontirat and Ja- 
ranthada Karnasuta, respectively, in their unpublished Ph.D. 
dissertations. I originally intended to include keys for all of the 
larger and more dilTicult groups but exigencies of time and prac- 
tical considerations forestalled this plan. Nevertheless the in- 
formation provided herein, supplemented with a few references, 
should facilitate identification of all freshwater fish species known 
from western Borneo. 



22 



CALIFORNIA ACADEMY OF SCIENCES 



Distribution. — Information on species distribution is given 
in the material examined, in summary descriptions under the 
hcadmg Distributions or as brief statements in the body of the 
species account, and in selected instances, on distribution maps. 
In many instances extralimital material has been examined for 
the express purpose of verifying or adding to information on 
species distributions. The information given under the heading 
Distribution comes largely from the literature. For Thailand I 
have consulted Smith (1945) and for Indonesia Weber and de 
Beaufort's Fishes of the Indo-Auslralian Archipelago. It should 
be noted that most of the distributional information in Weber 
and de Beaufort cannot be verified by re-examination of spec- 
imens because it is based on Bleeker's locality records for which 
documented specimens are unavailable. I have tried to utilize 
this information judiciously, but workers should be alert to its 
shortcomings. For many species it seems to me that many or 
most locality records may be based upon misidentifications, and 
therefore I have not attempted a summary statement of their 
distribution. The maps of species distribution, on the other 
hand, are based on information regarded as reliable. Scleropages 
formosus could hardly be confused with any other species; due 
to its presumed status as an endangered species a special effort 
has been made to obtain all available information on its dis- 
tribution. The distribution maps of Cyclochcilichthys and Do- 
ryichthys are based mainly on information provided by my col- 
leagues Sucbsin Sontirat and C. E. Dawson, who have revised 
the species involved. The rest of the distribution maps are based 
almost entirely on material examined by me. 

The work of Volz (1907) contains numerous potentially valu- 
able locality records of freshwater fishes in Sumatra. Some of 
his material (e.g., the holotype of Bagarius lica) is deposited in 
the Bern Natural Histor\' Museum and perhaps elsewhere in 
Europe, but many — perhaps most — of his locality records seem 
to be based on sight identifications in the field. Until the material 
basis for Volz's localities can be examined it seems best to ignore 
them. 

Meristic and Other Morphological Data. — Observations 
have been made on many genera or species-groups extending 
into adjacent areas of southeast Asia, Burma, and India in efforts 
to elucidate species problems. For several of the larger and more 
complex groups extensive meristic data, including vertebral 
counts obtained from radiographs, are presented. This should 
facilitate future systematic study of such difficult groups as Ras- 
hora. Mysius, Kryptoplerus. Mastacembclus. and others. Very 
little of such data has been recorded previously. I have not found 
gill raker or vertebral counts previously recorded for any species 
ofRasbora or for most of the Sundaland catfishes. Some authors 
(e.g.. Sufi 1956, for Mastacembelidae) have recorded valuable 
meristic data on Asian freshwater fishes without indicating the 
source of the specimens from which it was taken, thus greatly 
diminishing the utility of their data. In the present account the 
source of the specimens from which meristic and other data has 
been taken is directly stated or can be readily deduced. Data 
from type specimens are invariably presented separately, rather 
than lumped with data from non-types. 

Remarks on Counts and Measurements. — Counts and 
measurements used herein generally are those of Hubbs and 
Lagler (1947), and conform to their definitions. The standard 
measure of size, except in Dasyatidae, is standard length, and 



proportional measurements (unless otherwise stated) are ex- 
pressed as times in standard length. The standard measure of 
size in Dasyatidae is disc width (at widest point). For specimens 
under 100 mm, standard length is given to the nearest 0.1 mm; 
in specimens of 100 mm or more, to the nearest mm. Relatively 
few new measurements have been used. One of these is humeral 
spine (=postcleithral process of Lundberg) length in catfishes, 
measured from base of pectoral fin spine to tip of humeral 
process. Body width, especially useful in catfishes, is measured 
at widest point across the pectoral girdle. Only a few new or 
unfamiliar counts are used, most of them self-explanatory. 
"Transdorsal scale rows," used in Cyprinidae, refers to the max- 
imum number of scale rows on the anterior part of the body 
lying dorsal to the lateral line scale row of either side. I find 
vertebral counts helpful in various groups, and it is often im- 
portant to indicate the number of abdominal and caudal ver- 
tebrae as well as the total number of vertebrate. I define the 
division between abdominal and caudal vertebrae as the rela- 
tionship of the hemal spines to the anal fin pterygiophores. The 
first caudal vertebra is that with its hemal spine posterior to the 
anteriormost anal fin pterygiophores. All vertebrae anterior to 
this (including those of the Weberian complex) are abdominal. 
This method obviously will not work for fish without an anal 
fin, but I have found it works well with all fish groups herein 
for which vertebral data were taken. In some groups, especially 
catfishes and mastaccmbclids, the vertebral column is often 
anomalous (with complex or multiple central fusions). Speci- 
mens with abnormal vertebral columns should be omitted from 
the data base. This is particularly a problem in Mastacembel- 
idae. in which fin-ray as well as vertebral counts may be distorted 
by abnormality which may not be externally evident. Thus, all 
fin-ray and vertebral counts presented herein for Mastacem- 
belidae are based on specimens with normal vertebral columns. 

Dasyatidae 

Dasyatidae (whiptailed stingrays) comprises about six genera 
and 40 species. The great majority of the species are restricted 
to marine or marine and estuarine habitats, but the Indo-west 
Pacific Hyp(^lophiis scphcn {¥orsk&\ 1775) frequently enters fresh 
water, sometimes ascending nvers for hundreds of kilometers, 
and two genera — Dasyatis and Himantum—are represented by 
endemic freshwater species. A detailed account of southeast 
Asian freshwater Dasyatidae is given by Compagno and Roberts 
(1982). A single freshwater species of Hinianliira, discovered 
during the Kapuas survey of 1976, is known from western Bor- 
neo. 

Dasyatidae appears to be the only family of cartilaginous 
fishes or Elasmobranchii present in the fresh waters of western 
Borneo. No sharks or sawfishes have been reported from the 
Kapuas, and fishermen I interviewed in 1976 unanimously in- 
dicated their absence. 

Himantura Miiller and Henle, 1837 

llimanlura Miiller and Hcnic, 18.^7:4110 (type species /Jk/ij iiarnak ForskSI, 1775. 
b\ subsequent designation of Garman 1^I.'*;.''75). 

Himantura differs from all other dasyatids in having a long 
slender tail without dorsal or ventral tail folds. 



ROBERTS-WESTERN BORNEO FISHES 



23 




Figure 4. Himanlura sigmfer. Kapuas 1976-19. Left. 312 mm male (MNHN 1981-1342, paratypc): nght. 294 mm female (MZB 3004, holotype). 



Himantura slgnifer Compagno and Roberts, 1982 

(Figure 4) 

Himanlura sigmfer Compagno and Roberts, 1982:333 (l>pe locality mouth of 
Sungai Ketungau near Kapuas mainstream). 

Material Examined.— Thailand: Chao Phr\a, Nakornsawan prov., 263 mm 
disc width adult male (KUMF 2862). Western Borneo: Kapuas 1976-54, 294 mm 
disc width (MZB 30304. holotype): Kapuas 1976-19. 6: 278-382 mm disc width 
(BMNH 1981.1 1.16.1. CAS 48777, MNHN 1981-1342, MZB 3005, RMNH 28800, 
USNM 229492, paratypes), 

A species of Himantura with a relatively oval disc and dis- 
tinctive brown and white coloration: overall color of dorsal 
surfaces of disc, pelvic fins, and tail base light brown or tan; 
dorsal surface of disc and pelvic fins with a narrow milk-white 
peripheral margin: sides of tail base and tail posterior to sting 
milk-white; more or less well defined milk-white spots imme- 
diately in front of each eye and behind each spiracle; ventral 
surfaces uniformly pale or white. Tooth rows in upper/lower 
jaws 35-45/38-46. Total pectoral fin radials 109-1 16, 

Distribution.— Western Borneo (Kapuas); tentatively iden- 
tified from Thailand (Chao Phrya), Malay Peninsula (Perak R.), 
and Sumatra (Indragiri) (see Compagno and Roberts 1 982:336). 
Apparently restricted to fresh water. 

Osteoglossidae 

The bony tongue or osteoglossomorph family Osteoglossidae 
comprises four genera, Osteoglossum and Arapaima in South 



America, Heterotis in Africa, and Sclewpages in southeast Asia 
and the Australian region. 

Scleropages Gunther, 1864 

Scleropages GUnther. 1 864: 1 96 (type species Scleropages leichariiii Gunther, 1 864, 

b\ monotypy). 
Delsmama Fowler, 1934:243 (type species Oneoglossum jitrmoium Miiller and 

Schlegel, 1844, by original designation and monotypy). 

Scleropages includes two species in the Australian region (Lake 
1971) and one species in southeast Asia (including western Bor- 
neo). 

Scleropages formosus (Miiller and Schlegel, 1844) 

(Figure 5) 

Osleoglossiim formosum Miiller and Schlegel. 1844:1 (type locality Barilo River). 
Scleropages formosus Weber and de Beaufort, 1913:13. 

M,^TERiAL Examined. -Western Borneo: Kapuas 1976-19, 300 mm (MZB 3010); 
Kapuas 1976-43, 2: 148-279 mm (CAS 49182, USNM 218603). 

This southeast Asian species of Scleropages is readily distin- 
guished from the two species of the Australian region by its 
much larger scales, of which there are only 21-24 in a lateral 
series (instead of about 35). 

Distribution (Fig. 6). — Thailand (southeast). Kampuchea 
(southwest). Southern Vietnam. Malay Peninsula. Sumatra. 
Borneo (Sarawak, Sambas, Kapuas, Barito, Mahakam). Banka. 



24 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 5. Scleropages formosus. Kapuas 1976-43, 148 mm (CAS 49182). 



Notopteridae 

The osteoglossomorph family Notopteridae or featherfins 
comprises three genera. Papyrocrauiis and Xcnoinysins in trop- 
ical Africa. Notoptents in India and southeast Asia. 



Notopterus Laccpede, 1 800 

Noloplenis LacepMe, 1800:190 (lypc species (iyninunn iinuiptciiis Pallas, 1769, 

by absolute taulonymy). 
Chilali) Fowler, 1934ii;244 (type species Mynus chilala Haiiiillon-liuchanan, 1 822, 

b\ original designation and absolute tautonym\) 



properly identified and their distributions worked out. There 
seems to be only one species in western Borneo. 

Clupeidae 

Apparently only two species of herrings occur in the fresh 
waters of western Borneo, Cliipeichihys blcekcrl and Clupcoides 
hypselosonia. Both belong to Pellonulinae, a predominantly 
freshwater subfamily with endemic species or genera in main- 
land southeast Asia, southern India and Sri Lanka, Madagascar, 
southeast Africa, and tropical west Africa (including the Zaire 
or Congo basin, where the subfamily is most diverse). 



Notopterus borneensis Bleeker, 1851 

Noliiplcriis horiwcnsis Bleeker. 1851c-:4.^7 (t>pe localit\ Sambas). 

Material Examined. — Western Borneo: Rapuas 1976-19. 28."! mm (MZB 3006): 
Kapuas 1976-20, 2: 247-248 mm (CAS 49181, MZB 3007): Kapuas 1976-35. 
329 mm (MZB 3008); Kapuas 1976-45. 172 mm (MZB 3009). 

The southeast Asian species of Notopterus are poorly known; 
the entire genus will have to be revised before they can be 




I3,«¥ -10° 



Scleropages formosus 



=5. U -10" 



Figure 6. Scleropages hrmosus Geographical distribution. 



Clupeichthys Bleeker, 1855 

Cliipculilhys Bleeker, 1 855/i:274 (type species Chipcichlhys goitiognalinis Bleeker, 
1855, by monotypy). 

Distinguished from all other clupeid genera in southeast Asia 
by having a divided anal fin. Two species known, both appar- 
ently restricted to fresh water. 

Clupeichthys bleekeri Hardenberg, 1936 

(Figure 7) 

C /»/)<'( j/(fes-/),«'Ha'c)/)/('nKVaillant. 1893:100 (nee Bleeker. 1852: Kapuas. Knapei). 

Clupeichthys goiuognathus VdiiWanX. 1902:31 (nee Bleeker. 1855; Kapuas). 

Corica iClupeichlhys) gitiiiogihithiis Weber and de Beaufort. 1913:55 (pro parte; 
fig. 21, Kapuas 4), 

Clupeichthys hiecken Hardenberg, 1936:229 (t\pc locality middle course of Ka- 
puas). 

Material Examined. — Western Borneo: Kapuas 1976-9, 28: 38.4-61.0 mm 
(CAS 49183. MZB 3011): Kapuas 1976-14. 41: 22.0-60.6 mm (AMNH 48878. 
BMNH 1979.3.21.145-152. FMNH 94193. MZB 3012): Kapuas 1976-15. 28: 
22.2-40.0 mm (CAS 49508, IRSNB 19726, MNHN 1982-650. MZB 3013); Ka- 
puas 1976-22. 13: 52.2-60.8 mm (MZB .3014, ROM 38625); Kapuas 1976-33.9: 
I 1.7-40. 1 mm (MZB 301 5. UMMZ 209884): Kapuas 1976-35. 34: 34.3-71.7 mm 
(CAS 49184, MZB 3016, HSNM 230152); Kapuas basin. Mendalan River, 26.3 
mmtZMA 100,949); Kapuas basin. PoetoesSibau, 6: 44,2-69.0 mm (ZMA 100.948. 
100.952), 

Clupeichthys bicckcri differs from C. goniogiiathus, its only 
congener, in lacking a dusky longitudinal stripe on the body and 
having a somewhat longer pectoral axillary scale (T. Wongra- 
tana. pers. comm.). 

Distribution. — Borneo (Kapuas. Pamangkat in southern 
Borneo; T. Wongratana. pers. comm.). 



ROBERTS-WESTERN BORNEO FISHES 



25 




Figure 7. Clupeichlhys hleeken Kapuas 1976-14, 58.0 mm (BMNH 1979.3.21.145, drawn by T. Wongralana). 



Clupeoides Bleeker, 1851 

Clupeoides Bleeker, 1 85 U '.274 (type species Clupeoides horneemis Bleeker, 1851, 
by monotypy). 



Clupeoides hypselosoma Bleeker, 1 866 

(Figure 8) 

Clupeoides hypselosoma Bleeker. 1866/):293 (type locality "Bandjcrmassing, in 
fluviis"). 

Clupea (Clupeoides) poiaiuophdiis Bleeker, 1866-72:101 (proposed as substitute 
name for Clupeoides hypelosoma. regarded as preoccupied by Harengula (or 
Clupea) hypselosoma Bleeker, 1855). Sec Whitehead et al. (1966:76). 

Material Examined.— Western Borneo: Rapuas 1976-42,24.2 mm(MZB3017), 
Southern Borneo: Bandjermassing, 4 1 . 5 mm (BMNH 1867.1 1.28.35, 57 mm total 
length, holotype). 



This species previously was known only from the holotype; 
identification of the single specimen obtamed in 1976 was made 
by T. Wongratana. For a redescription of the holotype see 
Whitehead et al. (1966:74). 

Distribution. — Borneo (Kapuas, Barito). 

Engraulidae 

The Engraulidae (comprising the anchovies) is represented by 
two freshwater species in western Borneo: Lycothrissa cwcodihis 
and Selipinna melanochir. Additional species, such as Coilia 
coomansi Hardenberg, 1934 and Selipinna breviceps (Cantor 
1850) evidently occur in the mouth and lower reaches of the 
Kapuas River but are not actually known from freshwater hab- 
itats and are therefore excluded from the present account. 




Figure 8. Clupeoides hypselosoma. Bandjermassing, 42.0 mm (BMNH 1867.1 1.28.35, holotype). 



26 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 4. Lycolhnssa croivtiilm Kapuas I'iTO-ai. 147 mm (MZB 3024). 



Lycothrissa Gunlhcr, 1868 

l.nolhns'^Mi Gunther. 1S68;385. 34>} {l\pc species Hni;rLiiiln cmintlihii Bleeker. 
1S3 1 . h\ monot\p>), 

Lyc(>!l!nssci is distinguished from all other southeast Asian 
cngraulids by its enlarged caniniform jaw teeth. A single species, 
restricted to fresh water. 

Lycothrissa crocodilus (Bleeker, 1851) 

(Figure <)) 

linf;riiulisiri>oHlilii.s Bleeker. 185 Id; 15 (type localitv "Banjcrmassing, in fluviis'")- 
L\inihii\\<i irmoililiis Bleeker, 18(i6-72: 1 25. 

Material Examined. — Wcslern Borneo; Kapuas 147b-14, 4; 93.6-148 mm 
(CAS 49185. MZB 3019); Kapuas 1976-15. 141 mm (MZB .3020); Kapuas 1976- 
19. 144 mm (MZB 3021 ); Kapuas 1976-33. 4; 1 1 7-1 72 mm (AMNH 4887Q. MZB 
3022. I'SNM 230153); Kapuas 1976-35. 1 19 mm (MZB 3023); Kapuas 1976-41. 
147 mm (MZB 3024). Kapuas 1976-44, 2: 117-170 mm (MZB 3025, LISNM 
209911). 



Distribution. — Thailand. Sumatra (Banjuasin, Palembang, 
Djambi). Borneo (Kapuas. Sinkawang. Banto). 

Setipinna Swainson, 1839 

Siiipiiuki Swainson, 1839;I86, 292 (l\pe species Sclipiiiiiu mcKdliira Swainson. 

\i}9=Clupcii i>hasu Haniillon-Buchanan, 1822, by suhseciuent designation of 

Swain I8S2;280; see Whitehead et al. 1966:128) 
Tclara Gunther. 1868i;;400 (type species Cliipca liiara Hamilton-Buchanan, 

\S22=Cliipca pha.sci Hamilton-Buchanan. 1822. by absolute tautonymy). 
Hett't'iHhnssa Giinlher, 1868d;385, 401 (type species Iiiii^iaiihs hivvuvps Cantor. 

1850. by monotypy). 

Setipinna melanochir (Bleeker. 1849) 

(Figure 10) 

Fiii;ruiilis inclamxhir Bleeker. 1849; 13 (type locality "in Freto madurae prope 

Kammal et Surabaya"). 
Ciiilia )>u'/uH<)W»r Bleeker. 1855;418. 
Selipmna melanochir Bleeker. 1866-72; 136. 




Figure 10. Sclipiniia incliuwilur Gull of Thailand, 153 mm (BMNH 1966.3.8.72, drawn by T. Wongratana). 



ROBERTS-WESTERN BORNEO FISHES 



27 




Figure 1 1. Sundasalanx microps. Kapuas 1976-33, 17.0 mm (CAS 44220, paralype). 



Material Examined. — Western Borneo: Kapuas 1976-33, 159 mm (MZB 3026); 
Kapuas 1976-52, 101 mm (MZB 3027). 

Distribution. — Widely distributed in brackish and fresh water 
in Thailand, Malay Peninsula, Sumatra, and Java. 

Sundasalangidae 

Although first collected in the Kapuas River by Maunce Chaper 
in 1890. and more recently in the Mekong River along the Thai- 
Laos border and in the Malay Peninsula, these minute trans- 
parent fishes were not recognized as taxonomically distinct until 
my study of Kapuas specimens demonstrated that they are sex- 
ually mature and constitute a separate family related to the east 
Asian noodlefishes or Salangidae (Roberts 1981, 1984). 

With both sexes ripe at 14.6 mm. the largest known specimen 
only 23.4 mm, Sundasalangidae are by far the smallest known 
salmoniforms. They differ from adult Salangidae in having a 
median pectoral girdle, pectoral fins without segmented rays, 
hypobranchials 1-3 evidently fused to basibranchials 1-3 in- 
stead of separate, pelvic fins with 5 instead of 7 rays, adipose 
fin absent, males without anal scales or enlarged anal fin, and 
total vertebrae only 37-43 (versus 48-77). A detailed account 
of the skeletal anatomy and systematics of Sundasalangidae and 
Salangidae has been published (Roberts 1984). 

The family comprises a single genus with two or more species 
from Thailand, the Malay Peninsula, and Borneo. 

Sundasalanx Roberts, 1981 

Sididasalaiix Roberts. 1 98 1 :297 (type specisi, Sundciialuii\ pi citrox Roberts. 1981, 
by original descnplion). 

Sundasalanx microps Roberts. 1981 

(Figure 1 1) 

Siiiuiasalaiix microps Roberts. 1 98 1 :30n (type localilN Rapuas River mainstream 
al Kampong Nibung, about 7 km NE of Sclimbau). 

Material Examined. — Western Borneo: Kapuas 1976-33, 35: 14.6-19.9 mm 
(CAS 44220. MZB 3000-3001, USNM 227793, holotype and paralypes). 

Distribution. — .S;(«(i't;5fl/a/;.v microps is reported only from 
the Kapuas. but further study may show that it is more, widely 
distributed. Sundasalanx (including undetermined species) also 
occurs in the Malay Peninsula and Mekong basin but is not yet 
known from Sumatra. 

Sundasalanx sp. undet. 

'!Behne caiidiinaculala Vaillanl, 1893:1 10-112. pi. 2. fig. 4 (Kapuas). 
Sundasalanx sp. undet. Roberts. 1981:300. 



Material Examined —Western Borneo: Kapuas. 
1891-596). 



17: 18.8-23.4 mm (MNHN 



Although these specimens were presumably collected near the 
type locality of 5. microps (see Roberts 1981:302), they have 
much larger eyes, fewer maxillary teeth, and perhaps slightly 
more vertebrae (about 42-45 instead of 41-43). 

Cyprinidae 

Cyprinidae, comprising the carps or minnows, is the domi- 
nant family of freshwater fishes in Asia and includes more genera 
and species than any other family of freshwater fishes. About 
one-third of all freshwater fishes in western Borneo belong to 
this family. Many genera and most species found in western 
Borneo are endemic to southeast Asia, and many species are 
known only from Sundaland or the Kapuas basin. All cyprinids 
have toothless oral jaws but well-toothed pharyngeal jaws with 
1-3 rows of large teeth. Despite lack of jaw teeth, diversity of 
feeding habits is very great. Among the new cyprinids discovered 
in the Kapuas in 1976 are Thryssocypris smaragdimis, an in- 
sectivorous new genus and species that looks remarkably like 
an anchovy, and Pectenocypris balaena. a small phytoplank- 
tonophage with over 200 gill rakers on its first gill arch. 

Albulichthys Bleeker, 1859 

Alhuhchlhvs Bleeker. 1 859i(: 1 53 (type speeics Sv\i<^nus ulhidoidcs Bleeker, 1 855, 
b\ monot\py) 

Albulichthys, with a small mouth and numerous tuberculate 
gill rakers forming a complex lattice-work, seems closely related 
to Amblyrhynchichthys. Kalimantania. and Punlioplnes (see ac- 
counts of these genera below). It differs most obviously from 
these genera in its rounded and relatively elongate snout, terete 
body form, and relatively small dorsal and anal fins. Body sub- 
cylindrical, snout obtusely rounded. Hyaline eyelid very well 
developed. Small rostral and maxillary barbels present. Lateral 
line complete. Scales in lateral series 36-38, predorsal scales 14, 
transdorsal scales 1 1. circumpeduncular scales 16. Scales with 
6-1 1 parallel radii. Dorsal fin with last simple ray very slender, 
almost flexible, very weakly serrated; branched dorsal-fin rays 
8; branched anal-fin rays 5. Pectoral-fin rays 17-19, pelvic 10, 
An elongate pelvic axillary scale. Pharyngeal teeth compressed 
with longitudinally grooved masticatory surface, 4,3,2/2,3,4 
(Weber and de Beaufort 1916:107; Chevey 1932, fig. 2). Total 
gill rakers on first arch 22 + 27=49. Vertebrae 22+ 12=34 (three 
Kapuas specimens). 

A single species restricted to southeast Asia. 



28 



CALIFORNIA ACADEMY OF SCIENCES 




FioLiRt 12. Albuluhthys albulutdcs- Kapuas 1976-52, 98.0 mm (CAS 49186). 



Albulichthys albuloides (Bleeker, 1855) 

(Figure 12) 

Svsloiiuis albuloides Bleeker. 1855(/:425 (lypc localits "Poniianak, in flumme 

Kapuas"). 
Illwluiithvi albidoides Bleeker, I860i;:3U6. 
Albuluhlhys Krempti Pellegnn and Chevey, 1927:304 (type localin Pnom-Pcnh), 

Che\cy. 1932:26. pi, 6. 

Material Examined, — Western Borneo: Kapuas 1976-52, 6: 71,2-98,0 mm 
(CAS 49186, MZB 3028, USNM 230154), 

Pellegnn and Chevey ( 1 927) and Chevey ( 1 932) distinguished 
their Mekong basin A. krempfi from A. albidoides on the basis 
of its having a larger eye, more truncate snout, dorsal fin less 
elevated, last simple dorsal-fin ray more feebly ossified, caudal 
peduncle more slender, and caudal-fin lobes more rounded. It 
would appear that these supposed differences were deduced from 
the only published figure of ,-J, albuloides (Bleeker 1863, repro- 
duced by Weber and de Beaufort 1916, fig, 47) rather than on 
actual specimens of ,-J, albuloides from Indonesia, This figure 
portrays the snout too sharp and dorsal fin too high. The other 



differences also seem to be non-existent or insignificant. Kapuas 
specimens of .-I, albuloides agree very well with the descriptions 
and both figures of ,-1, krempfi and I conclude they are the same 
species. 

DisTRiBiiTioN, — Laos, Vietnam, Kampuchea (Mekong basin, 
Tonle Sap), Thailand (Mekong, Chao Phrya; Smith 1945). Su- 
matra (Palembang), Borneo (Kapuas, Kahajan, Barito?). 

Amblyrhynchichthys Bleeker, 1859 

IniblvrlivihhiclilhV' Bleeker, 1859i;:153 (type species Burbus Inincalus Bleeker, 
1851, h\ monotypy). 

This monotypic genus and species is immediately recognized 
by its blunt, concave snout; antenorly situated nostrils; and 
small, distinctively shaped jaws. Hyaline eyelid well developed. 
Gill membranes broadly joined to flat, broad isthmus. Last 
simple dorsal-fin ray snout and serrate, branched dorsal-fin rays 
9, Branched anal-fin rays 5, Lateral line complete. Scales with 
strongly convergent radii lateral scale series 36; predorsal scales 




Figure 13, Aiublvihviuhuhlhv\ liiincalin Kapuas, Bunut (alter Weber and dc Beaulorl 1916, fig. 46). 



ROBERTS-WESTERN BORNEO FISHES 



29 




Figure 14. Balanliochedos miianoplcnis. Kapuas 1976-49, 1 13 mm (MZB 3037). 



13; transdorsal scales 1 1; circumpeduncular scales 16. Gill rak- 
ers on first arch 10-13 + 27-29=39-40. Vertebrae 21-22+12- 
13=34 (in three Kapuas specimens). 

The only other nominal species referred to Amhlyrhynchich- 
ifiys, A. alius Vaillant, 1 893, is a synonym of Pun! ioplites lawak. 
type species of the monotypic genus Kaliiiuuitania Banarescu, 
1980. 

Amblyrhynchichthys truncatus (Bleeker, 1851) 

(Figure 131 

BLirhiis InincatusBlteker. 1 85 la: 13 (type locality Banjcrmassing. m lluviis). 
Amblyrhynchichthys truncatus Bleeker, 1859ii:153. 

Material Examined.— Western Borneo: Kapuas 1976-19, 169 mm (MZB 3029); 
Kapuas 1976-33, 3: 36.3-107 mm (MZB 3030. I.ISNM 230155); Kapuas 1976- 
44, 6: 93.8-147 mm (CAS 49187, MZB .3031, HMMZ 209912); Kapuas 1976- 
53, 91.0 mm (MZB 3032); Kapuas 1976-54. 206 mm (MZB 3033). 

Distribution. — Thailand (Menam Pasak, Meklong, Chao 
Phrya, Mekong, Tapi). Sumatra (Palembang, Djambi). Borneo 
(Kapuas. Barito). 

Balantiocheilos Bleeker, 1859 

Balanliocheitos Bleeker, 1859i;:149 (type species Burlnis inetanopierus Bleeker, 
1851, by monotypy). 

This monotypic genus is readily identified by its lip mor- 
phology and fin coloration. 

Rostral and maxillary barbels present but small; anterior mar- 
gin of rostral barbel joined by a frenum to rostal cap. Rostral 
cap fleshy, deeply incised. Upper and lower lips thick and fleshy, 
their external surface with deep longitudinal or transverse 
grooves; lower lip posteriorly prolonged to form a deeply incised 
lobe with an < -shaped posterior border. Isthmus at point of 
posterior origin of incision of lower lip with a complementary 
< -shaped vertical incision; apex of isthmus posteriorly incised 
to form a small trapezoidal dermal wedge. This is conceivably 
homologous with a similarly shaped, somewhat larger dermal 
wedge bearing hypertrophied cutaneous papillae on the isthmus 
oi Cyclocheilichlhys. Last unbranched dorsal-fin ray moderately 



enlarged and strongly serrate. Distal margins of dorsal, anal, 
caudal, and pelvic fins with wide falcate marginal black bands; 
basal portion of caudal and sometimes other fins yellow. Gill 
rakers 7+ 10=17 (very short). Vertebrae 21 + 12=33 (three Ka- 
puas specimens). 

Balantiocheilos melanopterus (Bleeker, 1851) 

(Figure 14) 

Barbus melanopterus Bleeker, 1851a:l 1 (type locality Bandjermassing). 
Balantiocheilos melanopterus Bleeker, 1859ii:149. 

Material Examined.— Western Borneo: Kapuas 1976- 15, 8 1.8 mm (MZB 3034); 
Kapuas 1976-20, 4: 83.7-133 mm (CAS 49188. MZB .3035); Kapuas 1976-44, 
I 17 mm (MZB 3036); Kapuas 1976-49, 2; 87.9-1 13 mm (MZB 3037, USNM 
230156). 

Distribution.— Thailand (Mekong, Chao Phrya). Malay 
Peninsula (Perak). Sumatra (Palembang, Djambi. Kwantan R.). 
Borneo (Kapuas, Kahajan, Barito, Mahakam). 

Barbichthys Bleeker, 1859 

Barbichthys Bleeker, 1 859a: 147 (t> pe species Barbus lacvis Valenciennes in Cuvier 
and Valenciennes. 1842, by monotypy). 

This southeast Asian genus has a distinctively shaped head 
and mouth, numerous morphologically specialized gill rakers, 
and perhaps the most highly modified oral and pharyngeal ep- 
ithelia of any cyprinoid. Its relationships to other cyprinids are 
unknown. 

Mouth broad, subterminal. Rostral cap expanded posteriorly 
dorsal and ventral to rostral barbel, reaching comer of mouth, 
deeply incised, its posterior margin entire. Rostral barbel mod- 
erately elongate, lying (when adpressed) in shallow longitudinal 
groove on external surface of rostral cap. Maxillary barbel pres- 
ent. Upper lip similar to rostral cap, its margin entire and broad- 
ly continuous at comers of mouth with lower lip. Lower lip thin, 
moderately long, deeply incised posteriorly. Upper and espe- 
cially lower lips with a few irregularly placed flat tubercles of 
variable size near comer of mouth. A ventrally directed median 
papilla or barbel-like flap arising from lower lip at symphysis 



30 



CALIFORNIA ACADEM'i- OF SCIENCES 




FuRiRE I 5, Hurilnis bormvnsis Kapuas 1976-27, 38,0 mm (MZB 3U43. hololype). 



of lower jaw. Upper and lower homy jaw sheaths absent or 
rudimentary. Oral roof covered with large wrinkled folds or 
pleats, the largest longitudinally median, the others arranged at 
varying angles. Pharyngeal roof strongly arched, the apex of the 
arch forming a steep, narrow, median longitudinal groove and 
Its eaves a pair of almost vertically oriented, thickened lateral 
pads or cushions opposed to the raker bearing portions of the 
gill arches. Each lateral pharyngeal pad with about eight elevated 
longtudinal rows of fingerlike papillae or fimbriae. In smaller 
specimens the fimbriae are simple and similar in size and shape 
to the uniformly distributed fimbriae on the median pharyngeal 
cushion oi' Dangila. In the largest specimens examined (to 193 
mm), the fimbriae are muhifid, each with 5-6 fingerlike pro- 
jections which may themselves bear minute projections. 

Total gill rakers on first arch 36-48, their number apparently 
subject to little or no increase within size range of specimens 
examined. Rakers moderately large and fleshy, with numerous 
small fimbriate projections, those of successive arches inter- 
posing to form a highly complex lattice-work somewhat similar 
to that ofthe.)//'i(//c-/??/n'j--/';/"?/(Y'/"''-f group of southeast Asian 
cyprinids. 

Lateral line complete. Scales with moderately numerous par- 
allel radii; lateral scales 36-38, transdorsal 1 1, predorsal 13-14. 
Last simple dorsal-fin ray non-serrate, branched dorsal-fin rays 
8. Branched anal-fin rays 5. Pharyngeal teeth 2.4,5/5,4,2. Body 
silvery. Middle and dorsal fin with diagonal black band, upper 
and lower caudal-fin lobes with longitudinal submarginal black 
bands. Vertebrae 23+ 1 1 =34 (two Kapuas specimens). A single 
species is recognized. 

Barbichthys laevis (Valenciennes, 1842) 

Harhiis lucvis Valenciennes in CuvLcr and Valeneiennes, 1842:192 (type locality 

Builcn/org and Sarayevi, Java). 
Rarhus hrachviwmus Bleeker, 1850:18 (type loealit\ Surabaya, in tlumine K.ali- 

masl. .See Bleeker I860a:209. 
fiiiihus gobiiiulci Bleeker. Militi^l (type localit\ Palenibani;. in fluviis). See 

Bleeker ( 1 86O0: 209). 
Harhuhlhys kicvis Bleeker. 1859(;:147. 1860i;:209, 
liaihuluhvi laevn var, suiiuilrciiuis Volz. 1904:478. 
Harbuhthys «/f/(/i« Sauvage. 1878:9 (type locality Indo-C hinc). 
Barhichlhys lacvis nilidus Banarescu, 1980:98. 

Material Examined —Malay Peninsula: Perak. Chenderoh Dam. 5: 138-193 
mm (CAS-SH 34659), Western Borneo: Kapuas 1976-19. 5: 99.0-137 mm (CAS 
49 189, MZB 3038, LISNM 230157); Kapuas 1976-20, 85,2 mm (MZB 3039); 
Kapuas 1976-33, 106 mm (MZB 3040); Kapuas 1976-37, 1 10 mm (MZB 31.141). 
Kapuas 1976-44. 1 10 mm (MZB .3042). 



Distribution. — Thailand (Mekong, Chao Phrya, Patani). 
Malay Peninsula. Sumatra (Lampong, Pangahuang, Lahat, Pa- 
lembang, Kwantan R., Rampar R., Indragiri, Djambi). Borneo 
(Kapuas, Barito, Mahakam). Java (Batavia, Tangerong, Ran- 
kasbetong, Lebak, Buitenzorg, Parongkalong. Brantas R., Sura- 
baya, Gempal). 

Barilius Hamilton-Buchanan, 1822 

Ranlius Hamilton-Buchanan. 1822.266. 384 (type species Cypnnus bania Ham- 
ilton-Buchanan, 1822, by subsequent designation of Bleeker, 1863/':203). 

Opsaiiiis McClelland, 1 839:295, 4 1 3 (type species Cypniiin iBanhus) banIa Ham- 
ilton-Buchanan, 1822, by subsequent designation of Bleeker, 1863:203), 

Pachvilomus Heckel. 1843:1038 (type species Cypnnus {Bardius) shacra Ham- 
ilton-Buchanan, 1822, by subsequent designation of Bleeker, 1863ft:203; preoc- 
cupied by Pachvslonius Latrcillc, 1809, Diptera). 

Op'.andmm Peters. 1854:783 (type species Laiciiciis ranibcrcnsis Peters. 1854, 
b\ monotypy), 

Shutra Bleeker, 1860c;: 431 (t\pe species PLichvsloiniis schagra Heckel, 
l^4} = C'vp''inn\ {Buribin) shdini Hamilton-Buchanan. 1822. by absolute tau- 
tonymy 1 

Bcmiclisis Bleeker. I860u:431 (type species Cipnnns {Bunhus) bcndtiisis Ham- 
ilton-Buchanan, 1822, by absolute tautonymy and monotypy). 

Bi'la Gunther. 1868:293 (type species Cypnnus (Bardms) bold Hamilton-Bu- 
chanan, 1 822, by absolute tautonymy; preoccupied by Sola Hamilton-Buchan- 
an, 1822, Sciacmdae), 

Rauvnas Jordan. 19|9;.«4 (replacement name for Bola GUnther. 1868). For 
current status see Howes (1980. 1983). 

Sagiltabanbus Fowler, 1936:293 (type species Bardms suliniibums Nichols and 

Gnscom, 1917, by original designation and monotypy), 
I/A i</(;/»() Smith, 1945: 100 (type species flt;H;('(.-l//i>(/«"/t');)("i"c'H/H.s Smith. 1945, 
by original designation and monotypy). 

Chedms Sviaimon. 1839:285 (type species CVm/rHj,i;imv(Swainson. l839=ri'pr/;;Hi 
{Bardms) ihcdra Hamilton-Buchanan. \f,21=Crpnniis {Bardms) bcndcbsis 
Hamilton-Buchanan. 1822, by monotypy) 

The genus Banlius has not been reported previously from 
Indonesia. A single specimen of an undescribed species was 
obtained during the Kapuas survey of 1976. 

Barilius borneensis new species 

(Figure 15) 

HoLOTYPE. — MZB 3043. 38.0 mm. rocky channel in mainstream of Sungai 
Pinoh. 37 km S of Nangapinoh (Kapuas 1976-27), 

A Barilius with short rostral barbel but no maxillary barbel; 
gill rakers very short, 1 + 1+6=8; lateral line scales about 39, 
predorsal about 15, transdorsal about 15, circumpeduncular 
about 1 5 (numerous scales missing); dorsal-fin rays ii9, anal-fin 
rays iii7, pectoral-fin rays 14, pelvic-fin rays 8; vertebrae 
19+19=38. 



ROBERTS-WESTERN BORNEO FISHES 



31 



Horizontal diameter of eye 1 1.5, head length 3.4, length ros- 
tral barbel one-third of eye diameter, body depth 4.1, caudal 
peduncle depth 10.7. Lateral line sloping gently downwards be- 
hind head, separated from pelvic-fin origin by about I'h and 
from anal fin by 2 scale rows. 

Tubercles apparently absent on body and fins, but well de- 
veloped on head, especially on ventral surface of mandible; 
ventrolateral surface of mandible with 3-4 irregular longitudinal 
rows of conical tubercles with crowns laterally directed, ventro- 
median surface of mandible with 2-3 irregular longitudinal rows 
of similar but somewhat smaller tubercles with crowns ventrally 
directed; scattered fine tubercles near tip of snout. 

Side of body from immediately behind head to end of caudal 
peduncle with about 10 midlateral marks, more distinct ante- 
riorly but all dusky and poorly defined; anterior 4-5 marks 
vertically elongate, posterior marks becoming progressively more 
horizontal and indistinct; posterior marks superimposed on a 
dusky longitudinal stripe (most distinct on caudal peduncle) 
extending about from a vertical below dorsal-fin origin to cau- 
dal-fin base. Dorsal-fin rays darkly pigmented near distal margin 
of fin, forming a slightly oblique dark longitudinal stripe across 
entire fin. Pectoral, pelvic, and anal fins unpigmented or with 
only a few inconspicuous melanophores along some fin rays. 
Caudal fin with numerous fine melanophores along fin rays; tip 
of lower caudal fin lobe slightly darkened. 

Bahliiis horneemis is apparently most similar to Banlius pon- 
ticuhis (Smith, 1945), and B. nanensis Smith, 1945. described 
from mountain streams in the Mekong basin of northern Thai- 
land. The two nominal Thai species B. ponticulus and B. na- 
nensis (probably conspecific with each other) have body-shape, 
fin positions, body and dorsal-fin coloration similar to those of 
B. bornecnsis but differ in having fewer scales in the lateral series 
(31-33 instead of about 39), 10 instead of only 7 branched anal- 
fin rays, and ma.xillary as well as rostral barbels present. 

Etymology. —The name borneensis refers to the geographical 
origin of this species, the only member of the genus known from 
Borneo. 

Chela Hamilton-Buchanan, 1822 

Chela Hamilton-Buchanan, 1822:258. 383 (proposed as subgenus oi Cypnmiy. 
type species Cypniuis cachius Hamilton-Buchanan, 1822, by subsequent des- 
ignation of Bleeker, 1 863?: 264). 

Lauhuca Bleeker. l859-60i"438 (type species Cyprinm laiihiica Hamilton-Bu- 
chanan. 1822, by absolute tautonymy), 

Cflr/?/»5 Giinther. 1868:339 (type species CvpnniLs cachius Hamilton-Buchanan. 
1822, by absolute tautonymy). 

Allochela Silas, 1958:64. 87 (type species Chela tasciala Silas. 1958, by onginal 
designation; proposed as subgenus of Chela). 

Neocheta Silas. 1958:64, 93 (type species Laubtua dadybiirion [emended from 
misspelling dadihiinori] Menon, 1952, by original designation and monotypy). 
Proposed as a subgenus of Chela 

Atalayochela Banarescu, 1968:59 {lype species Kusnra niaasst Weber and de Beau- 
fort. 1912. by monotypy). 

Chela, as revised by Silas (1958), comprises 6-7 species dis- 
tributed in the Indian subcontinent, mainland southeast Asia 
west of the Mekong basin, the Malay Peninsula, Sumatra, and 
Borneo (Silas 1958; Banarescu 1968). 

Chela maassi (Weber and de Beaufort, 1912) 

Chela megalolepis Vaillant, 1893:91 (in part: misidcntihed from Kapuas; see 
Banarescu 1968:61). 



Haploeheilm panehax Vaillant, 1902:15 (Pontianak; misidentification). 
Chela acinaces Duncker, 1904:183 (Muar River; see Banarescu 1968). 
Eustira maassi Weber and de Beaufort. 1912:531 (type locality Gunung Sahilan 

on Kampar River. Sumatra), 
Lauhuca (Eustira) maassi Weber and de Beaufort, 1916:49. 
Chela (Allochela) maassi Silas. 1958:91. 
Chela {Atalayochela) maassi Banarescu, 1968:59. 
''Chela {Malavochela) cf maassi Kottelat, 1982:420 (Mcntaya basin). 

Material Examined. — Malay Peninsula: Pahang, Tasek Bera, 35.8 mm (CAS 
47619). Western Borneo: Kapuas 1976-7. 33.5 mm (MZB 3044); Kapuas 1976- 
37, 3: 28.5-34.5 mm (MZB 3045. ROM 38610); Kapuas 1976-42, 12: 22.0-33.1 
mm (CAS 49190, MZB .3046, USNM 230158); Kapuas 1976-43, 35.3 mm (MZB 

3047). 

Diagnosis. — A comparatively slender-bodied Chela, depth 
about 3.3, with a dark median longitudinal stripe, lateral line 
scales 28-34, predorsal scales 20-2 1 , and branched anal-fin rays 
10-1 1, Pharyngeal teeth in two rows, 4-5,2-3/4-5,2-3 (Bana- 
rescu 1968), It is the only Chela species known from Borneo. 

Distribution. — Malay Peninsula (Muar, Pahang). Sumatra 
(Kampar). Borneo (Kapuas). 



Cosmochilus Sauvage, 1878 

Cosmochilus Sauvage, 1878:240 (type species CosDiochilui haniiandi Sauvage, 
1878, by monotypy). 

Diagnosis. — Large, deep-bodied and laterally compressed; 
dorsal fin large and falcate, with 4 simple and 8-9 branched 
rays; last simple dorsal-fin ray greatly enlarged, its posterior 
border more or less strongly serrated for its entire length; anal 
fin relatively small, with 3 simple and 5-6 branched rays; head 
relatively small, compressed; snout truncate, without enlarged 
tubercles or pores; mouth small and inferior, its opening trans- 
verse; rostral and maxillary barbels large and relatively elongate; 
lips moderately thick, entirely covered with large, contiguous 
papillae; horny jaw sheaths transverse, moderately thick but 
with relatively weak cutting edge; gill rakers fleshy, relatively 
unspecialized, 1 5-1 8 on first gill arch; pharyngeal teeth triserial, 
morphologically generalized for Cyprinidae, usually 1,3,5/5,3,1 
or 2,3,5/5,3,2; lateral line almost perfectly straight; each lateral 
line tubule with a short ventroposterior branch terminating in 
small pore on exposed portion of posterior shield; scales in 
lateral line series 35-48; circumpeduncular scales 16-18; scales 
oblong, with relatively huge posterior shields; radii of posterior 
shield strongly convergent; radii of anterior shield frequently 
conjoined or bifurcate; vertebrae 35-43, 

In addition to C. harmandi from the Chao Phrya and Mekong, 
the genus includes a species from the Lancang-jiang or Mekong 
in Yunnan Province, China, described by Chu and Roberts 
(1985), and one species from Borneo, 

Cosmochilus falcifer Regan, 1906 

Cosmochilus falciler Regan, 1906:66 (type locality Baram River). 

Material Examined. -Kapuas 1976-19, 316 mm (MZB 3048); Kapuas 1976- 
52. 143 mm (MZB 3409). 

Life color of C. falcifer has not been reported previously. The 
fresh 316 mm specimen caught in the Kapuas and photographed 
in the market at Sintang had overall color white or milk-white, 
especially ventrolaterally and ventrally; dorsolaterally and dor- 
sally distinctly brownish or violaceous brown; posterior margins 
of scales, especially on upper parts of body, with broad dark 



32 



CALIFORNIA ACADEMY OF SCIENCES 



margins: dorsal surface of head faintly yellowish and entire gill 
cover distinctly yellow; iris and ventral portion of head milk- 
white; entire dorsal fin rosy pink or faintly orangish except black 
at tip; pectoral fin white, pelvic white or pinkish; anal and caudal 
fins dusky, caudal very dark, its posterior margin almost black. 

Fishermen at Sintang, where large individuals of this species 
are frequently marketed, informed me that they are familiar 
only with full-grown or nearly full-grown adults. The only small 
individual (143 mm) 1 observed in 1976 was purchased at the 
market in Pontianak. 

Distribution. — Sarawak (Baram, Rejang). Western Borneo 
(Kapuas). 



Crossocheilus cobitis (Bleeker, 1853) 

(Figure 16) 

Lobncheilos cobilis Bleeker, 1853/:523 (t\pe locality Padang, Sumatra and Batavia, 

Java). 
Croisochcilos iCrossochcilichlhys) cabins Bleeker. 1860ii:125. 

Material Examined. — Western Borneo; Rapuas 1976-14, 34.0 mm (MZB 3050): 
Kapuas 1976-24. 3: 66.6-94.3 (CAS 49191. MZB 3051); Kapuas 1976-26, 5: 
70.3-99.8 mm (BMNH 1 982.3.29. 1 . MNHN 1982-65 1 , MZB 3052. RMNH 28822): 
Kapuas 1976-29. 2; 63.9-76.1 mm (MZB 3053, USNM 230159); Kapuas 1976- 
31. 2: 34,8-45 I mm (MZB .3054. USNM 230160). 

Distribution. — Sumatra (Padang), Borneo (Kapuas, Maha- 
kam). Java (Batavia, Surabaya), 



Crossocheilus Kuhl and van Hassell, 1823 

Crossocheilus Kuhl and van Hasselt iii van Hasselt, 1823:132 (type species Cros- 
socheilus oblongus Kuhl and von Hassell in van Hasselt, 1823, by monotypy). 
Crossocheilos Bleeker, 1853:525 (unwarranted spelling emendation). 
Owsin/i//i« Giinther. 1868:73 (unwarranted spelling emendation). 

It is debatable whether the brief original account of the new 
genus and species Crossocheilus oblongus in van Hasselt, 1823, 
qualifies as a valid description. It is tentatively accepted as valid 
because it mentions one taxonomic character ("mouth opening 
downwards in the form of a parallelogram") and because its 
recognition is in keeping with most current practice and seem- 
ingly will not be contrary to nomenclatural stability. The earliest 
unquestionably valid description of the genus is by Bleeker 
(1853); that of the generic type species is by Valenciennes in 
Cuvier and Valenciennes ( 1 842), 

Rostral cap very thick, its margin fimbriate. Rostral lobe pres- 
ent between rostral barbel and sublacrimal groove. Sublacrimal 
groove extending from origin of rostral lobe to behind rictus of 
jaws. Upper lip well developed, continued in a fold at comer 
of mouth, its entire margin uniserially papillate, .laws somewhat 
narrow, horny jaw sheaths present but not greatly thickened, 
without sharp cutting edge. Lower lip fleshy, fitting into a shal- 
low depression on homy sheath of lower jaw (as in Epalzeorhyn- 
chos). its anteroventral margin with several rows of lobate pa- 
pillae. Rostral and maxillary or rostral barbels only present. 
Males with dorsal surface of posterior half of first 8 or more 
pectoral-fin rays covered for their whole length with numerous 
close-set fine breeding tubercles. 



Key to Crossocheilus of western Borneo 

la Rostral and maxillary barbels present; mouth relatively 
narrow; rostral cap with 14-15 fimbriae; upper lip rela- 
tively thick, with large marginal papillae; papillae on lower 
lip relatively large C. cobilis 

lb Rostral barbels only present; mouth relatively wide; ros- 
tral cap with about 18-23 fimbriae; upper lip thin, with 
very small marginal papillae; papillae on lower lip rela- 
tively small, rounded 2 

2a Body with midlateral longitudinal stripe relatively pale, 
its margins poorly defined, ending in a small round spot 
at base of middle caudal-fin rays C. oblongiis 

2b Body with midlateral longitudinal line very dark (black), 
its margins sharply defined, extending two-thirds of length 
nearly to end of middle caudal-fin rays before ending 
abruptly C sp, undet. 



Crossocheilus oblongus Kuhl and van Hasselt, 1823 

(Figure 16) 

Crossocheilus Oblongus Kuhl and van Hassell in van Hasselt. 1 823; 1 32 (type 

locality Java). 
Labeo oblongus Valenciennes in Cu\ier and Valenciennes. 1842:357. 
Crossochilus oblongus Popta. 1906:120. 
Epalzeorhynchus suimensis ?,m\\\i. 1931:20. fig. 9 (Tapi Stream, near Nakon Sri- 

tamarat. Peninsular Thailand). See below. 
Epalzeorhynchus kalliurus Inger and Chin, 1962:99. fig, 46 (nee Epalzeorhynchus 

kalliunn Smith. 1945'') (Kinabatangan). 

Material Examined, — Peninsular Thailand; Pallani R. 3; 46.2-1 II mmlNIFI 
uncat). Malay Peninsula: Pcrak. Batang Padang, 5; 93.8-1 13 mm (CAS-SU 39348); 
Pcrak. Plus R., 97,8 mm (CAS-SU 68638); Pahang, Benas R., 3: 81,1-95.0 mm 
(CAS-SU 34667). Western Borneo; Kapuas 1 976-6. 66.3 mm (MZB 3055); Kapuas 
1976-14, 7; 40.5-48.6 mm (CAS 49192. MNHN 1982-652, MZB 3056, RMNH 
28823); Kapuas 1976-24, 102 mm (MZB 3057); Kapuas 1976-27, 62.5 mm (MZB 
3058); Kapuas 1976-31, 54,9 mm (MZB 3059); Kapuas 1976-35. 2: 42.3-49.0 
mm (BMNH 1982,3.29,2, MZB .3060); Kapuas 1976-55,4; 3 1.0-39.7 mm (MZB 
3061. USNM 230161), 

Distribution, — Peninsular Thailand (Tapi), Malay Penin- 
sula (Perak, Pahang), Sumatra (Padang, Lahat-Enim, Palem- 
bang. Taluk, Upper Langkat, Deli), Borneo (Kapuas. Mahakam, 
Kinabatangan''). .lava (Bantam. Batavia, Butenzorg, Tjampea, 
Lebak, Tjandjur, Parongkalong, Surabaya, Ngantang). 



Crossocheilus species undetermined 

(Figure 16) 

''Croimchilus oblongus var, nigriloba Popta, 19(14:200 (type locality Bo R., Ma- 
hakam basin). 

Material Examined— Western Borneo; Kapuas 1976-42, 4: 42.9-50.9 mm 
(CAS 49193. MZB 3062), 

The four juvenile specimens of Crossocheilus under consid- 
eration are quite distinct from juveniles of C cobilis and C. 
oblongus of similar size obtained by the Kapuas survey of 1 976, 
and do not seem identifiable with any recognized species in the 
genus. In having rostral barbels only they agree with C. oblongus 
rather than C. cobilis: in having the midlateral longitudinal black 
stripe extending well onto the middle portion of the caudal fin 
(more on the lower lobe than on the upper) they differ from 
both of these species and call to mind the series of three larger 
specimens 119-139 (RMNH 7589) from the Mahakam de- 
scribed as C oblongus var. lugnloba by Popta (1904; see also 
Popta 1906:120. pi. 8, fig. 27), In these Mahakam specimens, 
which I have not examined, the midlateral longitudinal stripe 
is darker and more sharply defined than in C. oblongus. and it 
extends on the middle caudal fin and particularly over the lower 



ROBERTS- WESTERN BORNEO FISHES 



33 




Figure 16. Cwssocheilus. Above, C cobilis. Kapuas 1976-27, 99.8 mm (MZB 3052); middle, C. oblongiis. Kapuas 1976-6, 66.3 mm (MZB 3055); below, C sp. 
undet.. Kapuas 1976-42, 50.9 mm (MZB .3062), 



caudal-fin lobe much more extensively but in a manner sugges- 
tive of these Kapuas juveniles. 

Cyclocheilichthys Bleeker, 1859 

Cyclocheilichtim Bleeker. 1859a:148 (type species Baihiis ciiopliis Bleeker. 1850, 

by subsequent designation of Bleeker. 1863ft; 199). 
Siaja Bleeker, 1859u:149 (type species Capocia mtcrotepis Bleeker, 1851, by sub- 
sequent designation of Bleeker. 1863/': 199 [not Capoeia siaja Bleeker 1851, a 

nomen nudum not validated until Bleeker, 1860a]. 
Aneinatichthys Bleeker, 1859u;149 (type species Barbus apogon Valenciennes in 

Cuvier and Valenciennes, 1842, by subsequent designation of Bleeker, 1863ft: 

199). 
Oxybarbus Vaillant. 1893:83 (type species Burhm hcli'wncma Bleeker, 1853, by 

monotypy). 
Neobaiynottis Banarescu. 1980:475 (type species Capoeia miovlt'pis Bleeker, 1851, 

by monotypy). Objective synonym o( Siaia Bleeker, 1859; also, see present 

account of Cyclocheilichthys niicrolepis 



NoMENCLATURAL NoTE. — Blccker ( 1 859a) indicated Siaja and 
Aneinatichthys as subgenera of his new genus Cyclocheilichthys. 
Although he did not define the subgenera he referred two or 
more described species to each of them. 

Bleeker ( 1 860a) treated five species in Siaja and four in Ane- 
inatichthys. So far as I have been able to determine a type species 
has not been previously designated for Aneinatichthys prior to 
Bleeker (1863ft). 

This southeast Asian genus includes nine species, seven of 
which occur in western Borneo. Four of these. C. enoplos. het- 
eronema. janthochir, and microlepis. are very distinct and can- 
not be confused with any other species. The remaining three, 
C. apogon, armatiis. and repasson, are very similar and difficult 
to distinguish. They are apparently closely related to each other; 
they share a distinctive color pattern not found in other species. 



34 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 17. C'ycliidu-ilichlln'S a, C. apogon. Kapuas 1976-37, 63.6 mm (MZB 3068); b. C drnialtn. Kapuas 1176-30. 78.0 mm (MZB 3077); c, C armalus, Kapuas 
1976-3:^, 78.5 mm (CAS 4')|y8); d. C aniiutin. Kapuas 1476-49, 78,4 mm (MZB 3080), 



The characters used lo separate them in the following key may 
not always work. In preparing the present account I have drawn 
freely on the unpublished doctoral dissertation of Sontirat (1976). 



Key to Cyclocheilidnliys of western Borneo 

la Middle of caudal peduncle with a large, round, dark or 
dusky spot; body with about 9-1 1 thin longitudinal stripes 
formed by small round spots centered on scales 

lb Caudal peduncle without spot; body without thin longi- 
tudinal stripes or with only a single longitudinal stripe on 
lateral line scale row 



2a Circumpeduncular scales 16 4 

2b Circumpeduncular scales usually 20 C. repasson 
3a Barbels absent C. apogon 
3b Maxillary barbel present; rostral barbel sometimes pres- 
ent _ C. armatus 

4a Barbels present _ _ 5 

4b Barbels absent _ „ -., 6 

5a Barbels multifid; last simple dorsal-fin ray slender, not 

greatly enlarged; lateral line tubes simple C. heteroncma 
5b Barbels simple; last simple dorsal-fin ray enormously en- 
larged and stout; lateral line tubes usually bifid 

C. enoplos 



ROBERTS- WESTERN BORNEO FISHES 



35 



70° 80° 90° 100° 110° 120° 130° 




^^^^, 




C- armatus 



O^ -II 



'^^^^ 



'^ -„ 



Figure 18. Cychcheilichlhysapogon. Geographical distribution (after Sontiral 
1976). 



Figure 19. Cychxheilichthys armatus. Geographical distribution (after Son- 
tirat 1976). 



6a 



6b 



Dorsal-fin rays 10-1 1; lateral line scales 32-33; a longi- 
tudinal line composed of spots centered on lateral line 

scales C. janihochir 

Dorsal-fin rays 18-22; lateral line scales 55-59; no lon- 
gitudinal line(s) on body C. micwlepis 



Cyclocheilichthys apogon (Valenciennes, 1842) 

{Figure 17) 

Barhus apogon Valenciennes in Cuvier and Valenciennes, 1 842:392 (type locality 

Java). 
Systomus apogon Sleeker. 185h:428^29. 
Syslomus apogonoides Bleeker. 1855c:410 (type locality Java). 
Cyclocheilichthys {Anematichthys) apogon Bleeker, 1859a;149; 1860fl:378. 
Cyclocheilichthys (Anematichthys) apogonides Bleeker, 1859u:149; 1860a:379. 
Cyclocheilichthys rubripmnis Fowler. 1934t:343-344. fig. 7 (type localit\ Ban 

Thuang Luang, Thailand). 

Material Examined.— Western Borneo: Kapuas 1976-7, 3: 18.9-28.7 mm(MZB 
3063); Kapuas 1976-8, 5: 16.7-52.0 mm (MZB 3064, USNM 230162), Kapuas 
176-16, 3: 84.0-100 mm (CAS 49194, MZB 3065); Kapuas 1976-17, 4; 41.0- 
99.1 mm (CAS 49195, MZB 3066); Kapuas 1976-28, 30.7 mm (MZB 3068); 
Kapuas 1976-37. 63,6 mm (MZB 3068); Kapuas 1976-42, 53.5 mm (MZB 3069); 
Kapuas 1976-44. 2; 72. .1-79. 2 mm (MZB 3070, USNM 230163). 

Distribution (Fig. \S,). — Cyclocheilichthys apogon is widely 
distributed in Thailand, Malay Peninsula. Sumatra, Borneo, and 
Java. 



ICyclocheilichthys lineatiis Weber and de Beaufort, 1916:162, See Sontiral (1976). 
Cyclocheilichthys dezwaani Weber and de Beaufort m Maass. 1912:525 (type 

locality Solok and Lake Singkarah, Sumatra), 
Cyclocheilichthys tapiensis Smith, 1931:1 1 (type locality Tapi River, peninsular 

Thailand). See Sontirat (1976). 
Cyclixheilichthys mekongensis Fowler. 1937:187, fig, 126, 127 (type locality Ke- 

marat=Mekong basin). See Sontirat (1976), 

Material Exampmed,- Western Borneo: Kapuas 1976-6, 2: 51.6-59.0 mm (CAS 
49196, MZB. W71); Kapuas 1976-10, 60,0 mm (MZB 3072); Kapuas 1976-14,2: 
29,3-35.7 mm (MZB 3073); Kapuas 1976-21. 19.0 mm (MZB 3074); Kapuas 
1976-22, 4; 46,5-56.7 mm (CAS 49197, MZB 3075); Kapuas 1976-25, 80,3 mm 
(MZB 3076); Kapuas 1976-30, 78.0 mm (MZB 3077); Kapuas 1976-32, 8: 45,4- 
78.5 mm (CAS 49198, MZB .W78, USNM 2.30164); Kapuas 1976-44, 45: 52.8- 
113 mm (AMNH 48918, BMNH 1982,3.29.3-5. CAS 49199, FMNH 94194, 
IRSNB 19727, MNHH 1982-653, MZB ,W79, ROM 38612, UMMZ 209913, 
USNM 230165); Kapuas 1976-49, 78.4 mm (MZB 3080). Philippines: Palawan 
Isd.. west coast Iwahig Rixer (CAS 53949). 

Distribution (Fig. 19).— This species is widely distributed 
in Thailand. Malay Peninsula. Sumatra, Borneo, and Java. It is 
also known from the single specimen reported above from Pa- 
lawan; this represents the only known record of the genus Cy- 
clocheilichthys in the Philippines. The specimen, collected by 
Robert B. Fox in 1963, was found by me among unidentified 
material in the Bureau of Fisheries and Aquatic Resources in 
Intramuros. Manila, in 1975, It is curious that C. armatus has 
not been found in northern Borneo, 



Cyclocheilichthys armatus (Valenciennes. 1842) 

(Figure 17) 

Barhus armatus Valenciennes m Cuvier and Valenciennes, 1 842: 1 63 (type locality 
Java), 

Barhus valenciennesii Bleeker, 1850:17 (type locality Surabaya, in flumine kali- 
mas). 

Capoeta siaja Bleeker, 1851;:432 (nomen nudum). 

Capoeia deventeri Bleeker, I855<::413 (type locality Java, Grati). 

Cyclocheilichthys {Siaja) siaja Bleeker 1859a:149 (nomen nudum). 

Cyclocheilichthys iSiaja) Deventeri Bleeker, 1859a: 148; 1860a:365, 375. 

Cyclocheilichthys (Cyclocheilichthys) armatus Bleeker, 1860a:364, 368. 

Cyctocheilichlhys (Siaja) siaja Bleeker, 1 860o;365, 374 (ty pe locality western Bor- 
neo, Sumatra). 

''Barhus UneatusVopu. 1905:171 (type locality Bo River, Mahakam basin; preoc- 
cupied by Barhus Imealus Dunckcr, 1904). See Sontirat (1976). 

''Barbus Fonlerii Popta, 1906:218 (replacement name for Barhus Imealus Popta, 
1905). 



Cyclocheilichthys enoplos (Bleeker. 1850) 

Barhus enoplos Bleeker, 1850:16 (type locality Surabaya, m flumine Kalimas), 
Capoeta enoplos Bleeker, 1851;:431 (description of specimen from Sambas), 
Barhus macracanthus Bleeker. 1853/:516-517 (type locality Palembang). 
Cyclocheilichthys (Cyclocheilichthys) enoplos Bleeker, 1859c;: 148; 1860u:366. 
Cyclocheilichthys (Cyclocheilichthys) macracanthus Bleeker, 1859a; 149; 1860a: 

367. 
Cyclocheilichthys dumerili Sauvage. 1881:182 (type locality Bangkok). 
Barhw. enoploides Tirant, 1885:157 (type localit> "Thu-dau-mot"). See Sontirat 

(1976). 

Material E.xamined — None. 

Last simple dorsal-fin ray greatly enlarged and very stout, and 
lateral line tubules usually bifid (rather than simple). Lateral 
line scales 35-37; circumferential scales 26-27; circumpedun- 



36 



CALIFORNIA ACADEMY OF SCIENCES 




Fu.uKi 2(1. ( viiochcilichllivs luiithnchir Above, Kapuas 1476-32. 59 4 mm IMZB 4iiii:i, tx-low. uncuhreroiis upper Up. Kapuas l976-4<), 76.8 mm (CAS 44180). 



cular scales 16. Gill rakers on first arch 15-20; vertebrae 36- 
37. 

DisTRiBiiTioN. — Laos. Kampuchea. Vietnam (Mekong basin). 
Thailand (Mekong. Chao Phrya, Meklong, Pasak). Borneo 
(Sambas). Sumatra (Palembang). Java (Surabaja, Ngawi). The 
only Borneo recorid is that of Bleeker (1852:431-432). Appar- 
ently unknown from Malay Peninsula. 

Cyclocheilichthys heteronema (Bleeker, 1853) 

Harhu\ helcroiienia Bleeker. I853i':446 (type locality Sambas, m fluviis). 
CYcl.nhcilwhlliYS{Swia) hclcrniwnui Bleeker, 1854i;;149; l860</;.377. 
Owhiirhii', hctcrmicina Vaillant, 1893:8.3 

Material Examined —Weslern Borneo: Kapuas 1976-9, 5; 46.9-63.(3 mm 
(AMNH 48919. BMNH 1982.3.29,6-7. MZB 3081): Kapuas 1976-15,20: 42.9- 
58.3 mm (CAS 4920(3. FMNH 94195. IRSNB 19728. MCZ 58339. MZB 3082). 
Kapuas 1976-33, 6: 46.6-83.0 mm (MNHN 1982-654. MZB 3083. CAS 349509): 
Kapuas 1976-34. 5: 40.7-57.6 mm (MZB 3084, RMNH 28824): Kapuas 1976- 
35, 7: 518-712 mm (MZB 3085, ROM 38607. HMMZ 209892); Kapuas 1976- 
37. 5: 60.2-73.1 mm (MZB 3086. LISNM 230166). Kapuas 1976-41. 41.5 mm 
(MZB 3087): Kapuas 1976-44, 4: 55,1-69,8 mm (MZB 3088, ZMA 116 516): 
Kapuas 1976-49, 4: 49 6-78.2 mm (MZB 3089, CAS 49201) 

DifTers from all other Cyclocheilichthys in having maxillary 
barbel multifid instead of simple. Rostral barbel absent. Lateral 
line scales 32-34, Circumferential scales 1 1/2/13=26; circum- 
peduncular scales 16. Gill rakers on first arch 8-12. Total ver- 
tebrae 33-35. 

Distribution. —Thailand (Tale Noi, Pattani; Smith 1 945: 1 42- 
143). Malay Peninsula (Muar Johore, Kuala Pilah, Negri Sem- 



bilan; Herrc and Myers 1937:61 
Kapuas). 



Western Borneo (Sambas. 



Cyclocheilichthys janthochir (Bleeker, 1853) 

(Figure 20) 

Syslonuis lanlhochir Bleeker. 1853i':448 (type locality Kapuas River), 
Cn-liKheilichlhys l-Uwimiliclulivs) lamhncliir Bleeker, I859u:I49; 1860u:381, 

Material ExAMiNED-Western Borneo: Kapuas 1976-20. 9: 100-119 mm 
(BMNH 1982,3.29.8, FMNH 94196, KIIMF2848, MNHN 1982-655, MZB 3090. 
RMNH 28825, LIMMZ 209862): Kapuas 1976-32. 59,4 mm (MZB 4002): Kapuas 
iq7fi-49, 8: 67 9-78 2 mm (CAS 44180, KUMF 2849, MZB 3091 , USNM 230167). 

Barbels absent; circumpeduncular scales 16; circumferential 
scales 12/2/13=27, Lateral line scales 32-33. Last simple dorsal- 
fin ray stifiTcned and finely serrate, but very slender. Coloration 
unique for the genus: live fish with body white laterally and 
ventrally, with bluish reflections dorsally; a thin black mid- 
longitudinal line straddling lateral line canal; dorsal and caudal 
fins carmine red. with blackish margins; other fins colorless. Gill 
rakers 1 1. Vertebrae 22+ 1 1=33 (3. Kapuas). 

Distribution. — Borneo (Kapuas; southern Borneo. Kottelat 
1982). 

Cyclocheilichthys microlepis (Bleeker, 1851) 

(Figure 21) 

Capocia inurolepis Bleeker. 1 85 I /j, 206 (l\pe loealily Bandjermassing). 
CvclocheihchtlmiSiaia) niicmlcris Bleeker, 1859^:149: I860</:37I, 
Rarvnolus i>ucriilcpis Gun[hcr. 1868:61. 
\'v<>hiirvn<>fn\ >uicriilcpi\ Banarescu. 1980:477. 



ROBERTS-WESTERN BORNEO FISHES 



37 




Figure 21. Cyclocheilichlhys inicrolepis. Kapuas 1976-54, 256 mm (MZB 309-1), 



Material Examined.— Western Borneo: Kapuas 1976-34, 332 mm (MZB 3092); 
Kapuas 1976-44, 3: 1 12-144 mm (CAS 49202, MZB 3093, USNM 230168). 

Differs from all other Cyclochelllchlhys in having 18-22 dor- 
sal-fin rays (vs. only 11-13) and lateral line scales 55-59 (vs, 
30-40). Circumferential scales about 13/2/18 = 33, Circumpe- 
duncular scales 22, Last simple dorsal-fin ray stiffened and finely 
serrate but very slender. Gill rakers on first arch 10, Vertebrae 
22+12=34, 

Distribution,— Sumatra (Palembang, Djambi). Borneo (Ka- 
puas, Barito). 

Cyclocheilichthys repasson (Bleeker, 1853) 

Barbus repasson Bleeker. 1853/):295 (lype locality Pangabuang, Lampong prov- 
ince. Sumatra). 
Cyclocheilichthys (Cychcheilichihys) repasson Bleeker. 1859ii:149; 1860ii:370, 
Cyclochelllchlhys inegalops Fowler. 1905:483 (type locality Baram River). See 
Sontirat (1976). 

Material Examined.— Western Borneo: Kapuas 1976-24. 102 mm (MZB 3095); 
Kapuas 1976-33, 128 mm (MZB 3096); Kapuas 1976-36,61.7 mm (MZB 3097); 
Kapuas 1976-39, 85.5 mm (MZB 3098); Kapuas 1976-43, 47.9 mm (MZB 3099). 

Distribution (Fig, 22), — Vietnam, Laos, and Kampuchea 
(Mekong); Thailand (Mekong, Chao Phrya, Meklong, Pattani, 
Tale Sap); Malay Peninsula (Perak), Sumatra (Lake Singarah; 
Moesi R.). Borneo (Baram, Kapuas, Mahakam?), 

Dangila Valenciennes. 1842 

Lahioharhiis van Hasselt, 1823:132 (unidentifiablc). 

Dangila Valenciennes in Cuvler and Valenciennes, 1 842:229 (type species Dangila 

lcpiochcila= Dangila cuvieri Valenciennes in Cuvier and Valenciennes, 1 842, by 

subsequent designation of Bleeker, 1863/):193). 
Cyrene Meckel, 1843:1024 (type species not designated). 

Most authors, following Cuvier and Valenciennes ( 1 842). have 
recognized Dangila. Smith (1945:221). however, unwilling to 
accept its rejection, employed Lahioharhus. Inger and Chin ( 1 962: 
94) acknowledged Smith's claim of priority for Labiobarbus but 
treated it as a nomen oblitum and retained Dangila. Ichthyol- 
ogists of the last century (including Bleeker) obviously knew 
about the "priority" of Labiobarbus but set it aside in favor of 
Dangila. The paragraph in van Hasselt (1823:132) treating La- 
biobarbus reads as follows: 



Het genus Lahioharhus Mihi bestaat mit Labiones met 4 barhillons. 
met eene dorsal vin. waarvan de 2de slraal met getand is, verecnigt 
dus de karakteren van Labw. en barbus. waarom ik den naam La- 
biobarbus het aangenomen. De namcn L Leplocheilus M. en Lipochei- 
lus M. anderschieden de twee spccien, die heide bij Batavia in de nvier 
gevonden worden, en onder mij afgebeeld zijn. 

The genus Labiobarbus Mihi consists of Labiones with 4 small bar- 
bels, with a single dorsal fin of which the second ray is not serrated 
and thus combines the characters of Labia and barbus, wherefore I 
adopted the name Labiobarbus. The names L. Leplocheilus M. and 
Lipocheilus M. distinguish the two species which were both found in 
the river near Batavia and figured under me (after .•Xlfred 1961). 

Many, perhaps most, members of the family Cyprinidae and 
subfamily Labeoinae have 4 barbels and the dorsal fin without 
a serrated ray; the identities of the two species mentioned by 
van Hasselt are known only because of the subsequent publi- 
cation by Valenciennes ( 1 842) of descriptions based on the orig- 
inal figures referred to by van Hasselt, Valenciennes used the 
species names originally proposed by van Hasselt. but coined 
the new generic name Dangila. presumably to avoid confusion 
or synonymy with Labcobarbus Riippell. 1836. which he treated 
in the same volume. 

The names Labiobarbus. L. leplocheilus. and L. lipocheilus of 




Figure 2 
tirat 1976). 



Cyclocheilichthys repasson. Geographical distnbulion (after Son- 



38 



CALIFORNIA ACADEMY OF SCIENCES 



van Hasselt, 1823 are regarded as unidentifiable and best for- 
gotten. 

Dorsal fin with 21-30 branched rays, last simple ray slender, 
flexible, and non-serrate. Two pairs of barbels, both moderately 
elongate. Gill rakers on first arch 38-47. Soft roof of mouth 
swollen, covered with hundreds of macroscopic villi, and fitting 
snugly against raker-bearing portions of gill arches. Scales in 
lateral series 39-68. Pharyngeal teeth 3,3,5/5,3,3 (Meckel 1843: 
34) or 2,3-4,5/5,3-4,2 (Beaufort 1916:113). 

Dangila cuvieri Valenciennes, 1842 

''.Lahioharbus Icplochedus van Hassell. IS2.1;1.^2 (unidenlifiable). 

Dangila Curieri Valenciennes in Cuvicr and Valenciennes, 1 842:230 (type locality 

Batavia?!. 
Diingila Icptocheila Valenciennes in Cuvier and Valenciennes. 1842:2.14 (type 

locality Java). 
Dangila kocdiem Popta. 1 904: 1 92 (t> pc locality "le Boelif"='.'Sungai Bulit, Kapuas 

basin, at 0°48'N, II3M4'E). 
Dangila rosea Popta. 1904:193 (type localit\ "le Bo". Mahakam basin). 
Lahioharhiis leplocheilus Smith. 1948:227, 

Material Examined. — None. 

Dangila fasciata Bleeker, 1853 

Dangila lasciala Bleeker. l853/):297 (type locality Pangabuang. Lampong Prov). 
Dangila taeniata Giinther, 1868:38 (unwarranted substitute name). 

Material Examined.- Western Borneo: Kapuas 1976-14, 37,1 nim(MZB3l01); 
Kapuas 1976-16, 5: 47.1-77.1 mm (CAS 49204. MZB 3102. HSNM 230169); 
Kapuas 1976-37. 80,6 mm (MZB 3 103): Kapuas 1976-39. 2: 63,0-68,6 mm (MZB 
3104. HMMZ 209897), 

Dangila festiva (Meckel, 1843) 

Cvrcne fcsliva Meckel, 1843:1025 (t\pc locality Borneo), 
Dangila lesuva Bleeker. 1857o:l6 (Kahajan), 

Dangila fcslira var. sieniis-iniisi.arniii Vaillanl. 1902:87 (t\pc locality "Ponti- 
anak": "Smitau. riviere Sibau"), 

Material Examined —Western Borneo: Kapuas 1976-19, 172 mm (MZB 3105). 

Distribution. — Malay Peninsula (Pahang. Selangor, Johore). 
Borneo (Kapuas, Rahajan, Barito). 

Dangila lineata Sauvage, 1878 

Dangila liiwala Sauvage, 1878:237 (type locality Stung-Strang, Laos), 
Lahmharhin liiicatin Smith. 1945:223, 

Material Examined. — Western Borneo: Kapuas 1976-14, 2: 72.6-84.3 mm 
(CAS 49205. MZB 3 106); Kapuas 1976-15,67.1 mm (MZB 3 107); Kapuas 1976- 
34, 4: 69,6-78.8 mm (MNHN 1982-656. MZB 3108, USNM 230170); Kapuas 
1976-35. 2: 79.7-80,4 mm (CAS 49206. MZB 3109); Kapuas 1976-37, 72,6 m 
(MZB 3110); Kapuas 1976-39. 72,9 mm (MZB 3111) 

Distribution. — Laos. Thailand (Mekong, Chao Phrya, Tapi, 
Rayong). Malay Peninsula. Borneo (Kapuas). 

Dangila ocellata (Meckel, 1843) 

Cvrcne oa'llala Meckel. 1843:1025 (type localit\ Borneo). 

Dangila inierolepis Bleeker. IS52i/595 (type locality Palembang), See Bleeker 

(186(.)ij:I95), 
Dangila ocellala Bleeker. 1860i7:194, 

Material Examined —Western Borneo: Kapuas 1976-20. 6: 63,7-110 mm 
(AMNM 48920. BMNH 1982,3.29.9. CAS 49207, FMNM 94197, MZB 3112); 
Kapuas 1976-31,58,1 mm (MZB 31 13): Kapuas 1976-33, 2: 81, 9-93.0mm(MNMN 
1982-657, MZB 3114); Kapuas 1976-34, 2: 114-120 mm (IRSNB 19729, MZB 
3115); Kapuas 1976-39, 97.3 mm (MZB 3116); Kapu.as 1976-43. 3: 47.4-54.5 



(MZB 31 17. IIMMZ 209914. USNM 230171): Kapuas 1976-44. 90.3 mm (MZB 
3118); Kapuas 1976-49. 67.7 mm (MZB 3119), 

Lateral line scales 65-68 (30-53 in all other Dangila). 

Body plain except for a black spot surrounded by a light 
colored ring (yellow in life?) on shoulder and a plain black spot 
on middle of caudal peduncle. 

Distribution. — Sumatra (Pangabuang, Palembang, Lahat, 
Lematang Enim, Gunung Sahilan, Sungei Mahe, Danau Sialong 
Lotong, Kwantan R., Indrajiri, Djambi). Borneo (Kapuas, Ka- 
hajan, Mahakam). 



Eirmotus Schultz, 1959 

Eirmotus Schultz, 1959:10 (type species Einnoms octozona Schultz. 1959. by 
monotypy). 

Small cyprinids (largest known specimen of only known species 
33 mm) with several longitudinal and transverse rows of ce- 
phalic cutaneous papillae; barbels absent; lateral line incom- 
plete, with only 5-6 pored scales anteriorly; scales relatively 
larger than in any other southeast Asian cyprinid, lateral scale 
series 20, predorsal scales 7, circumpeduncular scales 8, Dorsal- 
fin rays 8, last simple ray serrate. Anal-fin branched rays 5. 
Pharyngeal teeth unknown. Gill rakers absent or extremely re- 
duced. Vertebrae 15+15-16=30-31. 

Eirmotus. Orcichthys. and Cydochcilichthys are the only 
southeast Asian cyprinid genera with parallel rows of enlarged 
cephalic cutaneous papillae. Whether they are closely related 
(perhaps forming a monophyletic group) or independently de- 
rived from cyprinid genera lacking papillae is an open question. 
It is noteworthy that numerous species of tropical African "Bar- 
bus" (sometimes placed in separate genera on this account) have 
parallel rows of enlarged cephalic cutaneous papillae which are 
at least superficially very similar to those observed in these Asian 
genera. 

Eirmotus octozona Schultz, 1959 

(Figure 23) 

Eirmoliis oclnzona Schultz, 19591 I (t\pc locality, possibly erroneous, Bung Bo- 
rapet, Thailand; described from aquarium specimens). 

Material Examined -Western Borneo; Kapuas 1975-16 (CAS 49208, MZB 
3120. IISNM 230172); Kapuas 1976-17. 32.1 (MZB 3121); Kapuas 1976-18. 2: 
19,8-19,9 mm (CAS 49209, MZB 3122); Kapuas 1976-51. 25,4 mm (MZB 3123). 

The color pattern of this species, while somewhat similar to 
that of some Puntius (including P. cndecanalis. with which it 
occurs sympatncally) is quite distinctive. It includes all of the 
vertical bars found in the species of Puntius with barred color 
patterns, but in addition there are usually one or more midlateral 
roundish spots between the bars, and separate midventral spots 
on the belly, straddling the pelvic-fin base (and extending onto 
the pelvic fins), and on the caudal peduncle. Separate midventral 
spots are not found in any other cyprinid in western Borneo. 

Distribution,— Thailand'' (Bung Borapet?), Borneo (Kapuas; 
southern Borneo, Kottelat 1982). 

Epalzeorhynchos Bleeker, 1855 

Epalzeorltyiichos Bleeker, 1855/i:258. 270 (type species Barhiis kaloplerus by 

monotypy). 
Epalzeorhynchiis Bleeker, 1860u:1 18 (unjustified spelling emendation). 



ROBERTS- WESTERN BORNEO FISHES 



i^ 




Figure 23. Einnolus ociozotm Kapuas 1976-17. 32.1 mm (MZB 3121). 



Epalzeorhynchos belongs to a group of Asian genera (includ- 
ing Crossochcilm and Paracrossochilus) which have the upper 
jaw entirely covered by a greatly enlarged fimbriate rostral cap. 
It differs from all other genera in havmg a large fleshy rostral 
papilla ("moveable lateral lobe") between rostral barbel and 
sublacrimal groove. Posteriorly directed distal tip of rostral pa- 
pilla almost invanably with a single large sharply pointed tu- 
bercle; additional tubercles sometimes on other parts of rostral 
papilla. Sublacrimal groove extending from origin of rostral 
papilla to behind rictus of jaws. Upper lip absent or vestigial. 
Horny jaw sheaths with straight or gently curved and broad 
margins with sharp cutting edges. Lower lip a broad, non-pa- 
pillose fleshy lobe lying in concavity on ventral surface of lower 
hornyjaw sheath. Rostral and maxillary barbels well developed; 
rostral barbels black, often in sharp contrast to rostral cap; max- 
illary barbels pale. 

The rostral cap and papilla and other features of the mouth 
and snout of E. kalopterum are illustrated by Weber and de 
Beaufort (1916, fig. 94). Their illustration erroneously depicts 
the middle portion of the rostral cap with a horizontal groove 
extending between the rostral barbels. The rostral cap is entire 
in all species of Epalzeorhynchos. 

Epalzeorhynchos is closely related to Crossocheilus, which 
differs from it in having a well developed upper lip (fimbriate), 
somewhat narrower jaws, and numerous papillae on lower lip, 
as well as in lacking a discrete rostral papilla. In Crossocheilus 
as in Epalzeorhynchos there is a well developed sublacrimal 
groove, and lying between its anterior portion and the rostral 
barbel is a round rostral lobe similar to the free rostral papilla 
of Epalzeorhynchos. 

The essential diagnostic features of the genus were clearly 
recognized by Bleeker (1855). 

This genus, which is being studied by J. Kamasuta. comprises 
at least four species, including Labeo bicolor Smith, 1934, and 
Lahco frenatus (with which Labeo munensis Smith, 1934 and 



Labeo erythrurus Fowler, 1934 are synonymous), both from 
Thailand (R. M. Bailey and J. Kamasuta, pers. comm.), as well 
as the more widely distributed type species, E. kalopterum. the 
only species known from Borneo. 

Epalzeorhynchos kalopterum (Bleeker, 1850) 

Barbus kaloptenis Bleeker. 1850:13 (type locality Banjcrmassmg. m fluviis). 
Epalzeorhynchos kalopleriis Bleeker. 1855/i:270. 

Epalzforhynchus kallopicrus Bleeker. I860i(;ll8 (unjustified spelling emenda- 
tion). 

Material Examined. — Malay Peninsula: Johorc. near Kulai. 204: 20.2^5.6 
mm (CAS-SU 39350). Western Borneo: Kapuas 1976-14. 3: 52.7-72.1 mm (AMNH 
48291. BMNH 1982.3.29.10. MZB 3124): Kapuas 1976-15, 3: 52.8-55.5 mm 
(CAS 49210. MZB 3125): Kapuas 1976-16, 3: 46.1-64.5 mm (FMNH 94198, 
IRSNB 1 9730. MZB 3 1 26): Kapuas 1 976-24. 113mm (MZB 3127): Kapuas 1976- 
27. 121 mm (MZB 3128): Kapuas 1976-29. 6: 63.0-92,5 mm (MCZ 58340, 
MNHN 1982-658. MZB 3129, RMNH 28826. ROM 38608): Kapuas 1976-33. 
4: 55.2-74.0 mm (MZB 3130. CAS 4921 1. HMMZ 209885): Kapuas 1976-36. 
71.9 mm (MZB 3131): Kapuas 1976-43. 4: 41.1-60.5 mm (MZB 3132. USNM 
230173. ZMA 116.517). 

Note: As the gender of Epalzeorhynchos is neuter, the correct spelling of the 
species name is kalopterum (R. M. Bailey, pers. comm.). 

This handsome species is distinguished from all congeners by 
Its very distinctive coloration. Side of body with a broad lon- 
gitudinal black stripe with sharply defined very straight margins, 
continued less distinctly to snout-tip and on middle caudal-fin 
rays to end of caudal fin. A broad longitudinal stripe above 
midlateral stripe and entire body below pale greyish or white. 
Dorsal surface of body with dark pigmentation with a sharply 
defined straight ventral border. Middle of dorsal, pelvic, and 
anal fins with a black scythe-shaped mark. 

Sexual dichromatism and dimorphism apparently absent. 
Coloration as described above and rostral papilla with terminal 
or distal tubercle fully developed in smallest specimens exam- 
ined of both males and females. 

Distribution. — Peninsular Thailand (Tapi R.; Smith 1945: 



40 



CALIFORNIA ACADEMY OF SCIENCES 



264). Malay Peninsula (Johore). Sumatra (Palembang, Lahat, 
Batang Hari. Kwantan R.). Borneo (Kapuas. Kahajan, Barito). 

Garra Hamilton-Buchanan, 1822 

Garra Hamilton-Buchanan, 1822:343. 3'J3 (type species Cypnmis (Garra) lamta 

Hamilton-Buchanan. 1822. by subsequent designation ol'Bleeker. 1863^:192). 
Plalnara McClelland, 1 839:947 (type species Plalycara nasula McClelland, 1839. 

by subsequent designation of Jordan. 1919:195). 
Discognalhus Heckel, 1843:1027 (type species Discognalhus ranahilis Meckel, 

1843. by subsequent designation of Sleeker, 1863/): 192). See Hera, 1921:648. 
Lissorhynchus Bleeker, 1859-60c:422; 1860a:85-86 (as Lissorhynchiis on p. 86) 

(type species Plalycara lissorhynchus McClelland, 1 839, by absolute tautonymy). 
Discognalhichthys Bleeker. 1859-60i:423: 1860u:128 (type species Discognathus 

variabilis Heckel, 1843, by subsequent designation of Bleeker 1860i;:128). 
Parapsilorhynchus Hora, 1921:13 (type species Psilorhynchm icntaculatus An- 

nandale, 1919, by original designation). 
Discolabeo Fowler, 1937:210 (type species Discolabeo fishcn Fowler. 1937, by 

original designation and monotypy). 

Menon (1964:187) and Jayaram (1981) stated that Cyprinus 
lamta Hamilton-Buchanan, 1822 is type species of Garra Ham- 
ilton-Buchanan, 1822 by monotypy. Reference to the original 
publication, Hamilton-Buchanan (1822:343-349, 393-394), 
however, shows that no fewer than seven species were included 
in this "subgenus" Garra (only one with a question mark). Cy- 
prinus lamta. the first species mentioned, seems to be the only 
species indicated as type species by subsequent authors, of whom 
Bleeker (1863/') apparently is the earliest. 

Rostral cap with unculiferous fimbriate margin. Upper lip 
varying from well developed with unculiferous fimbriate margin 
to vestigial or absent. Homy jaw sheaths of both jaws well 
developed. Lower lip with tuberculate anterior margin, enor- 
mously expanded and incised posteriorly to form a thick callous 
pad or mental adhesive disc; ventroposterior margin of adhesive 
disc with unculiferous tubercles. 

Rostral and maxillary barbels present, rostral barbels only 
present, or both absent. In some species the mental adhesive 
disc is weakly developed, vestigial, or perhaps even absent; this, 
coupled with ontogenetic changes associated with the mental 
disc (see Hora 1921) and the highly variable timing of its ap- 
pearance, has contributed to considerable systematic confusion 
at the genus as well as species level. It seems likely that all 
species of African and Asian Labeoinae with a mental adhesive 
disc belong to this one genus. Menon (1964) recognized 34 
species in his revision of the genus. 

A rostral cap with unculiferous fimbriae similar to that of 
Garra is found in Crossochcilus. Epalzcorhynchos. and Para- 
crossochiliis. Garra differs from these and all other Bornean 
cyprinid genera in having a mental disc (missing in very young 
fish). The posterior border of the disc is heavily papillose. The 
fringes of the rostral cap and papillae of the mental disc are 
heavily unculiferous (Roberts 1982a) and presumably function 
in rasping food from rocky substrate as well as adhesion in the 
swift-flowing mountain streams typically inhabited by Garra. 
Only a single species is known from Borneo, and the genus is 
unknown from Sumatra and Java. 

Garra borneensis (Vaillant, 1902) 

Discognathus borneensis Vaillant. 1902:91 (type locality Bluu River. Mahakam 

basin). 
Garra borneensis Fowler, 1905:482, 

Material Examined. — Western Borneo: Kapuas 1976-24. 6: 46.0-69.8 mm 
(BMNH 1982.3.29.11-12. CAS 49212. MZB 3133); Kapuas 1976-25, 8: 49.7- 



109 mm (FMNH 94199. MNHN 1982-659. MZB 3134. RMNH 28827, UMMZ 
209867, USNM 230174). 

This is the only species of Garra known from Indonesia: 
according to Menon (1964), it is closely related to G. taeniata 
Smith, 1931 from Thailand. 

Distribution. — Borneo (Baram, Kapuas, Mahakam, Kina- 
batangan, Tempasuk). (Tempasuk is a small coastal drainage 
flowing N from the W side of Mt. Kinabalu.) 

Hampala Bleeker, 1859-60 

Hampala Kuhl and van Hasselt in \an Hasselt. 1823:132 (nomen nudum). 

Hampala Bleeker. 1859-60(:430. 1860a:275. 307-311 (type species Hampala 
macrolcpulota van Hasselt. \i2'i=Capoela macrolepidola Valenciennes in Cu- 
vier and Valenciennes. 1 842. by subsequent designation of Bleeker. 1 863/i:200). 

Superficially similar to Puntius but with a characteristically 
pointed snout not seen in any member of that genus, and at- 
taining greater size (to 70 cm total length). Pharyngeal teeth 
usually in three rows, 1,3,5/5,3,1, cochleariform. 

Hampala bimaculata (Popta, 1905) 

Barbus hampal var. himaciilala Popta. 1905:173 (type locality le Bo. Mahakam 

basin). 
Hampala bimaculata Weber and de Beaufort. 1916:146. 

Material Examined. — Western Borneo: Kapuas 1976-24. 71.2 mm (MZB 3135); 
Kapuas 1976-25.2: 1 2.3-1 24 mm (CAS 492 1 3. MZB 3 1 36). 

For a discussion of this taxon, its geographical distribution 
and relationship to Hampala macrolepidota. see Inger and Chin 
(1962:78-82). 

Hampala macrolepidota Valenciennes, 1842 

Hampala macrolepidota Kuhl and van Hasselt. m van Hasselt. 1823:132 (nomen 

nudum). 
Cap<>eta macrolepidota Valenciennes //; Cuvier and Valenciennes. 1842:280 (type 

locality Java). 

Material Examined —Western Borneo: Kapuas 1976-6. 4: 37.6-58.4 mm (CAS 
49214. MZB 3137); Kapuas 1976-27. 3: 31.5-78,3 mm (MZB 3138. USNM 
230175); Kapuas 1976-33. 123 mm (MZB 3139). 

Juvenile H macrolepidota exhibit a temporary color phase 
with 5 of the 6 vertical bars found in some vertically barred 
Puntius species (only the post-anal bar is absent); Taki and 
Kawamoto (1977, fig. 3). 

Kalimantania Banarescu, 1980 

Kalimanlania Banarescu. 191S1I:471 (type species Syslomus lanak Bleeker, 1855, 
by monotypy). 

As discussed above, this genus appears to be closely related 
to Albulichthys. Amhlyrhynchichihys. and Puntioplites. It is per- 
haps closest to Amblyrhynchichthys, from which it differs in 
having snout less truncate and nostrils not so far anterior, gill 
rakers much fewer, scales with divergent rather than convergent 
radii, and last simple anal-fin ray greatly enlarged ('"spmified"). 
It difliers from all of its presumed close relatives in having 7 
instead of only 5 branched anal-fin rays. 

Kalimantania lawak (Bleeker, 1855) 

Svstomus lawak Bleeker, 1855i:4l I (type locality "BataMa. in llumine Tjiliwong; 

Surabaja. in Huminc Kalimas"). 
Barbus lawak Gunthcr. 1868: 149. 



ROBERTS-WESTERN BORNEO FISHES 



41 



Amblyrhymhichlhys alius Vaillam, 1893^:59 (lype locality Kapuas). See Ban4- 

rescu ( 1 980), and below. 
Punlius lawak Weber and de Beaufort, 1916:203. 
Puniiopbles la»ak Banarescu, 1978:116. 
Kaliinanlaiim lawak Banarescu. 1980:475. 

Material Examined. — Western Borneo: Kapuas. 77.0mm(MNHN 1891:355, 
holotype of .-1. alnisY. Kapuas 1 976-34, 2: 39.3-40. 1 mm (CAS 49253, MZB 3268). 
Java: Tjiliwong River. Balavia or Kalimas River, Surabaya. 59.7 mm (BMNH 
1866.5.2.203, syntype). 

The holotype of Amblyrhynchichthys alius has dorsal-fin rays 
iii(or iv)8-l/2, anal-fin rays iii7-l/2. Last simple anal-fin ray 
stout, spine-like, deeply concave posteriorly with sharp lateral 
edges, non-serrate. Scales with striae parallel or slightly diver- 
gent, lateral series 34 or 35. transverse scales 8/1/6, predorsal 
about 13. and circumpeduncular 16. Gill rakers on first arch 
26. Coloration largely silvery on head and body; no vertical 
streak marks as in Pitntioplites bulit. A few days after examining 
this holotype in Paris, I examined the syntype of B. lawak in 
London, and concluded that they are conspecific. The BMNH 
syntype has 7-1/2 branched anal-fin rays and 23 gill rakers on 
first arch. 

Distribution. — Apparently known only from western Bor- 
neo (Kapuas) and Java. Records from Sumatra (and Riouw?) 
(Weber and de Beaufort 1916:204) are based on misidentified 
Piintioplires waandersi according to Banarescu (1978:1 16). 

Leptobarbus Bleeker, 1859-60 

Leplobarbus Bleeker, l859-60c:435; 1859u:285, 432 (type species Barbiis Hoe\- 

ciiii Bleeker, 1851. by monotypy) 
Filirasbora Fowler. 1937:172 (type species Fihrasbora rubnpiiifiu Fowler. 1937, 

by monol\py= Leplobarbus hocvcnii Bleeker, 1851). 

In general appearance like a very heavy-set Rasbora but with 
rostral and maxillary barbels well developed, longer than eye 
diameter; top of head broad and flat; lower jaw without sym- 
physeal knob. Last simple dorsal-fin ray non-serrate. Lateral 
line complete, curved downward but slightly, lateral line scales 
34-38; transdorsal 9-11; circumpeduncular 14-15. Gill rakers 
on first arch 14-17. Pharyngeal teeth cochleariform with cren- 
ulate margins, in three rows. 2.3.5/5.3.2. 

Four species, all but L. hoevenii confined to Borneo. The two 
species found in the Kapuas differ strikingly in coloration. 



Key to Leplobarbus 



L. hoevenii 



1 a Transdorsal scales 9 

lb Transdorsal scales 10-1 1 2 

2a Caudal-fin lobes with submarginal longitudinal black 

stripes (Kapuas) L. inelanoplerus 

2b Caudal fin dusky or plain „ 3 

3a Body with a narrow, black, mid-lateral zig-zag stripe 

(northern and eastern Borneo) 

L. melanolaenia Bouknger, 1894 
3b Body plain (northern Borneo) L. hosii Regan, 1906 

Leptobarbus hoevenii (Bleeker, 1851) 

Barbiis Hoevenii Bleeker, 185l/;:207 (type locality Bandjermassing, in fluviis). 
Leplobarbus Hoevenii Bleeker, 1860<i:433. 

Filirashora rubripnma Fowler, 1937:172 (type locality Kemrat, Siam). See remarks 
by Bailey in Bohlke (1984:90). 

Material Examined. -Kapuas 1976-15, 10: 18.6-36.3 mm (CAS 49215, MZB 
3140); Kapuas 1976-20. 3: 91 .6-1 36 mm (FMNH 94200. MNHN 1982-660. MZB 
3141): Kapuas 1976-44. 3: 81.2-158 mm (MZB 3142. RMNH 28828. UMMZ 



209915): Kapuas 1976-49, 3: 77.3-109 mm (CAS 49216, MZB 3143, USNM 
230176). 

In life, pelvic and anal fins blood-red; other fins dusky or 
colorless. 

Distribution.— Thailand (Mekong, Chao Phrya). Sumatra 
(Palembang, Indragiri, Kwantan, Taluk, Danau Sialong Lotong, 
Djambi. Bagan Api Api). Borneo (Baram?, Kapuas, Barito, Ka- 
hajan, Pengaron. Mahakam). 

Leptobarbus melanopterus Weber and de Beaufort, 1916 

Leplobarbus inelanoplerus Weber and de Beaufort, 1916:97 (type locality Kapuas 
R. at Selimbau), 

Material Examined. — Western Borneo: Kapuas 1976-32, 3: 66.1-71.1 mm 
(MZB 3144, USNM 230177); Kapuas 1976-49, 4: 120-131 mm (CAS 44179. 
MZB 3145, ZMA 116,518). 

In life, caudal fin deep magenta or purple with black sub- 
marginal longitudinal stripes on each lobe; other fins dusky or 
pale, colorless. 

Distribution. — Known only from the Kapuas. 

Lobocheilos Bleeker, 1853 

Lobocheilus Kuhl and van Hasselt m van Hasselt. 1823:132 (nomen nudum). 

Lobocheilos Bleeker, 1853/:520 (type species Lobocheilus falcifer Kuhl and van 
Hasselt in van Hasselt, lS2i= Labeo falcifer Valenciennes in Cuvier and Va- 
lenciennes. 1842. by subsequent designation of Bleeker, 1863:194). 

Gobionichthys Bleeker. I859<7:145 (type species uncertain). 

A genus of Labeoinae with rostral cap greatly thickened, its 
margin entire; upper lip also greatly thickened, its margin entire. 
Sublacrimal groove extending from rostral barbel (when present) 
to beyond rictus of jaws. Inner margin of upper lip with a valve- 
like transverse flap. No separate upper homy jaw sheath. Lower 
homy jaw sheath enormously expanded, its dorsal portion ex- 
tending as a convex pad into roof of mouth, its ventral portion 
with a broad, nearly straight, sharp-edged, heavily comified 
wedge. Lower lip fleshy, deeply incised, connected to ventral 
surface of lower jaw by a pair of large internal freni. Short rostral 
and maxillary or maxillary barbels only present. 

The species of Lobocheilos need systematic revision. More 
information is needed about ontogenetic changes in coloration 
and other characters, and the numerous nominal species de- 
scribed from Thailand by Fowler (1934, 1935, 1937, 1939) should 
be compared with each other and with the species known from 
Indonesia. 

Lobocheilos bo (Popta, 1904) 

(Figure 24 below) 

Tylognalhus bo Popta, 1904:199 (type locality Bo R., Kapuas basin). 
Lobocheilus bo Smith, 1945:239. 

Material Examined —Western Borneo: Kapuas 1976-14, 5: 59.8-71.6 mm 
(CAS 49218. MZB 3147); Kapuas 1976-15, 7: 17.9-22.2 mm (CAS 49219, MZB 
3148): Kapuas 1976-40, 23.9 mm (MZB 3149). 

Distribution. — Borneo (Kapuas, Mahakam; also north Bor- 
neo according to Inger and Chin 1962). 

Lobocheilos hispidus Valenciennes, 1842 

(Figure 24 above) 

Labeo hispidus Valenciennes m Cuvier and Valenciennes. 1842:356 (type locality 
Buiten/org). 



42 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 24. Lohocheilm Above, L hispiihis. Kapuas H76-6. 60,7 mm (CAS 44217); below, /, hi: Kapuas 147b-14, b5.7 mm (MZB ,M47). 



Chtiiuiroslomii hpochcilos Valenciennes //; C u\ ler and Valenciennes. 1844:400 

(tspe locality Java). 
Gnbio lavaniciis Bleeker, 1857/i;358 (i>pe localiu Bata\ia, m fluviis). 
Lohoclu'ilus {(lohiiinuhlhyi) jiumiicKS Bleeker. 185yu:l45. 
Liibochi'ilus {Gohi(vmliihys) lipochcihis Bleeker. 1860u:154. 
Tvlognallnis hispuiiis GOnlher, 1868i;:66. 
Lithochcilii'i hispuliis Fowler. 1<J05 :48-'' 

Material Examined — Wcslem Borneo: Rapuas 1476-6. 8: 55, 4-44. .3 mm (CAS 
44217. MNHN 1482-661, MZB 3146, USNM 2,^0178), 

Distribution. — Borneo (Baram. Kapuas, Mahakam). Java. 
Luc'osoma Bleeker, 1855 

Liiciosonia Bleeker, I855/i:263 (type species Barbiis sfiincriis Valenciennes in 
Cuvier and Valenciennes. 1842. by subsequent designation of Bleeker, 1863/i: 
204), 

riiih-iihiluhlln-. Bleeker. 1860i;:283. 412. 416 (proposed as subgenus of Liicio- 
si'njii: t\pe species Lt'iui^cii^ rniicma Bleeker, 1852, b> monotypy). 

DiAGNosis. — An cntomophagous bariliin genus with body 
elongate, subcylindrical or moderately compressed; head point- 
ed; snout elongate, with two (sometimes three?) obliquely ori- 
ented. S-curved shallow grooves (open lateral line canals?) be- 
tween anterior nostril and middle of snout; mouth obliquely 
ascending, with a long, wide gape, extending posteriorly well 
beyond anterior margin of eye; lower jaw with a moderately 
large symphyseal knob fitting into concavity behind tip of upper 
jaw; upper and lower lips with transverse ridges for their entire 
length, ridges separated by deep but narrow grooves, with oral 
ends more or less free (lobate); a single pair of rostral barbels 



and of maxillary barbels (absent or weakly developed in one 
species); gill rakers short, widely set. 1-3 + 9-11 = 11-13; pha- 
r>'ngeal teeth uncinate, in three rows. 4-5.4.2/2,4,4-5; lateral 
line complete; scales in lateral series 36-47. predorsal 20-24, 
transdorsal 1 1, circumpeduncular 12-14; dorsal-fin rays ii7-l/ 
2, anal iii6-l/2, pectoral i 14-1 6, pelvic i8; side of body with a 
longitudinal row of more or less evenly spaced roundish or 
vertically elongate oval spots; middle of dorsal and anal fin 
sometimes with a thin dark horizontal mark; both caudal-fin 
lobes with dark submarginal (never diagonal) band (sometimes 
absent in two species); vertebrae 22-25+ 16-18=39-43. 

For osteological characters see Howes (1980). Relationships 
of Liiciosonia are discussed below under L. spiloplcura. 

Liiciosoma is widely distributed in Thailand (Mekong, Chao 
Phrya). Malay Peninsula, Sumatra. Borneo, and Java, There are 
five species, three in western Borneo. 



Key to Luciosoma of western Borneo 

Head moderately pointed; numerous close-set tubercles 
in a broad semi-circle on snout-tip and on entire ventro- 
lateral surface of mandible; pelvic and anal fins without 
filamentous extension (pelvic fin sometimes with a very 
short prolongation of simple ray); body with a row of 
vertically elongate roundish or oval spots extending to 
base of caudal fin and continued as solid horizontal stripe 
on middle rays of caudal fin; upper and lower lobes of 



ROBERTS-WESTERN BORNEO FISHES 



43 




Figure 25. Luciosoma- Above. L aeligenini. Kapuas 1976-24. 102 mm (MZB 3152), below. L spiloplcura. Kapuas 1976-31, 45,5 mm (USNM 2301)531. 



2b 



caudal fin with submarginal stripe originating on dorsal 
and ventral margin of caudal peduncle L. spilopleura 
Head strongly pointed; tubercles entirely absent; pelvic 
and sometimes anal fin with long filamentous extension; 
body with row of roundish spots extending to caudal pe- 
duncle and continued as solid bar on caudal peduncle and 
submarginal stripe on upper caudal fin lobe; lower caudal 
fin lobe with submarginal band originating on lower half 
of caudal fin base; middle caudal fin rays colorless 2 

Rostral and maxillary barbels well developed, length to 
twice as much as eye diamater L. seligenim 
Barbels absent or vestigial in adults, sometimes present 
but very small in young L. trinema 



Luciosoma setigerum Valenciennes, 1842 

(Figure 25 above) 

Bariiioiff/gerHi Valenciennes //I Cuvier and Valenciennes. 1842:203 (type locality 

"riviere Pebak 4 Java"). 
Barbus podonemus Sleeker. 1850:18 (type locality "Surabaya, in tlumine Kali- 

mas"). See Bleekcr (1855/1:264). 
Luciosoma seligenim Sleeker. l855/i:264, 
Luciosoma Weheni Popta. 1905:177 (type locality "le Boelit, affluent du Bon- 

gam'"=Mahakam basin). See Weber and de Beaufort (1916:88). 
Luciosoma trinema Herre and Myers, 1937:58 (Pcrak. River Plus: misidentili- 

cation) 

Material Examined. — Malay Peninsula: Perak, Plus R., 3: 140-1 70 mm (CAS- 
SU 31 168-69: previously identified as L innema). Negri Sembilan. Kuala Pilah. 
119 mm (CAS-SU 31167). Western Sorneo: Kapuas 1976-17, 6: 31.9-119 mm 
(CAS 49220. MNHN 1982-662, MZB 3150, USNM 230182); Kapuas 1976-20, 
157 mm (MZB 3151): Kapuas 1976-24, 102 mm (MZB 3152): Kapuas 1976-30, 
121 mm (MZB 3153). 

See remarks under L. trinema. 

Distribution, — Malay Peninsula (Perak. Pahang. Malacca, 
Negri Sembilan), Sumatra (Solok, Sidjungdjung, Lahat, Batang 



Hari, Musi R,). Borneo (Kapuas), Java (Batavia, Bekassi, Ran- 
kasbetong, Lebak, Kediri, Surabaya, Gempol), 



Luciosoma spilopleura Bleeker, 1855 

(Figure 25 below) 

Luciosoma spilopleura Sleeker. 1855/i:265 (t\pe locality "Lahat. in nu\iis" = Musi 
basin. Palembang). 

Materials Examined —Sumatra: Lahat. 81.9 mm (BMNH 1866.5.2.97. ho- 
lotype). Western Borneo: Kapuas 1976-14. 6: 28.6-19.8 mm (BMNH 1982.3.29.13, 
CAS49221, MZB 3154. RMNH 28829); Kapuas 1976-22. 47.7 mm (MZB 3155); 
Kapuas 1976-31.4: 37.6-45.5 mm (MZS 31 56. flMMZ 209879, USNM 230183); 
Kapuas 1976-50. 67.8 (MZB 3157). 

Diagnosis. — Evidently a relatively small species, largest spec- 
imen 8 1 ,9 mm (all other Luciosoma attaining 1 35 mm or much 
more); neither pelvic nor anal fin with long filamentous exten- 
sions, pelvic fin sometimes with short prolongation of outermost 
ray (pelvic and sometimes anal fin with long filaments in all 
other Luciosoma): snout moderately pointed (strongly pointed 
in all other Luciosoma); numerous moderately large, close-set 
tubercles present in semicircle on snoul-tip between nostrils, tip 
of chin, and ventrolateral surface of mandibles (tubercles absent 
in all other Luciosoma except L. pellegnnii Popta, 1905, in 
which their distribution is like that in L. spilopleura); body with 
continuous lateral row of 16-18 vertically elongate oval or 
roundish spots continued to caudal-fin base; caudal fin with 
horizontal submarginal bars on each lobe and middle rays with 
a dark stripe continuous with body spots; vertebrae 39-41, 

The following observations are from the 67,8 mm Kapuas 
specimen: head 3,7; snout 11.1; eye diameter 16,1; rostral barbel 
almost twice as long as eye diameter, length 8.9; maxillary barbel 
about half as long, 15.8; gill rakers 2+10=12; scales in lateral 
series about 39, predorsal 23, transdorsal 1 1, circumpeduncular 
13; dorsal-fin rays ii7-l/2. anal iii6-l/2. pectoral ii4. pelvic i8; 



44 



CALIFORNIA ACADEMY OF SCIENCES 



dorsal-fin origin much nearer hypural fan than snout-tip, pre- 
dorsal length 1.6; pectoral fin reaching to or slightly beyond 
pelvic-fin origin, length 4. 1 ; pelvic-fin origin somewhat anterior 
to vertical through dorsal-fin origin, prepelvic length 3.7; pelvic- 
fin length 5.7; anal-fin origin below middle of dorsal-fin base. 

Luciosoma spilopleura is known only from Bleeker's holotype 
from Lahat, Sumatra and 1 2 specimens obtained by the Kapuas 
survey of 1976. It difl^ers m several respects from other species 
of Luciosoma and may be the most generalized or primitive 
member of its genus. Further study of the species might well 
contribute to a reassessment of the relationships of Luciosoma. 

Relationships of Luciosoma. —In my view Luciosoma may 
be more closely related to Barilius (scnsu lalo) than to any other 
Cyprinidae. This is contrary to the opinion of Howes, who ex- 
cludes Luciosoma from his "bariliine assemblage" and places 
it in a "luciosomine assemblage" with Parluciosoma (Rasbora. 
partim) and Mcgarasbora. I think Luciosoma is not so closely 
related to Rasbora. Readers are referred to the discussion of 
osteological characters by Howes (1980. especially pp. 180- 
184), particularly those concerning the ethmoid region. 1 have 
not studied these fishes osteologically and have not prepared a 
critical discussion of this aspect. It should be noted, however, 
that the osteological observations available are based on rela- 
tively few species, and that osteological examination of L. spi- 
lopleura could be particularly rewarding. Two general consid- 
erations bear noting: the elongate jaws of all Luciosoma and 
well developed rostral and maxillary barbels of Luciosoma ex- 
cept L. trincma strongly resemble the more or less elongate jaws 
of all Barilius and well developed rostral and maxillary barbels 
of many Barilius. In contrast, all Rasbora have relatively short 
jaws and lack both pairs of barbels (maxillary barbels weakly 
developed in .Mcgarasbora). Three specific features of L. spi- 
lopleura indicative of relationships to Barilius rather than to 
Rasbora are as follows: 1) Bariliins as a group, Barilius partic- 
ularly, are tuberculate, many species having tubercles on snout 
and mandible more or less comparable to those of L. spilopleura 
and L. pellegrinii. Tubercles are absent or but weakly developed 
on the snout and mandible oC Rasbora (including Parluciosoma) 
and .Mcgarasbora (mandibular tubercles of " Rasbora" a.xelrodi 
are highly specialized and may indicate this species is misiden- 
tified generically). 2) Coloration of Luciosoma and especially L. 
spilopleura agrees in general and in detail with coloration found 
in Bariliinae including Barilius (sensu lato) but not in Rasbora 
or any other Cyprinidae. I refer to the continuous lateral row 
of spots on the body, a primitive color feature of most African 
and Asian species of Barilius and of all Luciosoma. In most 
Luciosoma the spots tend to be round, but in L. spilopleura. as 
in Barilius usually, they are vertically elongate. Submarginal 
bands on caudal fin. present in three species of Luciosoma and 
several Banlius. are found in relatively few cyprinids and are 
absent in all species of Rasbora and Mcgarasbora. (Several Ras- 
bora have a diagonal mark across distal portion of each caudal- 
fin lobe.) 3) The shape of the compressed and moderately point- 
ed head of L. spilopleura is very much like that in various 
Barilius and less like the shape observed in nearly all Rasbora 
(this distinction is admittedly somewhat subjective). 

Distribution. — Sumatra (Lahat. upper Musi basin). Western 
Borneo (Kapuas). Records of L. spilopleura from north Borneo 
by Weber and de Beaufort ( 1916:89) presumably are all based 
on Luciosoma pellegrinii Popta, 1905, a species found in north. 



northeast and west Borneo (including Mahakam basin); see In- 
gerand Chin (1962:62). 

In the Kapuas basin L. spilopleura was collected only in high 
gradient rivers with rocky, stony or gravel bottoms, whereas L. 
setigerum and L. irinema were found in the Kapuas mainstream 
and low gradient tributaries with logs and leaf-litter on the bot- 
tom. 

Luciosoma trinema (Bleeker, 1852) 

Loicisciis inneinu Bleeker, I852(/:tiOO (lype locality "Palcmbang, in fluviis"). 
Luciosoma {.Tnnemalichlhys) Innema Bleeker, 1860a:4I3, 416. 

Material Examined. — Sarawak: 2: 122-135 mm (CAS-SU 31170); western 
Borneo: Kapuas 1976-16,68.5 mm(MZB 3158); Kapuas 1976-20,2: I41-I52mm 
(CAS 49222. MZB 31 59). Kapuas 1976-33. 84.6 mm (MZB 3160); Kapuas 1976- 
49, 63.6 mm (MZB 3161). 

Remarks. — L. Irinema and L. setigerum are very closely re- 
lated. They share a unique feature of coloration, i.e., union of 
spots on side of body with solid submarginal band on upper 
caudal fin lobe. Apart from presence or absence of barbels (which 
may be present but reduced in size in juvenile L. trinema), there 
seems to be almost no way to tell them apart. Reported dis- 
tinctions between L. setigerum and L. trinema (other than bar- 
bels) do not hold up under closer examination. For example, 
Howes (1980:171) reported "modal number of vertebrae" in L. 
setigerum as 43 (4-1-21 -H 17-1- 1) and in L. trinema as 39 
(4+184-16+1). This suggests a clear difference between the 
species. But Howes did not indicate number of specimens ex- 
amined in determining his modes, and they may not have been 
numerous. L. setigerum have vertebrae 23-25+17-18=41(2), 
42(2). 43(2), and L. trmema 23-25+ 16-18=40(6), 41(1). 43(2). 
Weber and de Beaufort (1916:87-88) reported scales in trans- 
verse series 3-1/2/1/4 in L. trincma and 5-1/2-6/1/4 in L. se- 
tigerum: I find 5- 1 /2/ 1 /2- 1 /2 in both species. Perhaps L. trinema 
and L. setigerum are morphological varieties of one species; the 
problem merits further study. 

Distribution. — Sumatra (Palembang, Kwantan R., Upper 
Langkat, Batang Hari). Borneo (Kapuas, Baram). Presence of 
L. trinema in Malay Peninsula is unconfirmed; a record from 
the River Plus, Perak (Herre and Myers 1937:58) is based on 
L. setigerum. 

Macrochirichthys Bleeker, 1859 

Macrochinclithys Bleeker. 1 859i;: 1 55 (type species Lcuiiscin iininoscopiis Bleeker. 
1851 =Leiicisciis macrocluni.'; Valenciennes in Cuvier and Valenciennes. 1 844. 
b\ monotypy; second species doublfulh included). 

Body extremely elongate, highly compressed. Barbels absent. 
No hyaline eyelid. Jaws strongly upturned, nearly vertical. Man- 
dibular symphysis strongly uncinate. Gill rakers rudimentary, 
pharyngeal teeth in two rows, usually 4,4 '4,4. Pectoral fin greatly 
enlarged, falcate, with 17 rays. Pelvic fin small, with 8 rays. 
Dorsal fin with 8 rays, anal with 25-28, both set far back. Lateral 
line complete, slightly curved downwards; scales in lateral line 
series 120-130. 

I tentatively regard this genus as comprising a single species. 

Macrochirichthys macrochirus (Valenciennes, 1844) 

(Figure 26) 

I.ciillscus macroclnrifi Valenciennes m CuMer and Valenciennes. 1844:348 (type 
locality Java). 



ROBERTS-WESTERN BORNEO FISHES 



45 




Figure 26. Macrochiruhlhys macrochirus- Kapuas 1976-20, 212 mm (CAS 49223). 



Lcuasciis uranoscoptis Bleeker, 1851i;;.^14 (type locality "Banjermassmg, m flu- 

viis"). 
Maciochirichlhys uranoscopus Bleeker, 1860a:476. 
.\tacrnchinchthysT?='Leiicisciis inaimchinis Bleeker, 1860u;477. 
Mi2crochirichihys snyden Fowler, 1905:487, fig. 7 (type locality Baram). 
MacriKhinchlhys laosensis Fowler, I934/i:l 12 (type locality Mekong at Chieng- 

sen). 

Material Examined. — Western Borneo: Kapuas 1976-20, 10: 134-221 mm 
(BMNH 1978,1.10:1^, CAS 49223, MZB3162, USNM 230184): Kapuas 1976- 
44, 166 mm (MZB 3163); Kapuas 1976-49,2: I 32-168 mm(MZB 3164, UMMZ 
209925). 

Diagnosis.— See generic diagnosis above, 

Weber and de Beaufort (1916:54) noted a total length of over 
500 mm for this species. One I observed in a fisherman's catch 
in 1976 was about 1 m total length. 

Distribution, — Laos, Cambodia, Vietnam (Mekong), Thai- 
land (Mekong, Chao Phrya, Tapi. Tale Sap, Tale Noi). Sumatra 
(Palembang, Batang Hari, Danau Sialong lotong), Borneo (Ka- 
puas, Kahajan, Pengaron, Barito. Mahakam, Baram), Java, 



Morulius Hamilton-Buchanan. 1822 

Monihus Hamilton-Buchanan. 1822:331, 391 (type species Cyprimis (Morulius) 
inomla Hamilton-Buchanan, 1822, by subsequent designation of Bleeker, 1863/i: 
195), 



Morulius chrysophekadion (Bleeker, 1850) 

Rohila chrysophekudu^n Bleeker, 1850:20 (type locality Surabaya). 
Rohiia cyanomelas Bleeker, 1852(/:597 (type locality Palembang). 
Rohila polyporos Bleeker, 1853/:519 (type locality Moarah Kompeh, Sumatrae 

onentahs; Batavia). 
Rohila koilogeiwion Bleeker, 1857/i:359 (type locality Batavia), 
Chrysophekadion polyporos Bleeker, 1859<;:145. 
Morulius chrysophekadion Bleeker, 1860i/:188. 
Laheo chrysophekadion Giinther, 1868(v:52, 

Material Examined. — Western Borneo: Kapuas 1976-19, 2: 170-319 mm (C.\S 
49224, MZB 3165). 



Mystacoleucus Giinther, 1868 

Myslacoleucus Giinther. 1868:206 (type species Capocia padangensis Bleeker, 

1852, by original designation and monotypy). 
.•lfan(/!o«ori«Tickell/HDay, 1888:807 (type species .-Icanz/wnofui argpnr««Tick- 



cll in Day. 1888. by monotypy). Preoccupied by Acanihonotiis Bloch, 1795; 
Acanlhonolus Gray. 1831. 
.\falsva Day, 1 889:292 (replacement name for . Icaiiihonoius Tickell in Day, 1888), 

Mystacoleucus marginatus (Valenciennes, 1842) 

(Figure 27) 

Barbus ohlusiroslris van Hasselt, 1823:132 (nomen nudum). 

Barhiis marginatus Valenciennes in Cuvier and Valenciennes, 1842:164 (type 

locality "'riviere de Tjicanigui," Java), 
Barhus ohlusiroslris Valenciennes in Cuvicr and Valenciennes, 1842:167 (type 

locality Buiten/org). 
Myslacoleucus niarginaliis Smith, 1931:185. 

Material Examined. — Malay Peninsula: Tapah, Perak, 8: 66.6-94.1 mm (CAS- 
SU 39356). Western Borneo: Kapuas 1976-24, 29: 39.0-79.4 mm (AMNH 48922, 
BMNH 1 982. 3. 29. 14-1 6, CAS 49225, FMNH 9420 LlRSNBl 973 l.MCZ 58341, 
MZB 3166); Kapuas 1976-27, 35: 45.7-80.8 mm (MNHN 1982-663, MZB 3167, 
NlFl uncat., RMNH 28830, ROM 38614, UMMZ 209869, USNM 230185, ZMA 
116-519). Java; Buiten/org, 3: 77.1-98.4 mm (CAS-SU 20485). 

Osteochilus Giinther, 1868 

Deplochcilos van Hasselt, 1823 (nomen nudum). 

Diplocheilichlhys Bleeker, 1 860ii: 1 42 (type species Lolmcheilospleuroiaeiua Bleek- 
er. 1855, by monotypy; nomen oblilum), 

Diplochcilos Bleeker, 1 860u: 1 39 (as subgenus of Laheo: type species Laheo ro- 
hiloidcs Bleeker, \S51=Lohnchilos piciiroiaeniu Bleeker, 1855, by monotypy; 
nomen oblitum), 

(9s/['nW»/!/s Giinther, 1868:40 (type species Rohila mclanopleura Bleeker, 1852, 
by subsequent designation of Jordan, 1919351). 

Lips large, everted, sucker-like, covered with more or less 
elongate folds or plicae bearing numerous unicellular keratinous 
projectionsorunculi (Roberts 1982a), Pharyngeal teeth in three 
rows 2,4.5/5.4.2. Scales 27-53. 

The genus includes some 23 species in Burma, Thailand, 
southern China, the Malay Peninsula. Sumatra. Borneo, and 
Java, Many are important food fishes, Borneo is inhabited by 
20 species, 13 of which have been found in the Kapuas. One 
species, O. kappeni. is known only from the Kapuas, The ex- 
tensive material of Osteochilus obtained from the Kapuas in 
1976 has been identified by me and by Jaranthada Kamasuta, 
who studied most of the specimens in connection with his doc- 
toral dissertation on syslematics of Oi/coc/H/iw (Kamasuta 1 98 1 ). 
Much of the information presented here is based on Kamasuta 
(1981). 



46 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 27. Atyslacolcticus Diar^inatm Kapuas l'^76-27, 70.5 mm (MZB 3167) 



Ostcoiiiiliis may be divided into three groups based on length 
and shape of the longest folds or plicae on the greatly expanded 
lateral portion of the upper lip. The groups are also partly dis- 
tinguished by the distribution of unicellular keratinous projec- 
tions or unculi and taste buds on the plicae, and by the size and 
shape of the unculi. The groups seem to represent species adapt- 
ed to similar feeding ecology or habitat rather than species which 
are phyletically related. 

The three groups may be outlined as follows: 1 ) Endomorphs: 
Main plicae on lateral portions of upper lip divided into more 
or less numerous oval or mound-shaped sections. Unculi and 
taste buds tend to be mixed together. Unculi low-lying, polyg- 
onal, to 4-1 8 iim long, usually present in furrows between plicae 
as well as on plicae. This includes the largest and most deep- 
bodied members of the genus (endomorphs). Generally inhabit 
lakes and large rivers with slow current. Absent from mountain 
streams. Osteochihts horneensis. O. hasseltt, O. mclanopleura. 
O schle^cli. Maximum standard length 224-366 mm. This group 
includes some of the most important freshwater food hshes of 
southeast Asia. Ostcochilus melanopleura is especially note- 
worthy for its upturned mouth; in all other Osteochiliis the mouth 
is subinferior or inferior. It is also the largest species. 

2) Mcsomorphs: Main plicae on lateral portions of upper lips 
all divided into at least two or three elongate, ridge-like seg- 
ments. Taste buds generally distributed along anterior margin 
of unculiferous apex of plicae. Unculi intermediate in size, fre- 
quently expanded distally, to 16-24 ^m long, generally weakly 
developed or absent from furrows. More or less intermediate 
in size and body depth. Maximum standard length 120-205 
mm. Mainly inhabit lowland rivers of moderate gradient. Os- 
tcochilus kappcni. O. tnponis. and O. waaiuicrsi. 

3) Ectomorphs: Main plicae on lateral portions of upper lip 
forming very long, high, unbroken ridges. Unculi confined to 
summit of ridges, taste buds generally form well defined rows 
parallel to unculiferous portions of ridges. Unculi relatively large, 
to 20-27 ^m long, elongate. Maximum standard length 75-225 
mm. Tend to inhabit smaller forest streams and mountain 
streams with relatively strong current. Ostcochilus cnncaponis. 
iutciiuedius, kahcyancnsis. microccphalus, plcurotaenia and spi- 
liinis. In this group O. plcwfltacuia and O. spiliirus are partic- 
ularly noteworthy. Oslcnchilns plcuroiucnia apparently has the 



largest lips of any Ostcochilus and its mouth is strongly inferior. 
The habitat is not well known, but from the localities recorded 
I suspect It inhabits large mountain streams and probably prefers 
swift currents. Ostcochilus spilurus. attaining a maximum stan- 
dard length of only about 75 mm, is by far the smallest species 
of Ostcochilus. Some populations apparently mature at much 
smaller sizes, and the largest adults in such populations may 
attain only about half this length. 

The above divisions can be extended to include the rest of 
the species oi Ostcochilus which are not found in western Bor- 
neo. The quality of the plicae is readily observable with a dis- 
secting microscope or in most instances with the unaided eye. 
The unculi, however, are best observed by scanning electron 
microscopy, and thus far such observations have been made 
only for O. horneensis. enneaporus. hasseltt. intennedius. 
melanopleura. microccphalus. sarawakensis. irtporus. and 
waandersi (K.2irr\&suXz 1981, and herein). 

The different types of lips in Ostcochilus cannot yet be cor- 
related with difltrences in feeding behavior or food. Ostcochilus. 
like other Labeoinae, probably feed on aufwuchs. Species of 
forest streams presumably feed on aufwuchs growing on sub- 
merged tree trunks or logs and roots, those of mountain streams 
on epilithic aufwuchs. Scanning electron micrographs of the lips 
of a 58.5 mm O. hasseltt (Fig. 30) reveal numerous straight 
fragments of about 4-28 i^m which may possibly be chopped 
up bits of filamentous algae. These pieces appear to be cut off 
at right angles at both ends. Examination of the stomach con- 
tents of the same specimen with a compound light microscope 
reveals large numbers of fragments from 1.0 ;um to 1.5 mm, 
many of which are segments of filamentous algae with 2-1 1 
cells. Even more abundant are irregular masses of apparently 
organic material, at least some of it possibly from higher plants. 
More information could certainly be gleaned from examining 
stomach contents of more specimens, but field and laboratory 
study of live fish would be much more profitable. 



Key to Ostcochilus from western Borneo 

la Lateral line scales 45-53; dorsal-fin rays 20-23 2 

lb Lateral line scales 27-35; dorsal-fin rays 14-21 3 



ROBERTS-WESTERN BORNEO FISHES 



47 




Figure 28. Osleochilus borneemis. Kapuas 1976-16. 84.9 mm (CAS 49226). 



2a Mouth upturned; snout non-tuberculate; a large dark 
vertical mark on body above pectoral fin; gill rakers 27- 
35; circumferential scales 22-23/2/23-24 

O. melanopleura 

2b Mouth inferior; snout with three large tubercles; no mark 
on body above pectoral fin; gill rakers 20-25; circum- 
ferential scales 17-19/12/17-20 0. bomeensis 

3a Circumpeduncular scales 20 O. schlegeli 

3b Circumpeduncular scales 12-16 — 4 

4a Large, round, peduncular spot 5 

4b No large, round peduncular spot _ 8 

5a Lateral line scales 27-29; circumferential scales 9/2/11 

O. spihtrus 

5b Lateral line scales 29-35; circumferential scales more 

than 9/2/11 6 

6a Two large tubercles on snout; lateral plicae of upper lip 

long and undivided O. kahajanensis 

6b No tubercles on snout; lateral plicae of upper lip short, 

divided in two or three 7 

7a Circumferential scales 13/2/15; spots on sides weak or 

indistinct O. kappeni 

7b Circumferential scales 11/2/13; spots on sides darker 

and more distinct O. hasselti 

8a Midlateral stnpe 9 

8b No midlateral stripe 1 2 

9a Midlateral stripe continued on middle caudal-fin rays to 

end of caudal fin O. waandersii 

9b Midlateral stnpe stopping more or less abruptly at base 

of caudal fin 10 

10a Gill rakers 27-35 „ _._.._ O. microcephalus 

10b Gill rakers 40-60 11 

1 la Snout non-tuberculate or with numerous fine tubercles; 
sides of body with little or no pigmentation except me- 
dian longitudinal stripe O. pleuwtaenia 

1 lb Snout with 1 or 3 tubercles; sides of body heavily pig- 
mented, so that median longitudinal stripe is obscured 
(particularly anteriorly) O. enueaponis 

12a No rows of spots on scales; no dark mark on base of 
anterior portion of dorsal fin O. inlennedms 



13a 



13b 



Body with several longitudinal rows of spots on exposed 
portion of scales; a large dark mark on base of anterior 

portion of dorsal fin 13 

Plicae on lateral portion of upper lip undivided; no tu- 
bercles on snout; upper and lower lobes of caudal fin 

with thin dark margins O. iiUermediiis 

Plicae on lateral portion of upper lip divided into two 
or three; snout with three tubercles; caudal fin with dark- 
ened margins O. triporus 



Osteochilus borneensis (Bleeker, 1857) 

(Figure 28) 

Rohiia borneemis Bleeker. 1857(i:17 (type locality Kapuas River al Pontianak). 
Osleochilus borneensis Giinther, 1 868£j:4 1 . 

Material Examined. — Western Borneo: Kapuas 1976-16. 2: 66.3-84.9 mm 
(CAS 49226. MZB 3168); Kapuas 1976-17, 2: 57.2-82.1 mm (MZB 3169. NIFI 
uncat); Kapuas 1976-33. 70.4 mm (MZB 3170). 

Endomorph. Body depth 3.3-3.8. Dorsal-fin rays 20-23, pec- 
toral 15-16. Scales in lateral series 41-49, circumferential 17- 
19/2/17-20, circumpeduncular 22-24. Gill rakers 28-31. Mouth 
subinferior. Lateral plicae of upper lips irregularly arranged, 
short, mound-shaped. A large, dark round circumpeduncular 
spot. Body with about 15-17 faint longitudinal stripes. Fins 
dusky or plain. Maximum standard length 282 mm. 

Distribution. — Sumatra (Djambi). Borneo (Kapuas basin 
only). 

Osteochilus enneaporus (Bleeker, 1852) 

(Figure 29) 

Rohila enneaporos Bleeker. 1852(/:596 (type locality Padang, Sumatra). 

Osleochilus v;rMf;« Giinther. 1868(;:44 (in pan). 

Osleochilus ritlalouies Popta, 1904:195 (type locality Howong River. Mahakam 

basin, eastern Borneo). See Kamasuta (1981). 
Osteochilus scapulans Fowler. 1939:69. fig, 17. IS (type locality Trang, southern 

Thailand). .See Karnasula (1981). 

MaterialExamined. — Western Borneo: Kapuas 1976-6, 7: 62.0-1 38 mm (CAS 
49227, MZB 3171, NIFI uncat., USNM 230186); Kapuas 1976-24, 86.3 mm 
(MZB 3172): Kapuas 1976-25. 2: 50.0-61.8 mm (BMNH 1982.3.29.17. MZB 
3173): Kapuas 1976-27, 6: 52.9-105 mm (FMNH 94202, MNHN 1 982-664, MZB 



48 



CALIFORNIA ACADEMY OF SCIENCES 







FfGURE 2y. Osicochilm cnncaponis Above. Kapuas 1976-6. 121 mm (CAS 49227). below, unculiferous lips, Kapuas 1976-6, 82.7 mm (CAS 49227). 




.M74. RMNH 288.'(I. ROM 38616); Rapuas 1976-29. 4: 72.3-136 mm (MZB 
3175, LIMMZ 209874. USNM 230187). 

Eclomorph. Body depth 2.9-3.4. Dorsal-fin rays 15-18. pec- 
toral 15-16. Scales in lateral series 31-32, circumferential 11/ 
2/13, circumpcduncular 16. Gill rakers 40-60. Snout tubercles 
1 or 3 (if 3, median tubercle largest). Median lateral stripe on 
body, usually distinct on posterior half of body, obscured by 
dark melanin. Lateral surface of upper lip with long undivided 
plicae. Maximum standard length 192 mm. 

Osteochilus hasseiti (Valenciennes, 1842) 

(Figure 30) 

Ri^hiUi Ihiwflli Valenciennes in C~UMer and Valeneiennes, 1842:209 (t\pe locality 
.laval. 

Rnhila rnsnilalia Valenciennes in Cuv icr and Valenciennes, 1842:250 (type lo- 
cality Rangoon, Burma). 

Riihilti {Riihila) kiilili Bleeker, 1860t;:l77 (i\pe localit\ Palenihang. Sumatra), 

O'.icoc lulus Ihissclu Giinther. lS68i;:41. 

OsIcikIiiIus kiihlii GUnlher. 1868i;:43. 

Osleochilus ncilli Day. 1870:99 (type locality Sittang River. Burma) 

nsicihiiiliis (lui>siif;iiM Fowler. 1937:182, fig. 121 (type locality kemaiat, north- 
eastern Thailand). 

(Hhwhilus hassclti tnvniici Menon. 1954 12. hg 3 (type locality Kuala Tahan, 
Pahang. Malay Peninsula). 

Material Examined.— Western Borneo: Kapuas 1976-6. 5: 43.5-96,0 mm (CAS 
49228. MZB 3176): Kapuas 1976-7, 43.5 mm (MZB 3177): Kapuas 1976-8. 9: 
56 4-95,2 mm (MNHN 1982-665. MZB 31 78. NIF! uncal., RMNH 28832, USNM 
230188): Kapuas 1976-13, 96 3 mm (MZB 3179) 

Endomorph. Body deep, 3.3-4.1. Dorsal-fin rays 16-21. pec- 



toral 14-16. Scale in lateral series 30-33, circumferential 1 1/2/ 
13. circumpcduncular 16. Gill rakers 27-35. No tubercles on 
snout. Lateral plicae of upper lip short, moundlike. Sides of 
body with about 9 longitudinal rows of close-set round spots. 
.\ large circumpcduncular spot. Fins plain. Maximum standard 
length 224 mm. 

Osteochilus intermedius Weber and de Beaufort. 1916 

(Figure 31) 

Oslcoihiliis inlcrmcdiin Weber and dc Beaufort. 1916:133-134 (type locality 
Djambi, Sumatra, and Kapuas River at Putus Sibau). 

Material Examined, — Western Borneo: Kapuas 1976-17. 2: 66.4-69.8 mm 
(CAS 44182. MZB 3180): Kapuas 1976-42. 4: 56.4-64,2 mm (MZB 3181. NIFI 
uncat.. RMNH 28833, USNM 230129): Kapuas 1976-43, 3: 46.7-49,9 mm (MZB 
3 1 82. CAS 49229); Kapuas 1976-47. 46.4 mm (MZB 3183), 

Weber and de Beaufort (1916:133) gave only a summary de- 
scription of this species and expressed doubt as to its distinct- 
ness. Hardenberg (1936) suggested it is only a variety of O. 
triporus. Karnasuta and 1 find that it is a distinct species, dif- 
ferent in many respects from O. triporus. which appears to be 
its closest relative. 

Ectomorph. Body slender, compressed. Dorsal-fin rays 17- 
1 8. pectoral 1 4-15. Scales in lateral series 30-32. circumferential 
9/2/13, circumpcduncular 16, Gill rakers 39-49, 

No tubercles on snout. Plicae on ventral portion of upper lip 
undivided. Dorsal fin with a large but indistinct dark spot an- 
teriorly; otherwise dusky. Dorsal-fin margin nearly straight. Pe- 



ROBERTS- WESTERN BORNEO FISHES 



49 




Figure 30. Oslcochihis hasschi- Above. Kapuas 1976-8. 95.2 mm (USNM 230188); below, unculifcrous lips. Kapuas 1976-6. 58.3 mm (CAS 49228). 



duncular spot absent. Upper and lower lobes of caudal fin with 
a diffuse, thin, longitudinal band extending from procurrent rays 
to end of uppermost and lowermost caudal fin rays (absent in 
O. triporus). Side of body with six rows of dark spots, one on 
lateral line scale row, two on the two scale rows above lateral 
line and three on the three scale rows below lateral line; row of 
spots on lateral line scales noticeably fainter than other rows 
(rows of spots on lateral line equally distinct in O. triporus). 
Maximum standard length 154 mm. 

Distribution. — Sumatra (Djambi; Batang Hari). Borneo 
(Kapuas basin only). 

Osteochilus kahajanensis (Bleeker, 1857) 

(Figure 32) 

Rolnia kalmjaiiemis Bleeker. 1857i;:18 (type locality Kahajan River). 
Osteochilus kahajanensis Giinther, 1868a:44. 

Material Examumed.— Western Borneo: Kapuas 1976-17. 78.3 mm(MZB 3184); 
Kapuas 1976-39. 62.8 mm (MZB 3185). 

Endomorph. Body depth 3.2-3.9. Dorsal-fin rays 1 9-2 1 . Scales 
in lateral series 31-32, circumferential 11/2/13, circumpedun- 
cular 16. Gill rakers 33-46. Differs from all other Osteochilus 
in having two large tubercles on snout (instead of 0,1,3, or 
numerous small tubercles). A diffuse or poorly defined median 
longitudinal stripe. Ventral surface of upper lip with moderately 
long undivided plicae. Maximum standard length 220 mm. 



Osteochilus kappenii (Bleeker, 1857) 

(Figure 33) 

Ruhila kappcnii Bleeker. 1857a: 19 (type locality Kapuas River at Pantianak). 
Osteochilus kappciiti Gunlhcr. 1868a;42. 

Osteochilus hre\iciiuda Weber and de Beaufort, 1916:138 (type locality Kapuas 
River at Putus Sibau and Putus Genting). See Karnasuta (1981). 

Material Examined. — Western Borneo: Kapuas 1976-28, 3: 46.5-79.8 mm 
(CAS 49230. MZB 3186. NIFI uncat.): Kapuas 1976-37. 2: 65.2-65.9 mm (MZB 
3187. USNM 230190); Kapuas 1976-43. 50.5 mm (MZB 3188). 

Mesomorph. Body compressed, depth 3.5—4.5. Dorsal-fin rays 
19-21, pectoral 15. Lateral line scales 31-33, circumferential 
1 3/2/ 1 5, circumpeduncular 1 6. Gill rakers 26-34. Lateral plicae 
of upper lip short, each divided into two or three. About nine 
rows of indistinct spots on sides of body (spots darker and more 
distinct in O. hasseiti). A large circumpeduncular spot. Fins 
plain. Maximum standard length 175 mm. 

This rare species of restricted distribution is very similar to 
the common and very widely distributed O. hasseiti. They can 
be separated by the circumferential scale count (13/2/15 in O. 
kappeni vs. 11/2/13 in O. hasseiti). as well as the less distinct 
spots in O. kappeni. 

This species may be endemic to the Kapuas. Identification of 
specimens reported from Djambi, Sumatra, as O. kappeni (We- 
ber and de Beaufort, 1916:138) should be checked. A record 
from the Baram River (Fowler 1905) seems likely to be based 
on O. kahajanensis (Karnasuta 1981). 



50 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 31. OsIchcIiiIks inlcrnwciiiis Atiovc, Kapuas 1976-43. 49,9 mm (CAS 49:29); below, unculifcrous lips, Kapuas 1976-17, 69.8 mm (CAS 44182). 



Osteochilus melanopleurus (Bleeker. 1851 

(Figure 34) 



(CAS 49231. MZB 3189, USNM 230191): Kapuas 1976-49, 2: 72.8-119 mm 
(CAS 49232. MZB 3190), 



R..lula mclanopU-ura Bleeker, 1852/r430 (type luealny Bandjermassing. Borneo. Immediately distinguished tVom all Other OstCOchlltlS by itS 

and Paiemhang, Sumatra) upturned mouth, and by the large size of the vertical dark bar 

Osioichilus niclcvioplfiiniK GuMhcr. 1868(;:40, on body above pectoral fin. 

Material Examined.- Western Borneo: Kapuas 1976-2(1. 3: 83,3-137 mm Endomorph. Bodv depth 3.1-4.1. Dorsal-fin rayS 20-22, pec- 




FiGURE 32. Ch'leochilus kahajanensis. Kapuas 1976-39. 62.8 mm (MZB 3185). 



ROBERTS-WESTERN BORNEO FISHES 



51 




Figure 33. Osteochilus kappenii- Kapuas 1976-43, 50.5 mm (MZB 31 



toral 17-18. Lateral line scales 41-53, circumferential 22-23/ 
2/23-24, circumpeduncular 22-24. Gill rakers 25-40 or more 
(number increasmg with growth). Lateral plicae of upper lip 
irregularly arranged, mound-shaped. Fins and body otherwise 
plain. Maximum standard length 366 mm. 



Distribution. — Mekong basin (Laos, Kampuchea, southern 
Vietnam, and Thailand). Malay Peninsula (Chenderoh Dam, 
Perak; Kalantan). Sumatra (Djambi, Palembang, Mandan R. at 
Siak). Borneo (Kapuas, Baram). Thailand (Tapi R., Meklong 
R., Menam Mun, Chao Phrya, Meyom at Lampang). 




mC^^:^/ ,9-^ 





'""^^v^-*^ 



Figure 34. Osicochilus melanopleurus. Above. Kapuas 1976-49. 1 19 mm (MZB 3190); below, unculiferous lips, Kapuas 1976-49, 72.8 mm (CAS 49232 






52 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 35. Oslcivhiliis micnxephalus. Kapuas l'^76-45. 70,4 mm (MZB 3204). 



Osteochilus microcephalus (Valenciennes. 1842) 

(Figure 35) 

Ri'Jiiiu nucroccpliahis Valenciennes /// Cuvier and Valenciennes, IS42:275 (type 

locality Bantam River, Java). 
Rdhtla hmchynotopenis Bleeker, 1855/);266 (type locality Lahat, Sumatra). 
Riiluia vitlala Bleeker, 1860u:178 (type locality Java. Sumatra, and Borneo). 
(>\icihiulu\ muToccphaliis GiiMher 1868i/;43. 
Oslciichilns hrachvnoloplmis GUnther I868t(:43. 
OMciniiihc Mllalus Giinther I868i(:44. 

Material Examined. — Western Borneo: Rapuas I'?76-I5. 3: 36.0—44.1 mm 
(BMNH I')82.3.2').l8-iq. MZB 31'JI); Kapuas 1'^76-lb, 3: 47,0-53,5 mm (CAS 
41233. MZB 3192); Kapuas 1976-17, 2: 55,6-61,5 mm (FMNH 94203, MZB 
3193), Kapuas 1976-19, 73,1 mm (MZB 3194); Kapuas 1976-21, 3: 24,7-28,1 
mm (MZB 3195); Rapuas 1976-27. 2; 93,8-1 15 mm (IRSNB 19732, MZB 3196); 
Kapuas 1976-29, 3; 87,9-99,5 mm (MCZ 58342. MZB 3197); Kapuas 1976-31, 
5: 39,6-100 mm (MZB 3198. CAS 49234); Rapuas 1976-33. 61.0 mm (MZB 
3199); Kapuas 1976-35. 5: 53.8-77.6 mm (MZB 3200. MNHN 1982-666); Kapuas 
IQ76-37. 8; 44.4-60.3 mm (MZB 3201. RMNH 288.34. UMMZ 209893. USNM 
230192); Rapuas 1976-39, 59.1 mm. MZB 3202); Rapuas 1976-43. 7; 34.7^3.8 
mm (MZB 3203. USNM 230193, ZMA 116.250); Kapuas 1976-45. 70.4 mm 
(MZB 3204); Kapuas 1976-46, 43,3 mm (MZB 3205); Rapuas 1976-47. 34,8 mm 
(MZB 3206), 

Eclomorph. Body depth 2.8-3.7. Dorsal-fin rays 14-17, pec- 
toral 14-16. Lateral line scales 32-33, circumterential 1 1/2/13, 
circumpeduncular 16. Gill rakers 27-35. Snout tubercles 0, 1 
or 3. Midlaleral stripe from gill opening to end of caudal pe- 
duncle. Mouth subinferior. Ventral surface of upper lip with 
moderately long undivided plicae. Maximum standard length 
140 mm. 

Distribution. — Laos, Kampuchea, and probably southern 
Vietnam (Mekong basin). Thailand (Mekong, Chao Phrya, Kwai 
Yai R.. Pattani R. at Yala). Sarawak (Niah R., Baram R.). Bor- 
neo (Kapuas. Barito R). Malay Peninsula (Tahan R.. Pahang; 
Sadili R.. Johore). Sumatra (Djambi. Moesi, Singkarak, Sock- 
adana, Sindjungdjung. Palembang). 



Osteochilus pleurotaenia (Bleeker, 1855) new combination 

Liibivhilw, pU'iinnacnia Bleeker, 1855/);267 (type locality Lahat, Sumatra), 
LdluKhihii rohinvdes Bleeker. 1857/i;363 (type locality Rrawang, Batavia. Java), 
DipUichciluhihvs plcurolacnia Bleeker, 1860a;143 (description of Diplocheil- 
ii_hfhv\ with Li>/'<nliilit\ plcuroiacnta as type species). 



Label {Diploihcilii'.) roliilnulc^ Bleeker, 1860ij;139, 
Labeo pleiirolacnia GUnther. I868i(;58, 

Osleochilus icniinkii Popla. 1904;194 (type locality Bongon River=Mahakam 
Basin, eastern Borneo), 

Material Examined, — Western Borneo (Rapuas R. at PuUissihau). 3, and Nan- 
garaun, Mandai R,. Kapuas basin, 2 (RMNH 168.3-86) 

Ectomorph. Body slender, depth 2.5-3.0. Dorsal-fin rays 14- 
15, pectoral 16-17. Lateral line scales 30-31, circumferential 
11/1/13, circumpeduncular 16. Mouth strongly inferior, lips 
perhaps larger than in any other species oi Osteochilus. Ventral 
portion of upper lip with plicae long and undivided. Snout non- 
tuberculate or with numerous fine tubercles. Gill rakers 40-60. 
Juveniles with median longitudinal stripe from gill cover to base 
of caudal fin (disappears with growih). Ma.vimum standard length 
225 mm. 

This is the only species oWsteochihis known from the Kapuas 
basin not obtained during the Kapuas survey of 1976. J. Kar- 
nasuta and 1 independently examined material in Europe and 
came to the conclusion that it belongs in Osleochilus. The syn- 
onymy presented here is from Karnasuta (1981). 

Distribution. — Sumatra (Lahat, Moesi R.). Borneo (Kapuas, 
Mahakam). Java (Krawang. Batavia). 

Osteochilus schlegeli (Bleeker, 1851) 

(Figure 36) 

Rohila schlegeli Bleeker, 1851/;432 (type locality Bandjermassing). 
Osleochilus schlegeli Giinther, 1868u;42, 

Material Examined, — Western Borneo; Rapuas 197t)-19, 125 mm (MZB 3207); 
Rapuas 1976-44. 102 mm (MZB 3208), 

Endomorph. Body deep, 3.4-4.3. Dorsal-fin rays 16-18. Lat- 
eral line scales 32-33, circumferential 13/2/15, circumpedun- 
cular 20. Gill rakers 24-30. Lateral plicae of upper lip mound- 
shaped, irregularly positioned. Body and fins usually plain, 
except juveniles usually have a vertical black blotch on side of 
body above pectoral fin. and some specimens have a large black 
spot on anterior base of dorsal fin. Maximum standard length 
31 3 mm. 

Distribution. — Thailand (lower Chao Phrya, Meklong R.). 
Sumatra (Djambi. Batang Hari. Palembang). Borneo (Kapuas). 



ROBERTS- WESTERN BORNEO FISHES 



53 




Figure 36. Oslcochilus schlcgch. Rapuas 1976-19. 125 mm (CAS 49235). 



Osteochilus spilurus (Bleeker, 1851) 

(Figure 37) 

Dangila spilurus Bleeker, 185U;272 (type loealit\ Bandjcrmassmg, Borneo). 
Ritlula oligolepis Bkeker. 1853</:191 (type locality Marawang, Banka). 
O'iU'flchi/us oligolcpis Giinther, 1868u:45. 
Oslcochilus spilurus Giinther, 1868£7:45 



Material Examined, -Western Borneo: Kapuas 1976-5, 2: 18.8-39.2 mm (CAS 
49235, MZB 3209); Kapuas 1976-8, 2: 46.8-48,5 mm (CAS 49236, MZB 3210); 
Kapuas 1976-16.3: 13,0-580 mm (CAS 49237, MZB 3211 ); Kapuas 1976-17, 
2: 53,3-68,0 mm (CAS 49238, MZB 3212); Kapuas 1976-32, 6: 22,1-28,7 mm 
(MZB 3213, USNM 230194); Kapuas 1976-37. 39.9 mm (MZB 3214); Kapuas 
1976-42, 49: 24.8-42.7 mm ( AMNH 48923. BMNH 1982.3.29.20-23, CAS 495 10, 
IRSNB 19733, MCZ 58343, MNHN 1982-667, MZB 3215. NIFl uncat, RMNH 




Figure 37, Osteochilus spilurus- Above, Kapuas 1970-5, 39,2 mm (CAS 49235); below, uncuhfcrous lips, Kapuas 1976-42, 42,1 mm (CAS 49236). 



54 



CALIFORNIA ACADEMY OF SCIENCES 




XiTtnL 1^^ ^ .^_A*<K^ 



Figure 38. Osleochiliis triporos. Above. Kapuas l47b-49. 66.6 mm (MZB illl). below, uncuhlerous lips, Rapuas l'i76-42, 79.0 mm {USNM 230197). 



28835, ROM 3861 I): Kapuas 1976-46, 22: 25.2-49 I mm (MZB 3216, IJMMZ 
209920. USNM 230195). Kapuas 1976-47, 5: 25.4-29.7 mm (MZB 3217, USNM 
230196); Kapuas 1976-51. 7: 22,4-34.7 mm (CAS 49239, MZB 3218). 

Ectomorph. Body moderately deep, 3,1-3.5. Dorsal-fin rays 
14-15, pectoral 1 3-14. Lateral line scales 27-29, circumferential 
9/2/1 1, circumpeduncular 16. Gill rakers 25-28. A round dark 
spot on caudal peduncle. Ventral surface of lip with moderately 
long undivided plicae. Maximum standard length 75 mm. 

Distribution. — Southern Malay Peninsula. Southern Su- 
matra. Western and southern Borneo, .lava. Banka. Billiton. 

Osteochilus triporos (Bleeker, 1852) 

(Figure 38) 

Ruhita Inponii Blocker. 1852i';59S (type localiU I'alembang, Sumalra). 
f>sU'i kIiiIus inporiis Giinlher, 1868ij:44. 

M.ATERiAL Examined. — Western Borneo: Kapuas 1976-19, 6; 78.9-116 mm 
(BMNH 1982.3.29.24, CAS 49240, RMNH 94204, MZB 3219); Kapuas 1976- 
20. 4; 66,4-113 mm (MZB 3220, MNHN 1982-668, RMNH 28836); Kapuas 
1 976-42, 6: 7 1 .6-84.8 mm (MZB 322 I , UMMZ 209905. USNM 230197); Kapuas 
1976-49. 66.6 mm (MZB 3222) 

Mesomorph. Body elongate, slender, 2.6-3.5. Dorsal-fin rays 
15-16. Lateral line scales 29-30, circumferential 11/2/11-13, 
circumpeduncular 13. Ventral part of lip with long plicae di- 
vided into two or three parts. GUI rakers 27-32. Snout tubercles 
3, middle one largest. Long plicae on ventral portion of upper 
lip divided into two or three parts. A large, round, dark spot 
near base of anterior portion of dorsal fin, followed by a straight 
row of smaller round spots on interradial membranes. Sides of 



body with about 8 rows of dusky round spots centered on an- 
terior exposed portion of scales. Peduncular spot absent. Dorsal 
fin margin strongly falcate. 

Distribution. — Sumatra (Palembang.GunungSahilan). Bor- 
neo (Kapuas). 

Osteochilus waandersii (Bleeker, 1852) 

(Figure 39) 

Rohik! waamlcrsii Bleeker, 1852,i;:733 (type locality Toboali. Banka). 
Osicochiliis iiuumi'iTi// Gunlher. 1868a:43. 

Material Examined. — Western Borneo: Kapuas 1976-6. 7: 53.2-65.8 mm (CAS 
49241, MZB 3223, USNM 230198); Kapuas 1976-16, 3; 46,8-58.5 mm (MZB 
3224. UMMZ 209857); Kapuas 1976-36, 77,0 mm (MZB 3225); Kapuas 1976- 
39, 46 mm (MZB 3226); Kapuas 1976-47. 44 3 mm (MZB 3227). 

Mesomorph, Body depth 3.0-3.4. Dorsal-fin rays 16-17. Pec- 
toral-fin rays 15-17. Lateral line scales 32-33. Circumferential 
scales 1 1/2/13. Circumpeduncular scales 16. Gill rakers 40-60. 
Snout tubercle 1 or 3; if 3, median tubercle largest. Median 
lateral stripe from posterior border of eye (or from gill opening 
to end of caudal fin). Maximum standard length 205 mm. 

Distribution. — Central Thailand (Khao Phanom Bencha, 
Trang, Old Cheing Sen, Punnang Star, Yala, Chanthaburi, Pak- 
jong, Menam Mun, Kanjanburi. Surathani, Chumporn, Kao 
Saming R.. Trod). Malay Peninsula (Tahan R., Jelai R., Negri 
Sembilan. Pahang R.. Mawai. .lohore. Bukit Merah, Perak). Su- 
matra (Dell, Sockadana, Singkrah Lake, river at Solak, Sid- 
joengdjoeng, Patang). Western Borneo (Kapuas). Java. Banka 
(Tobali Province). 



ROBERTS-WESTERN BORNEO FISHES 



55 




Figure 3v. Uswochihis waandersu. Above. Kapuas 1976-36. 77.0 mm {MZB illi). bcluw. uiKulilcrous lips. Kapuas 1976-6. 65.6 mm (USNM 230198). 



Oxygaster of Gunther. 1868 

Oxygaster wSin Hasselt. 1823:133 (nomen dubium). 

Oxygaster Gunther 1868:332 (included species Cypnnus or Lcuciscus oxygaster 

Valenciennes in Cuvier and Valenciennes. 1844:349). Proposed as a "section" 

of C/jc/a Hamilton-Buchanan, 1822. 

The first ichthyologist to recognize Oxygaster van Hasselt, 
1823 as a valid taxon subsequent to its original publication as 
a nomen dubium is Gijnther ( 1 868). The entire original account 
reads as follows: 

OxygastcrMyhx. onderscheiden vooral door eene mesvormige carina 
aan de buiksvlakte, is misschienten enregte door mij onder de Cypn- 
naceen gesteld. waarom ik nader hierop /a\ tcrugkomen; Anonmlura 
heb ik als species naam onder de Tafel geschreven, 

Oxygaster Mihi. especially distinguished by a knife-like keel on the 
abdominal surface, is perhaps incorrectly placed in the Cypnnes by 
me. for which reason I will come back to this later on. 1 have written 
Anomaliira as the species name under the figure (after .Mfred 1961). 

As it stands, this description of Oxygaster anomalura could 
refer to Macrochirichthys macrochirus. also present in Java and 
obtained by Kuhl and van Hasselt (see Cuvier and Valenciennes 
1 844:348) although not mentioned by van Hasselt (1823). There 
is no mention of specimens or locality, only of a figure that has 
never been published, and even reason to doubt that the fish in 
question belongs to Cyprinidae. Its identity is known only be- 
cause of the account published by Valenciennes, based on a 
specimen obtained by Kuhl and van Hasselt at Batavia. 



Although some authors (e.g.. Smith 1945; Banarescu 1971) 
have recognized Oxygaster van Hasselt, 1823, it is best if O.vr- 
gaster and O. anomalura of van Hasselt are treated as nomina 
dubia. 

Oxygaster was rejected as a distinct genus by Valenciennes 
in Cuvier and Valenciennes ( 1 844), who treated it as a synonym 
oi Leuciscus Valenciennes in Cuvier and Valenciennes, 1844, 
by Bleeker (1860a:469, 1863e:264) and Weber and de Beaufort 
(1916:52), who treated it as a synonym of Chela Hamilton- 
Buchanan, 1822. The statement by Smith (1945:74-75) that 
"Bleeker 1863 [314] made Chela cachiiis of Hamilton the ge- 
notype" of Oxygaster van Hasselt is in error. Bleeker (1863/): 
264) placed Oxygaster as a synonym of Chela and indicated 
Cypnnus (Chela) cachia Hamilton-Buchanan as type species of 
Chela: he did not indicate any type species for Oxygaster. 



Oxygaster hypophthalmus (Bleeker, 1860) 

(Figure 40) 

Che/a hypophthalmus Bleeker. I860ii:47l (type locality Palembang. in fluviis). 

Material ExAMTNiD.— Western Borneo: Kapuas 1976-8, 41. 0-50.9mm(BMNH 
1982.3.29.26-28, CAS 49244. MZB 3232); Kapuas 1976-9. 15: 40.3-51.2 mm 
(FMNH 94206, MNHN 1982-670. MZB 3233. USNM 230199); Kapuas 1976- 
14. 2: 42.5-1 10 mm (CAS 49245. MZB 3234); Kapuas 1976-15. 45.3 mm (MZB 
3235); Kapuas 1976-19, 65.0 mm (MZB 3236): Kapuas 1976-32, 57,0 mm (MZB 
3237); Kapuas 1976-33.6: 28,4-65.1 mm (MZB 3238. UMMZ 209886); Kapuas 
1976-34. 12: 27.7-61.4 mm (CAS 49246. MZB 3239); Kapuas 1976-35. 8: 43.8- 
65,3 mm (MZB 3240, USNM 230200); Kapuas 1976-37. 8: 42.6-50.3 mm (AMNH 



56 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 40. Oxym'^lcr Above. O oxygaslroidi-s. Kapuas 1976-32, 63.2 mm (CAS 49427); middle, O. hypni'lulmlmus. Kapuas 1976-41, 117 mm (MZB 3243): 
below, O hypophlhahinis. Kapuas 1976-32, 57.0 mm (MZB 3237). 



48924. MZB 3241); Kapuas 1976-40. 45.6 mm (MZB 3242); Kapuas 1976-41, 2: 
103-1 17 mm (MZB 3243. ROM 38603); Kapuas 1976-44, 54.5 mm (MZB 3244); 
Kapuas 1976-49. 2; 81 9-92 3 mm (MZB 3245. ZMA 116.521). 

Distribution. — Southern Malay Peninsula, Central and 
northeastern Sumatra (Palembang, Kampar R,; Indragin; We- 
ber and de Beaufort 1916:52). Western Borneo and Sarawak. 



Oxygaster oxygaster (Valenciennes, 1844) new combination 

Oxygasler anomalura van Hassclt. 1823:133 (nomen dubium) 

Cvprinus oxygaster Valenciennes //; Cuvier and Valenciennes, 1844:349 (type 

locality Batavia). 
Leucisciis oxygasler Bleeker. 1853(':453. 
Chela anomahirus Bleeker, 1860u;473. 
Chela oxygaster Weber and de Beaufort. 1 9 1 6. 



ROBERTS-WESTERN BORNEO FISHES 



57 




Figure 41. Paracrossochilm vinatus. Kapuas 1976-24, 58.0 mm (NIFl). 



Material Examined. — Malay Peninsula: Kola Tinggi. Johore, 5: 89.7-104 mm 
(CAS-SU 34665). Western Borneo: Kapuas 1976-16, 7: 64.7-85.4 mm (BMNH 
1982.3.29.25, CAS49242, FMNH 94205, MNHN 1982-669, MZB 3228); Kapuas 
1976-17, 83.7 mm (MZB 3229): Kapuas 1976-37, 2: 64.4-81.7 mm (CAS 49243, 
MZB 3231, UMMZ 209898). 

NoMENCLATURAL NoTE. — Alternation of usage has charac- 
terized this taxon, as it has other names published by van Hasselt 
(1823), because of the difficulty in deciding whether van Has- 
selt's name has validity. 

Distribution. — Malay Peninsula (Malacca). Sumatra (Pan- 
gabuang, Palembang, Lake Tidor, Selapian R., Upper Langkat, 
Wampu R., Deli). Borneo (Kapuas, Baram, Pengaron R.). 

Oxygaster oxygastroides (Bleeker. 1852) 

(Figure 40) 

Leuascus oxygastroides Bleeker, 1852/):431 (type locality Sumatra, Borneo, and 

Java). 
Chela oxygaslroides Bleeker, 1860u:472. 

Chela megalolepis Giinther, 1868a:337 (unwarranted replacement name). 
Oxygasler oxygastroides Fowler, 1935ii: 1 10. 

Material Examined. — Western Borneo: Kapuas 1976-20, 2: 117-131 mm 
(BMNH 1982.3.29.29. MZB 3246); Kapuas 1976-32, 4: 55.1-83.7 mm (CAS 
49247, MZB 3247); Kapuas 1976-34. 109 mm (MZB 3248); Kapuas 1976-44, 5: 
70.2-128 mm (MNHN 1982-671, MZB 3249, UMMZ 209916, USNM 230201); 
Kapuas 1976-49. 1 1 1 mm (MZB 3250). 

Distribution. — Thailand (Mekong; Chao Phrya, Klong Sao 
Tong, Nakon Sritamarat, Tale Sap, Tale Noi, Krabi; Smith 1 945). 
Malay Peninsula. Sumatra (Palembang. Lahat. Lake Tidor, Ba- 
tang Han). Borneo (Kapuas. Sambas. Prabukarta, Makaham, 
Upper Riko). Java (Batavia, Krawang, Lake Kamodjing near 
Tjihampch, Kediri. Tulung Agung). 

Paracrossochilus Popta. 1904 

Paracrossochihis Popta, 1904:200 (type species Paracrossochilus biconiis Popta, 
\')Oi\=Crossochihis rittatiis Boulenger, 1894, by monotypy). 

Rostral cap thick, completely covering upper jaw, with fim- 
briate margin. Lateral lobes of upper lip apparently present 
(papillose), but entire transverse or median portion of upper lip 
absent or vestigial. Homy jaw sheaths well developed. Lower 
lip fleshy, entirely covered with papillae. Rostral and maxillary 
barbels present. Circumpeduncular scales 8-9. 

Endemic to Borneo; only two species known. 



Paracrossochilus acerus Inger and Chin, 1962 

Paracrossochilus acerus Inger and Chin, 1962:100, fig. 47 (type locality Sungei 
Dapu. Baleh River. Rejang basin, Sarawak). 

Material Examined. — Western Borneo; Kapuas 1976-24, 33; 45.8-75.7 mm 
(AMNH 48925. BMNH 1982.3.29.30-31. CAS 44178, FMNH 94207, MNHN 
1982-672. MZB 3251, NIFI uncat., ROM 3861 5); Kapuas 1976-29,6: 20.7-33.0 
mm (MZB 3252, UMMZ 209875, USNM 2.30202). 

Distribution. — Borneo (Rejang, Kapuas). 
Paracrossochilus vittatus (Boulenger, 1894) 

(Figure 41 ) 

Crossochilus vittatus Boulenger, 1894:247 (type locality Senah, Poeh, and Tagora 

Rivers, Sarawak). 
Paracrossochilus hiconus Popta. 1904:201 (type locality le Howong. Mahakam 

basin). See Weber and de Beaufort (1916:227), 

Material Examined —Western Borneo: Kapuas 1976-6, 10: 31.9-49.5 mm, 
AMNH 48926, BMNH 1982.3,29.32-35. MZB 3253); Kapuas 1976-24, 27: 44.7- 
62.7 mm (FMNH 94208, IRSNB 19734, MCZ 58344. MZB 3254, NIFI uncat.); 
Kapuas 1976-25, 7: 34,3-74.8 mm (CAS 49248, MZB 3255); Kapuas 1976-26, 
5: 32.8-52.4 mm (CAS 49249, MZB 3256); Kapuas 1976-27, 21: 19.8-63.4 mm 
(CAS49250, MNHN 1982-673, MZB 3257); Kapuas 1976-29, 27; 20.7-56.7 mm 
(MZB 3258, ROM 38599, UMMZ 209876. USNM 230203, ZMA 1 16.522); Ka- 
puas 1976-30, 21: 26.7-71.7 mm (CAS 53049. MZB 3259. USNM 230204). 



Pectenocypris Kottelat, 1982 

Pcctenocypris Kottelat, 1 982:42 1 (type species Pectenocypris korthausae Kottelat, 
1982, by original designation and monotypy). 

Diagnosis. — Small, slender Rasbora-\\ke cyprinids distin- 
guished from all other genera by extraordinarily elongate and 
numerous gill rakers; pharyngeal teeth expanded, bladelike, with 
incised crowns, in a single row (3/3); upper endentulous limb 
of pharyngeal bone (usually simple in cyprinids) with strongly 
divergent, strutlike anteromedian and anterolateral processes. 

Jaws moderately upturned and short, extending posteriorly 
to about midway between snout-tip and anterior margin of eye. 
Dorsal surface of head flat. Abdominal keel absent. Scales mod- 
erately large, 26-33 in lateral series, 1 1-13 predorsal, 8 trans- 
verse, 11-14 circumpeduncular; lateral line incomplete, with 5- 
10 pored scales anteriorly. Dorsal-fin origin directly above pel- 
vic-fin origin. Dorsal-fin rays ii7, anal iiiS, pectoral 13, pelvic 
8-9. Total gill rakers on first gill arch from 90 to over 200. Total 



58 



CALIFORNIA ACADEMY' OF SCIENCES 




Figure 42. Pccleiiocypris halacna Kapuas 1976-32, 32.3 mm ripe female {MZB 4006, holotype). 



vertebrae 31-32 (present observations) or 36 (Kottelat 1982). 
Largest known specimen 34.9 mm. 

Known only from the type species recently discovered m 
southeast Borneo and an undescribed species obtained during 
the Kapuas survey of 1976. 

Pectenocypris balaena new species 

(Figure 421 

Holotype. — MZB 4006, 32,3 mm. Danau Piam near Kclungau, 3S km NNE 
of Smtang, 5-6 August 1976 (Kapuas 1976-32), 

Paratypes.-BMNH 1982,3.29,107-108, CAS 49307, MZB 3423, USNM 
230238, ZMA 1 16,531, 25; 20,0-34.9 mm. collected with holotype: MZB 3422, 
26 6 mm. small forest streams flowmg into Kapuas mainstream within 10 km 
upri\er of Sanggau. 16-17 July 1976 (Kapuas 1976-17). Two specimens. 27,4- 
30.1 mm. cleared and stained with alcian blue-alizarin (CAS 49307), 

Diagnosis. — Pecfe/jon'pra balaena differs from its only con- 
gener, P. korthausae. in its even more elongate and numerous 
gill rakers, sharper or more slender snout, somewhat more slen- 
der caudal peduncle, and coloration. In many other respects the 
two species are closely similar and they are undoubtedly con- 
generic. 

Counts of gill rakers in two alcohol specimens off. balaena 
examined under water are about 40+115=155 in a 23.6 mm 
specimen and 45+135=180 in a 28.9 mm specimen. These 
counts are possibly somewhat too low due to the minute size 
of the gill rakers at the ends of the upper and lower limits of 
the gill arch. A count of 64+ 148=212 rakers was obtained from 
a 30.1 mm cleared specimen stained with alcian blue-alizarin. 
Kottelat (1982) reported gill rakers 20 + 75=95 in a specimen 
off. /:or?/;a»5af (standard length not indicated) and I find about 
22 + 75=97 in a 21.8 mm paratype of this species (CAS 50488). 
The unusual condition of the pharyngeal bone and pharyngeal 
teeth appears to be very similar in P. balaena and P. korthausae 
(Kottelat 1982, fig. 1). 

Lower jaw distinctly projecting beyond upper (jaws equal or 
lower jaw extending very slightly beyond upper in P. korthau- 
sae). Snout 12.4-13.0 (14.7-17.9 in P. korthausae). Eye 12.0- 
12.7. inlerorbital width 11.5-12.1. Head 3.3-3.5 (3.5-4.3 in f. 
korthausae). Depth caudal peduncle 10.2-10.7 (9.6-12.0 in P. 
korthausae). Pored scales in lateral series 5-8. Vertebrae 16- 
17+15 = 31(1), 32(1). 

Coloration in life not recorded. Preserved specimens with a 
narrow but conspicuous midlateral longitudinal stripe extending 
from snout-tip, across middle of gill cover and length of body. 
The stripe is nearly continuous but is interrupted for a short 
distance on the posterior (membranous) portion of the gill cover 



and body immediately underlying this part of the gill cover. It 
ends abruptly at the base of the caudal fin. In some specimens 
there is a small dark area at the very end of the stripe but none 
has a large oval spot on caudal-fin base like P. korthausae. Side 
of body below midlateral stripe almost entirely free of mela- 
nophores. All fins colorless or plain except for some fine me- 
lanophores narrowly aligned along fin ray margins. Dorsal sur- 
face of head and body with numerous melanophores which form 
a regular pattern covering all of scales except their posterior 
margins. A thin midaxial streak on posterior half of body above 
or along dorsal margin of midlateral stripe (midaxial streak 
extending entire body length in P. korthausae). Ventral surface 
of head and entire abdomen to anal-fin origin free of melano- 
phores. A median dorsal stripe from occiput to dorsal-fin origin; 
a dark stripe along base of anal fin on either side continuous as 
a median ventral stripe on caudal peduncle. 

Food Habits (Table 1). — An analysis of the gut contents of 
one specimen off. balaena from Kapuas 1976-32 was made 
by Albert Mahood, who also took the photomicrographs of 
phytoplankton (Fig. 43). From Table I it can be seen that nearly 
all of the food items range in size from 4 to 40 ^m (longest 
dimension). The principal food both in number of items and in 
biomass visually estimated is Peridinium of 16-18 Mm (appar- 
ently a single species), which account for 1 22 or more than one- 
fourth of the total food items and roughly half of the biomass. 
It is interesting that several Peridinium have intact chloroplasts 
(Fig. 43b), indicating they had been ingested in living condition. 
Other true phytoplankters arc the unicellular Chlorophyceae 
(including Telraedron. of which three species are present) and 
presumably the numerous but very small unicellular Chlorophy- 
ta. The diatoms, on the other hand, are not typical phytoplank- 
ton but rather forms usually associated with bottom sediments. 
These, as well as the pollen? and nearly all other unidentified 
and miscellaneous items encountered, judging from their small 
size, may well be "tychoplanktonic" or "chance plankton." They 
account for about one-third of the food items. It is noteworthy 
that the brown round things identified as pollen? (see Fig. 43i, 
typical of many observed) constitute a substantial item both in 
terms of numbers present and biomass visually estimated. The 
two nematodes may be parasites. Gut contents of two additional 
fish specimens from Kapuas 1976-32 cursorily examined also 
consist largely of Peridinium. 

Dustin Chi vers and I have examined the gut contents of one 
of the paratypes off. korthausae (CAS 50488, 21.0 mm). Its 
gut contents consist mainly of large numbers of the dissociated 
valves, furculae, and antennae of a single species of Cladocera 



ROBERTS-WESTERN BORNEO FISHES 



59 



a 






f 













Figure 43. Phytoplanklon from gut o( Pnlenocypris balaena (Kapuas 1976-32). a-b, Pendiiuuin ( 18 and 16 iim). c. Tctrucilnni (29 ;jm); d, Fuasintm (11 iim). 
e. Slauraslrum (38 Mm); f, Scenedesmus (16 Mm); g-h. fungi? (29 and 34 Mm); i. pollen'' (15 Mm), j, triangular unicellular plant',' (30 Mm), 



(valves 216 x 108 Mm. regularly loculose). together with a large 
amount of loose granular or flocculent material which appears 
to be the partially digested soft parts of the cladoceran. Also 
present are a few diatoms, which may have been ingested by 

Table 1. Pectenocypris bal-iena. Gut contents (Kapuas 1976-32). 







Mm 


n 


Chlorophyta 








unicellular, round 




4-6 


119 


Chlorophyceae 








Tetraedron 




29^2 


51 


Euaslruin 




22 


8 


Cosnianitm 




48 


1 


Slauraslrum 




38 


1 


Scenedesmaceae 








Scenedesmus 




16 


4 


filamentous 




169 


1 


filamentous 




33 


1 


Chrysophyta 








Bacillariophyceae (diatoms) 








Pinnularia 




36 


10 


Eunolia 






1 


Navicula 




22 


1 


Pyrophyta (dinoflagellates) 








Dinophyceae 








Pendiniuiu 




16-18 


122 


Unidentified and miscellaneous 








brown, round (pollen'?) 




9-28 


83 


brown, 4-6 unisenal. Ihick-walled ce 


lis (fungi'') 


18-29 


14 


ovoid, bipartite 




12-18 


25 


oval, clear 




21 


6 


triangular unicellular plant'' 




30 




sponge spicule 




213 




higher plant 




48 




"claw" (copepod?) 








"antenna" (copepod?) 








nematodes 






-) 


Total number of items 






455 



the cladoceran, and a single slender oligochaete (910 x 40 fxm. 
with about 10 very elongate segments). No other food items 
observed. 

Relationships of Pect^a'oo'fr/.s. — Under "relationships" 
Kottelat ( 1 982) stated: "The depressed and somewhat expanded 
ethmoid is typical of chelines. but unlike that group, the ki- 
nethmoid is rod-shaped and not triangular. In overall appear- 
ance, the fish most closely resembles Inlecyphs (see Howes 
1980/i), It appears to have a 'generalized" cheline morphology 
with a highly specialized branchial arch; the pharyngeal bone 
being unlike that known in any other cyprinid." In examining 
both species of Pectenocypris I have been struck by their resem- 
blance to Rashora and particularly to such species as R. subtilis 
which resembles Pectenocypris in its relatively small size, slen- 
der body, and relatively small, upturned mouth. While it does 
not have the extraordinary specializations of the branchial arch- 
es and pharyngeal bones of Pectenocypris. R. subtilis does have 
relatively elongate and numerous gill rakers for a species of 
Rashora. 

Distribution. — fec/CTR^n'/'/'/i balaena is known only from 
the Kapuas. 

Etymology. — The name balaena. also the scientific name of 
the baleen whale, refers to the numerous gill rakers of this species. 

Puntioplites Smith. 1929 

Punuoplitcs Smith, 1929:1 1 (type species Puulius proclo:]Mon Bleeker. 1865, by 

original designation). 
Adamaeypns Fowler, 1934:125 (t\pc species Punlnts proctozy^ron Bleeker, 1865, 

b\ original designation). 

Puntioplites has been revised by Banarescu ( 1 978) and by Taki 
and Katsuyama ( 1 979). 

Superficially similar to some of the species currently included 
in Puntius. but with last simple anal-fin ray greatly thickened 
("spinified"), its posterior border smooth or serrate; gill rakers 
25-42; scales with parallel striae. Barbels absent. Body very 
deep. Lateral line complete, lateral line scales 35-38, circum- 
peduncular 1 6-20. Last simple dorsal-fin ray strong and serrate; 



60 



CALIFORNIA ACADEMY OF SCIENCES 



anal-fin branched rays invariably 5? (at least some reported 
counts of 6 include the last half-ray counted as I [Banarescu 
1978:116]). Pelvic fin with 9 branched rays. Pharyngeal teeth 
in three rows, usually 2,3.4/4,3.2. Vertebrae 33-35. 

Key to Piintioplites of western Borneo 

la Circumpeduncular scales usually 16 (rarely 18); scales on 
side of body with variably darkened ellipsoidal marks 
parallel to their anterior borders, scales with darkest marks 
forming a series of thin, wedge-shaped vertical bars 

P.biiht 

lb Circumpeduncular scales 20; scales without ellipsoidal 
marks, coloration on side of body uniform P. waandersi 

Puntioplites bulu (Bleeker, 1851) 

Svsliiiiuis hull! Becker. 1851/;;207 (i\pe locality Bandicrmassing. in fluviis), 

Barhiis huh Gunlher, 1868<j:14'5. 

Punlius hulii Weher and de Beaulbrl. I o 1 h: 1 44. 

Piinlioplilcs hiilii Banarescu. 1478:1 17 

Material Examined. — Malay Peninsula Perak. Tclok .-Xnson, 1 1 1 mm (CAS- 
SU 31044); Perak. Chenderoh Dam. 3: 155-187 mm (CAS-Sll 3-1720). Western 
Borneo; Kapuas 1476-14. 17; 20.8-47.6 mm (AMNH 48427, CAS 44251, MCZ 
58345. MZB 3260); Kapuas 1976-15; 24.3-48.5 mm (CAS 55328. FMNH 44204. 
MNHN 1482-674, MZB 3261); Kapuas 1476-20, 3; 72.6-103 mm (CAS 44252. 
MZB 3262); Kapuas 1976-33, 2; 48.5-56.3 mm (MZB 3263. USNM 230205); 
Kapuas 1476-34, 43.2 mm (MZB 3264); Kapuas 1976-44. 46.9 mm (MZB 3265); 
Kapuas 1476-45, 3: 21,2-89.5 mm (MZB 3266. RMNH 28838; Kapuas 1976- 
4y, ,v 58..V80,7 mm (MZB 3267, USNM 230206). 

Distribution. — Sumatra (Palembang, Lahat, Rmgat, Danau 
Sialong Lotong). Borneo (Baram?, Kapuas, Kahajan, Banto. 
Kinabatangan). Also reported, perhaps incorrectly, from south- 
ern part of Indochinese Peninsula (Taki and Katsuyama 1979: 
260). 

Pundoplites waandersi (Bleeker. 1858-59) 

SyMiiDius If uaihlcrsi Bleeker. 1858-59/;35S (lype locality Ngawi, Java). 

riinliui {PiinHiis) IVaaudmi Bleeker, 1863-64:1 III. 

Harhiis uuuh(/(ts/ Gunlher, 1868d;l49. 

''Haihii\ proclozysron Volz. 1904:477 (Kwanlan R.. Sumatra) 

I'liniiiis unit Weber and dc Beaufort. 1 9 1 ft;202 (type locality Bunut, Kapuas). Sec 

Banarescu (1478:1 16). 
Svslninus iriiiinus) bulu Bleeker, l863-h4:l II), 
Punlius lawak Weher and de Beaufort, |9lh:2(l4 ( parti m; specimens from Djambi 

and Riouw''). See Banarescu (1978:16), 
Puntuiplilci fcilcatus Smith. 1924; I 1-12 Itype locality Mekong), See Taki and 

Katsuyama (1974:263), 
Punlinplilcs mhiutlcru Banarescu. 1478:1 16. 

Material Examined. — Western Borneo; Kapuas 1976-14. 2: 31.7-82.0 mm 
(CAS 44254. MZB 3269); Kapuas 1976-33. 3: 38.3-112 mm (FMNH 94210. 
MZB 3270. USNM 2.30207). 

Distribution. — Thailand (middle Mekong). Sumatra (Djam- 
bi. Kwanlan?. Palembang. Indragiri). Borneo (Kapuas. Maha- 
kam. Kota Bangun). Java (Ngawi). 

Taki and Katsuyama (1979) reported samples from the mid- 
dle Mekong of Thailand which they identify as P. waandersi. 
These have consistently fewer gill rakers and differ in some 
minor respects from P. waandersi from the East Indies but are 
otherwise closely similar to P. waandersi from Borneo. They 
regard the middle Mekong population as a relict population, 
and suggest that absence of the species elsewhere in continental 
southeast Asia may be due to the widespread occurrence there 



off. proto:ysrfln (a species absent from the East Indies). If this 
Mekong population represents a distinct species, the name Piin- 
tioplites falcatus Smith. 1929 is available for it, 

Puntius Hamilton-Buchanan, 1822 

Punlius Hamilton-Buchanan. 1822:310 (type species ( i /'///»( v .vo/i/id/c Hamilton- 
Buchanan. 1822. by subsequent designation of Bleeker. 1 863/); 1 99). 

Rarhoilc'i Bleeker, 1 860«:275, 3 1 3 (type species SysloniusiBarhodes) belinka Bleeker, 
1860, by subsequent designation of Bleeker. 1863/';200). 

The attempt by Smith (1945:165) to designate the virtual 
tautonym C')7>/v/;/w /»;//;/;() Hamilton-Buchanan, 1822 as generic 
type species of Puntius in place of C sophore is invalid according 
to the current ICZN. 

Puntius as currently understood comprises numerous species 
and may be polyphyletic. Barbels variable, rostral and maxillary, 
maxillary only, or absent. Dorsal fin with last simple ray serrate 
or entire, branched rays usually 8. Anal fin with last simple ray 
entire, branched rays usually 5 (sometimes 6. 8 in one species). 
Lateral line complete or incomplete, lateral line scales 17-36. 
Cephalic cutaneous papillae minute or absent. Phamygeal teeth 
in three rows, usually 2.3.5^5,3,2, Color pattern extremely vari- 
able, many species with up to six very regularly placed vertical 
bars. Young often with color pattern radically ditferent from 
adults. Maximum adult size 30 cm or more {P. schwanenfeldii) 
but most species less than 100 mm. 

Puntius anchisporus (Vaillant, 1902) 

liarhu':{Punliu\) Suniuliuiiu'. Martens. 1876:402. pi. 11. Iig. 2 (Bengkayang. Man- 

dhor. and Danau Sriang). 
Raihiis sunialranus Vaillant. 1893:82 (Kapuas). 
Barbus aiK-liisporus Vaillant. 1402:46. fig, 27 (type locality Kapuas). 
Punlius anciiispnnis Weber and de Beaufort. 1 9 1 6: 1 90, 
Punlius pulchcr KQnii3h\. 1422:203 (type locality Bulungan. northeast Borneo). 

See Alfred (1964:138) 

Material Examined, — Western Borneo: Kapuas 1976-7. 4: 17,4-21,7 mm (CAS 
44255. MZB 3271); Kapuas 1476-16. 5: 30.2-35.1 mm (MZB 3272, USNM 
230208); Kapuas 1976-37. I 5: 32,3-42,8 mm (FMNH 942 1 1, MZB 3273, MNHN 
1482-675. LIMMZ 204894); Kapuas 1476-34. 40,1 mm (MZB 3274); Kapuas 
1976-43. 26,4 mm (MZB 3275): Kapuas 1976-44. 40,0 mm (MZB 3276), 

Body deeper than in any other vertically barred Puntius. depth 
less than 2 (depth more than 2 in other species); dorsal bar 
absent; lateral line complete. Parietal fontanel closed. 

The color pattern of P. anchisporus is \irtually identical to 
that off. telrazona from Sumatra which differs from it in having 
an incomplete lateral line as well as a more elongate body (.Alfred 
1964). 

Distribution. — Known onh from Borneo (Kapuas. Maha- 
kam. Bulungan). 

Puntius binotatus (Valenciennes. 1842) 

Harhu\ inaciilalus van Hasscit, I 823' I 32 (nomen nudum) 

Haibu\ binolalu\ Valenciennes /h Cuvier and Valenciennes. 1842:168 (type locality 
.lava) 

Barbus maculatus Valenciennes ni Cuvier and Valenciennes, 1842:145 (type lo- 
cality Buiten/org), 

Barbus im'sigcncs Bleeker, 1 850; 1 7 (type locality Java), Sec Bleeker ( 1 860u:348), 

Barbus bhtoncnsis Bleeker. I852t(;46 (type locality Billiton), See Bleeker (I860u: 
348), 

Baibin kinan,'n\n Bleeker. 1852/i:429 (type locality Prabukarta. Kusan Rncr. 
southeast Borneo) See Bleeker (I860i;:348l. 



ROBERTS-WESTERN BORNEO FISHES 



61 



Barhus polyspiios Bleeker, 1857/):351 (lype locality "Perdana, Tjibiiiong, provin- 
ciae Bantam Java occidentalis. in fluviis"). 

Systomus (Barhodes) maciilaius Blocker, 1860a:346. 

Syslonnis [Barbodes) goniosoina Bleeker. 1860:349 (type locality Benkuelen. Su- 
matra). 

Biirbus goinosonm\2t\\\3n\. I8'^3:81 (Kapuas). 

Barhus palaranensis Boulenger, 18<)5:186 (type locality Palawan). See Weber and 
do Beaufort (1916:189), and below. 

Barhus quiiiijiiemaatlatiis Seale and Bean, 1907:229 (type locality "near Zam- 
boanga"). 

'!Barhodes hcmicleniis Jordan and Richardson. 1907:241. fig. 5 (type locality Min- 
doro). See below. 

Barhus ivis Seale. 1909:494. pi. 1 (type locality Balabac Isd.. Philippines). 

Barhus elongaliis Sea\e. 1910:265 (type locality Sandakan: preoccupied by Barhus 
cinngalus RiippeW. 1837). 

Barhus maculatus v. hagemi Popta, 191 1:9 (type locality Lombok). 

Puuiius hinoialus Weber and de Beaufort, 1916:186. 

Barhus binolalus var. palavancnsis Herre, 1924:292. 

Barhodes scalei Herre, 1 933 (replacement name for Barhus clungalus Seale, 1910). 
See below. 

Pumius sihukensis Fowler. 1940:799, fig. 25 (type locality Silinipopon R.. Sibuko 
Bay, Borneo). See below. 

Puniius himnatus hanksi Herre. 1940:31 (type locality Kuching. Sarawak, by sub- 
sequent restriction of Bohlke. 1953:37). See below. 

Material Examined. — Malay Peninsula: Singapore. 75: 12.8-79,4 mm {CAS- 
SU 31089). Western Borneo: Kapuas 1976-6. 5: 37,1-85.3 mm (CAS 49256. 
MNHN 1982-676, MZB 3277); Kapuas 1976-7, 18.0 mm (MZB 3278): Kapuas 
1976-25, 2: 45.6-54.6 mm (MZB 3279. USNM 230209). Sarawak: 18 mi E of 
Kuching. 22: 37.3-65.7 mm (CAS-SU 33900. 33903, holotype and paratypes of 
P. hmotatus hanksi). Northern Borneo: Balung R., Tawau, 29: 16,1-64.7 mm 
(CAS-SU 33604); Sandakan, 143: 22.6-1 14 mm (CAS-SU 27805). Java: Buiten- 
/org, 47: 16.7-72.6 (CAS-SU 20483). Palawan: Malatgau R., 5: 34.5-62.2 mm 
(CAS-SU 61641). Malabo. 2: 26.7-27.5 mm (CAS-SU 61640). Culion: stream 
llowing into Halsey Harbor. 35: 40.5-145 mm (CAS-SU 27622). Mindoro: 4: 
77.5-93.0 mm (CAS-SU 20213, 20443, types of P. hcnvacuus): Tarogin R„ 133: 
19,7-52.2 mm (CAS-SU 61643); Hacienda Waterous, Mangarin, 112: 10.7-66.1 
mm (CAS-SU 38099); 536: 18,5-100 mm (CAS-SLI 61644). Basilan: Burakan R.. 
10: 45.4-61 . 1 mm (CAS-SU 23762). Mindanao: Marata Bogan, Lanao Prov., 1 39: 
24.2-87.4 mm (CAS-SU 38098). 

Parietal fontanel closed. Rostral and maxillary barbels well 
developed, both usually longer than eye diameter. Lateral line 
complete; scales in lateral series 23-27. predorsal 8-9, trans- 
dorsal 9, circumpeduncular 12. Last simple dorsal-fin ray finely 
serrate; dorsal-fin branched rays 8; anal-fin branched rays 5. 
Large adults often immaculate or with two round spots, one at 
base of dorsal-fin origin, the other in middle of caudal peduncle. 
Juveniles usually and adults sometimes with 2-4 midlateral 
round spots, variable in position. For further discussion of color 
pattern see below. Attains 145 mm, but adults in most popu- 
lations are under 100 mm. 

This is the most widely distnbuted and perhaps most variable 
species of Puniius in southeast Asia. It occurs from sea level to 
at least 2.000 m, and is commonly found below waterfalls in 
isolated mountain streams and on small islands inhabited by 
few other freshwater fishes. It exhibits great geographical vari- 
ation in body depth and especially in coloration. Samples from 
Malay Peninsula, Borneo, and Java (and Sumatra?) exhibit rel- 
atively little variation; the greatest variation apparently occurs 
in the Philippine Islands; some Philippine Puntius regarded as 
distinct species, such as P heinictenus of Mindoro, may be only 
geographical variants of this species. Puniius binolalus has been 
regarded as the parent stock for all of the endemic genera and 
species of cyprinids in Lake Lanao and on the Lanao Plateau 
of Mindanao (Herre 1955). 

Large adults typically have the body with a dull golden sheen 
and two small round black spots, one at the base of the dorsal- 



fin origin, the other in the middle of the caudal peduncle. Some- 
times these spots may be accentuated, especially the anterior 
spot, which may extend dorsally onto the dorsal fin, or ventrally 
as a vertical bar on the dorsal half of the body. In some spec- 
imens, particularly longer preserved specimens, the spots may 
be very faint or absent. Young individuals have an irregularly 
spaced series of four or more dark round midlateral spots and 
a spot at base of the anal fin. which variably disappear with 
growth. Sometimes the spots are joined in a nearly continuous 
midlateral stripe; in large adults the midlateral stripe is faint or 
absent. Frequently in young fish the dorsal and caudal spots 
may be joined by a vertical bar to a midlateral spot; this con- 
dition is occasionally encountered in adults in some localities. 
In some specimens it forms a large more or less triangular blotch 
with ventrally directed apex (Inger and Chin 1962, fig. 32). In 
others it forms a slender vertical bar; P. binolalus banski from 
Sarawak is based on this color variety, which also occurs in the 
Kapuas, Singapore, and Johore. 

Puniius hemiclenus is endemic to Mindoro and is perhaps the 
only cyprinid species on that island (see Herre 1955). I have 
examined the type specimens (four large adults) and three large 
series all including numerous small young (material listed above 
under P. binolalus) and conclude that it is either a valid species 
closely related to P. binolalus or a geographical race of P. bi- 
nolalus characterized by young without midlateral spots. Jordan 
and Richardson's figure of the holotype depicts a very dark 
midlateral stripe extending from head to base of tail and in- 
corporating the peduncular spot, but in the holotype as in other 
large adults from Mindoro I find that the midlateral stripe is 
faint and only the peduncular spot very dark. Juveniles from 
Mindoro invariably have small dorsal and peduncular spots; 
those in one sample (CAS-SU 38099) also have small anal spots. 
None of the juveniles examined exhibits the midlateral spots 
which occur, so far as I am aware, in juveniles (and sometimes 
adults) of P. binolalus from throughout its range. 

Juveniles and adults of P binolalus from many Philippine 
localities (including Palawan and Mindanao) tend to have two 
large midlateral spots (in addition to the peduncular spot) of 
relatively constant position, one below the dorsal-fin origin and 
the other above the anal-fin origin. The midpeduncular spot 
tends to be equally large and vertically oval, rather than round 
or horizontally oval. Such characteristics are not met with else- 
where except in northeastern Borneo (Tawau, Sandakan). For 
additional information on color variation in the Philippines, 
see Fowler (1941:790-795, figs. 20-23); for other Philippine 
Puniius and their distribution see Herre (1955). Puniius scalei 
(=Punlius clongaius Scale and P. sihul<ensis Fowler), here placed 
as a synonym of P. binolalus. is based on specimens with this 
type of color pattern from northeastern Borneo. 

Distribution. — Thailand (see Smith 1945, for specific lo- 
calities). Malay Peninsula. Sumatra. Borneo (Prabukarta, Bang- 
kajang, Kuching, Kapuas, Tawau, Sandakan, Kinatabangan). 
Java. Nias. Bangka. Billiton. Bali. Lombok. Philippines (Bala- 
bac, Palawan, Culion, Busuanga, Tawi Tawi, Sulu province, 
Basilan, "all Mindanao except Lanao Plateau," Herre 1955: 
123). Mindoro? (See discussion off. hemiclenus above.) 

Puntius bramoides (Valenciennes, 1842) 

Barhus hramouifs Valenciennes ;/; Cuvier and Valenciennes. 1842:160 (type lo- 
cality Java) 



62 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 44. I'ltutim colhiifiwoodn Kapuas 1476-27, 55.7 mm (MZB 3288). 



runlnis {Barhodes) hramnulc Blocker. 1863-64:95. 

.See Weber and de Beaufort (1916:195) for extensive synon>m\. 

Material Examined. — Western Borneo: Kapuas 1976-9, 2: 29.2-50.4 mm (CAS 
49257. MZB 3280): Kapuas 1976-14, II: 18.7-54.6 mm (FMNH 94212. MZB 
3281. MNHN 1982-677): Kapuas 1976-15,376:7.8-50.1 mm (CAS 49258. MZB 
3282); Kapuas 76-31. 7; 20.4-36.8 mm (MZB 3283. USNM 230210); Kapuas 
1976-33. 3: 33.8-44.0 mm (MZB 3284. UMMZ 209888); Kapuas 1976-34, 2: 
23.3-27.8 mm (MZB 3285. CAS 49259); Kapuas 1 976-37. 5: 2 1 .5-34.5 mm (MZB 
3286. USNM 23021 1); Kapuas 1976-40. 42.5 mm (MZB 3991). 

A moderately (ieep-bodied and compressed Puntius: caudal 
fin deeply forked, with pointed lobes bearing submarginal lon- 
gitudinal stripes of equal intensity. Parietal fontanel closed. Ros- 
tral and ma.xillary barbels well developed, somewhat longer than 
eye diameter. Lateral line complete. Lateral scale series 34, pre- 
dorsal scales 13. transdorsal scales 16. scales between lateral 
line and pelvic fin origin 3-1/2. Last simple dorsal-fin ray mod- 
erately stout, serrate. Total gill rakers on first arch 9. 

This species superficially resembles and is perhaps closely 
related to two others found in western Borneo. P collingwoodu 
and P. schwanenfeldii . 

Distribution. — Thailand (Mekong, Chao Phrya). Malay 
Peninsula (Tembeling R.. Pahang; Bukit Merah. Perak). Borneo 
(Baram, Kapuas. Barito, Mahakam). .lava (Batavia, Tjibitong, 
Tandjong Oost. Buitenzorg. Tjampea, Tjikao. Parongkalong, 
Kuningan, Banjumas. Ngawi. Surabaya, Kedin). Unknown from 
Sumatra? 

Puntius collingwoodii (Giinther. 1868) 

(Figure 44) 

liarbiis ciillingmmdii Gilnlha. 1868ij;483 (t>pe localitv Sarawak). 
Harhiis slngatiis Vaillant. 1902:98 (not of Boulenger; Kapuas). 
Burhiis BoiilengeriiPopia. 1 905: 172 (type locality Bongan. Howang. and Bo rivers, 
Kapuas and Mahakam basins). 

Material Examined. — Western Borneo; Kapuas 1476-24. 5: 43.1-71.7 mm 
(CAS 49260. MZB 3287. BMNH 1982.3.29,36-37); Kapuas 1976-27. 3: 39.1- 
55.7 mm (MZB 3288. UMMZ 209870). 

A Puntius with compressed body and slender caudal peduncle; 
caudal-fin lobes with submarginal longitudinal stripes, that of 
lower lobe distinctly darker. Last simple dorsal-fin ray mod- 
erately stout, serrate. Lateral line complete. Lateral scale series 
34, predorsal 15. transdorsal 13, circumpeduncular 16. Parietal 



fontanel closed. Rostral and ma.xillary barbels well developed, 
about as long as eye diameter. Total gill rakers on first arch 9. 
Distribution. — Borneo (Sarawak. Baram. Tinjar, Senah, Ka- 
puas, Mahakam, Tempasuk R., Kota Belud). 

Puntius endecanalis new species 

(Figure 45) 

HoLOTVPE. — MZB 4003. 48-2 mm. small forest stream where it Hows into Sungai 
Mandai 2-3 km upstream from its confluence with Kapuas mainstream. 17 km 
WSW of Pulussibau (Kapuas 1976-37). 

Paratypes.-BMNH 1982,3 29,38-39. CAS 49261. FMNH 94213. MNHN 
1982-678. MZB 3289. RMNH 28840. UMMZ 209895. 16: 29.1-47.8 mm. same 
locality as holotype; MZB 3290. ZMA 1 16.523. 2: 34.6-36.2 mm. Sungai Mandai 
Ketchil near its confluence with Kapuas mainstream. 18 km WSW of Putussibau 
(Kapuas 1976-39); MZB 3291. USNM 230212, 4: 28.4-33.5 mm. small oxbow 
lake cut off from Kapuas mainstream opposite Empangau, 1 24 km NE of Sintang 
(Kapuas 1976-43). 

Diagnosis. — A f (//;//;« with 8 (or rarely 7) branched anal rays 
and a distinctive color pattern of six bars on the head and body. 
.^11 other Punlius have 5 (occasionally 6) branched anal-fin rays, 
and in the few species with a color pattern of six bars the shape, 
extent, and position of the bars differs from that in the new 
species. [Note: .According to R. M. Bailey (pers. comm., Nov. 
1 984). Puntius gonionotus usually has 6 branched anal rays and 
P. altus often has 6.] 

Dorsal profile from snout-tip to dorsal-fin origin straight. Pa- 
rietal fontanel open. Rostral and maxillary barbels present. 
Maxillary barbel well developed, its length from two-thirds to 
slightly more than eye diameter. Rostral barbel relatively weakly 
developed, much thinner than maxillary barbel and about one- 
third as long. Gill rakers very short, 1+6=7 in several speci- 
mens. Pharyngeal teeth (examined in two specimens 45.8 and 
47.8 mm) uncinate, in three rows. 2,3,4/4,3,2; tooth 4 in row 
3 (innermost row) greatly enlarged; teeth 1-3 in row 3 usually 
bicuspid, with a small but well formed minor cusp projecting 
from shoulder and especially prominent on teeth 2-3. Pharyn- 
geal bone (ceratobranchial 5) elongate and relatively slender 
compared to other Puntius (cf. Taki et al. 1978). its dorsal limb 
evenly cur\ed medially, dorsal angle absent. 

Dorsal-fin rays iv8-l/2. Last simple dorsal-fin ray with basal 
two-thirds bearing 12-22 serrae, distal one-third without serrae 



ROBERTS-WESTERN BORNEO FISHES 



63 




Figure 45. Punlius cndecanalis. Kapuas 1976-37, 48.2 mm (MZB 4003, hololypc). 



and relatively slender (some, perhaps most. Puntius have last 
simple dorsal-fin ray serrate to its tip). Anal-fin rays usually iiiS- 
1/2 (iii8-l/2 in 22, iii7-l/2 in 1). Pectoral-fin rays 15-16, pelvic 
8-9. Lateral line complete. Pored scales in lateral series 24-27. 
Scale rows between dorsal fin and lateral line 5-6, between 
lateral line and pelvic fin 3. Predorsal scales 9-1 1, circumpe- 
duncular 1 2. Proportional measurements very uniform through- 
out type series, 48.2 mm holotype and five paratypes 29. 1-46.7 
mm from Kapuas 1976-37 with head length 3.1-3.2; snout 9.9- 
10.6; eye 9.1-10.2; interorbital width 1 1.4-1 1.9; maxillary bar- 
bel 8.9-13.4; body depth 2.6-2.8; caudal peduncle depth 7.1- 
7.5. 

Color in life pale whitish with six black bars; fins generally 
colorless, except where bars extend from body onto basal por- 
tion of dorsal and anal fins near their origin; some specimens 
with a concentration of melanophores along serrated portion of 
third dorsal-fin ray. Color pattern in preserved specimens (29. 1- 
48.2 mm) is very uniform; all have six bars on head and body. 
These bars correspond to the orbital, pectoral, subdorsal, su- 
praanal, postanal, and caudal markings that are variably de- 
veloped in many species of Puntius. 

The orbital bar is complete dorsally. forming a broad, pos- 
teriorly directed, chevron-shaped mark on the posterior part of 
head and anterior portion of nape; the inverted portion of the 
chevron lies over the posterior half of the parietal fontanel, while 
skin overlying the anterior half of the fontanel is pale whitish; 
ventrally orbital bar continues as a short poorly defined vertical 
mark below middle of eye. The pectoral bar is complete dorsally; 
its dorsal half (above lateral line) is vertically oriented, while 
Its ventral half slants slightly posteriorly and ends abruptly about 
midway between and on a level with pectoral- and pelvic-fin 
origins. The subdorsal bar is complete dorsally (extending onto 
basal portion of anteriormost 3-4 dorsal-fin rays), vertically 
oriented, and very short, extending only 3^ scale rows below 
dorsal-fin origin. The supraanal mark is complete dorsally and 
ventrally, and vertically oriented; dorsally it lies on and im- 
mediately posterior to last anal-fin ray. and onto basal portion 
of last anal-fin ray; ventrally it extends a short distance onto 



anal fin near its origin. The postanal and the caudal bars are 
complete (joined both dorsally and ventrally with correspond- 
ing bars of opposite side of body), vertically oriented, and (es- 
pecially caudal bar) somewhat more slender than other bars. In 
some specimens midlateral portion of postanal bar is expanded 
into a round spot, but in all specimens postanal bar is continuous 
(i.e.. round spot is not isolated). There is no separate nuchal 
mark; it is either absent or has become part of chevron-shaped 
mark formed by dorsal junction of orbital bars. There is also 
no separate anal-basal mark; it is either absent or amalgamated 
with ventral portion of supraanal bar. There is a dark round 
spot on dorsum of snout anterior to eyes, and dark pigmentation 
on anterior margin of rostral flap (overlying upper lip); these 
marks have not been noted in other Puntius. 

The nomenclature of bars or marks used here is that of Taki 
et al. ( 1978), who recognized seven types of barred color patterns 
(A-G) in the 23 species oi Puntius they studied. The color pat- 
tern of P. cndccanalis does not correspond to any of these types. 
It is closest to their type C, defined as having cross-bars or 
roundish blotches in pectoral, subdorsal, supraanal, and caudal 
positions, and sometimes an orbital mark. Puntius endecanalis 
has all of these marks, and also a postanal bar. Thus its barred 
pattern seems to have all of the vertical marks recognized in 
PurUius. 

Distribution. — Pm/;//!« endecanalis is known only from the 
type series of 23 specimens collected in the Kapuas in 1976. 

The only other Punlius species with a full complement of six 
black marks on the head and body apparently are the southeast 
Asian P. foerschi Kottelat. 1982, P. pentazona. and P. rhom- 
boocellatus. These three species all occur in Borneo; they are 
very similar to each other but not to P. endecanalis. These three 
species agree with each other but differ from P. endecanalis in 
shape of dorsal-fin base, morphology of simple dorsal-fin rays, 
shape of snout and orbit, and in the shape, position, and extent 
of black marks on head and body. The fleshy portion of the 
dorsal-fin base in the three species is strongly elevated or convex, 
and covered by two scale rows rather than one as in P. en- 
decanalis and most other Puntius species in which the dorsal- 



64 



CALIFORNIA ACADEMY OF SCIENCES 




FiouRh 4h. I'limnis. Above, P. eugrammm. Kapuas 1976-16, 48.6 mm (CAS 49262); below. P. hnealus. Kapuas 1976-32, .14.4 mm (CAS 49266). 



fin base is not so elevated. The last or fourth simple dorsal-fin 
ray is serrate and relatively thick for its whole length; its anterior 
proximal portion is deeply incised and intimately overlain by 
the posterolaterally projecting margins of the third simple dor- 
sal-fin ray which is strongly concave posteriorly (in P. endeca- 
nalis fourth dorsal-fin ray not strongly incised near base, third 
dorsal-fin ray not notably concave posteriorly). At least in P. 
peiUazona and P. rhombooccllatus the orbital bars are not com- 
plete dorsally but extend dorsally only to lateral margin of pa- 
rietal fontanel (condition of orbital bars unknown in P. focrschi). 
In all three species the subdorsal bar extends farther ventrally, 
the supraanal bar lies farther posterior (especially relative to 
dorsal fin) than in P. cndecanalis. and there usually is a well 
developed isolated spot near base of last dorsal-fin rays. 

Etymology.— The name cndecanalis (Latin) refers to the 
characteristic number of anal-fin rays in this species. 



Material Examined— Western Borneo: Kapuas 1976-16, 3: 37.1—48.6 mm 
(CAS 49262, MZB 3292): Kapuas 1976-18. 2: 18,8-19.2 mm (CAS 49263, MZB 
3293): Kapuas 1 976-20, 6: 76. 1-88.1 mm (t AS 49264, MZB 3294, USNM 2302 1 3), 

A moderately large Puninis. attaining 100 mm, with long 
barbels; adults with 4-6 dark longitudinal stripes on side of 
body. Parietal fontanel open. 

Individuals of this species undergo a remarkable color change 
as they grow (Taki 1978, fig, 4c-e; pers, obs,). Young of 18-20 
mm ha\e fi%c broad vertical bars and no indication of the lon- 
gitudinal stripes characteristic of adults. In young of about 25 
mm the vertical bars start to break up and become replaced by 
longitudinal stripes, of which the midlateral is first to appear. 
By 50 mm the change in color pattern is complete. 

Distribution.— Sumatra. Borneo (Kapuas, Mahakam, Ba- 
rito, Akar?). Banka, 



Puntius eugrammus Silas, 1956 

(Figure 46) 

Barhw. fascialus Bleeker, 1853a':l90 (not ol Jcrdon, 1849: type locality "Mara- 
wang, m lluMis [Banka): Moara Kompeh, Sumatrae onentalis, m fluviis"), 

Syslnnuis iBiirhnilcs) fusiuiln.s Bleeker, 1860ij:344. 

Punliiis (.BcirhiMk's) famalii'. Bleeker. 1863-64:103. 

Barhiis fasciatus. Bleeker: (var. Chupcn) Vaillant, 1902:13 (nomen nudum). 

Pitnnus eiixnimniiit Silas. 1956:194 (replacement name for Barhtis {=Pimlim) 
fasciatus Bleeker, 1853, preoccupied by Cinhinus {=Piinlius) lascuitus Jcrdon, 
1849), 



Puntius everetti Boulenger, 1894 

Barhus lcliazi>na^ti\\\3n\. 1893:81 (not of Bleeker. 1855: Kapuas). Re-identi- 

licalion by Alfred 1964:140. 
Barlnis Everclii Boulenger. 1894:248 (type locality Poeh. Sarawak). 
Piinlnis cvcrcui Weber and de Beaufort. 1916:180. Imaki ct al. 1978:14. pi. 6 

(small rivers near Sanggau. Kapuas basin). 

Material Examined —None. 

This species was not obtained during the Kapuas survey of 
1 976. It IS included here on the basis of identifications of Kapuas 



ROBERTS-WESTERN BORNEO FISHES 



65 



specimens by Alfred ( 1 964) (MNHN 1 89 1.352-3) and Imaki et 
al. (1978). 

Puntius lateristriga (Valenciennes, 1842) 

Barbus lateristriga Valenciennes m Cuvier and Valenciennes, 1 842: 1 6 1 (type lo- 
cality Java). 
Sysloimis (Barbodes) lalcristnga Bleeker, 1860a:342. 
Punlius lateristriga Weber and de Beaufort. 1916:179. 

Material Examined, — Western Borneo: Kapuas 1976-6, 13: 23.5-60.3 mm 
IC'^S 49265. FMNH 94214. MNHN 1982-679, MZB 3295); Kapuas 1976-25.2: 
42.0-53.7 mm (MZB 3296. RMNH 28841); Kapuas 1976-30. 7: 25.8-92.4 mm 
(MZB 3297, ROM 38619, UMMZ 209877. USNM 230214). 

A moderately large (to 1 50 mm), heavy-set or chunky, mod- 
erately deep-bodied Punlius: predorsal profile increasingly steep 
with growth; body generally with two broad vertical bars in 
anterior half and a broad midlateral longitudinal stripe on pos- 
terior half; on an overall background color of light lemon yellow 
in life. Often a roundish black spot above anal-fin origin. Lateral 
line complete, lateral scales 23. 

Geographical variation in color pattern in Malay Peninsula 
has been described by Tweedie (1961). 

Distribution. — Peninsular Thailand, Malay Peninsula 
(Tweedie 1961), Sumatra (numerous localities cited by Weber 
and de Beaufort 1916), Borneo (Bangkajang, Seminis, Lumar, 
Sadong, Senah, Kapuas). Java (Batavia, Buitenzorg, Tjampea, 
Sadingwetan, Tjipanas). Banka, Billiton, Singkep, 

Usually inhabits clear mountain streams strewn with rocks 
and boulders; frequently found below waterfalls. 

Puntius lineatus (Duncker, 1904) new combination 

(Figure 46) 

Barbus lineatus Duncker, 1904:180 (type locality Muar River at Tubing tinggi). 

Material Examined— Malay Peninsula; Tasek Bcrah, Pahang, 2: 39.2—41.9 
mm (CAS-SLF 31097). Western Borneo: Kapuas 1976-32. 6: 29,9-38.5 mm (CAS 
49266, MZB 3298. USNM 230215). 

This very distinctive species seems not to have been reported 
or recognized since its original description, perhaps because 
Duncker (1904:180) suggested that it might be only a geograph- 
ical race or variety of P. fascialus (=P. eugrammus). Its linear 
color pattern, with 5-6 longitudinal stripes on the body, is strik- 
ingly similar to that of P. eugrammus but in other respects the 
two species are so different as to indicate they are not very closely 
related. 

Punlius lineaius. the largest known specimen of which is a 
sexually mature female of only 41.9 mm, evidently is a much 
smaller species than the somewhat heavier-bodied P. eugram- 
mus. which attains at least 100 mm. In P. lineaius the rostral 
barbels are absent and the maxillary barbels are absent (ac- 
cording to Duncker) or relatively short, their length less than 
half eye diameter (material examined); both pairs of barbels are 
very well developed in P. eugrammus. as long or longer than 
eye diameter. Puntius lineatus has gill rakers on first gill arch 
moderately elongate, 18-19; in P. eugrammus they are very 
weakly developed, 10, Parietal fontanel open. There is no in- 
dication that P. lineatus ever exhibits a vertically barred color 
pattern or goes through anything like the complex and protracted 
change in color pattern from vertical bars to longitudinal stripes 
observed in P. eugrammus at 18-50 mm. The 39.2 mm spec- 
imen off. lineatus from Pahang is apparently a sexually mature 



male; the dorsal surface of the first four pectoral-fin rays is 
densely covered with fine breeding tubercles. The 41.9 mm 
specimen from the same collection is a female with ripening 
ovaries and non-tuberculate pectoral fins. Breeding tubercles 
have not been observed on the pectoral fins of sexually mature 
P. eugrammus males. 
Distribution, — Malay Peninsula, Kapuas, 

Puntius pentazona (Boulenger, 1894) 

Barbus pcniazona Boulenger. 1894:248 (type locality Baram R.). 

Barbus {Barbodcs) hcxazona Weber and de Beaufort. 1 9 1 2:527. pi. 11. fig. 2 (type 

locality Taluk and Gunung Sahilan. Sumatra). See Alfred (1963). 
Puntius pentazona Weber and de Beaufort. 1916:182 (pro parte). 
Puntius hexazona Weber and de Beaufort. 1916:181. fig. 73. 

Material Examhmed. — Western Borneo: Kapuas 1976-1. 34.8 mm (MZB 3299); 
Kapuas 1976-1 1. 8: 19.4-34.9 mm (BMNH 1982.3.29.40-41, CAS 49267. MZB 
3300. USNM 230216); Kapuas 1976-16. 23.9 mm (MZB 3301). 

This species has the maximum complement of six vertical 
bars seen in Puntius. In naming it "pentazona" Boulenger did 
not consider the vertical bar on the head. Alfred (1963) recog- 
nized two subspecies, based on the presence of a round black 
spot at the base of the dorsal-fin termination (the nominal sub- 
species, P. pentazona pentazona. from Sarawak) or its absence 
{P. pentazona Johorensis {Duncker. 1904), from the Malay Pen- 
insula and Sumatra). On this basis the Kapuas material is re- 
ferable to the latter subspecies. 

Distribution, — Malay Peninsula (Malacca, MuarR., Johore, 
Singapore, Selangor, Perak, Trengganu, Alfred 1963), Sumatra 
(Gunung Sahilan; Indragiri, Rawang, Djambi; Taluk), Borneo 
(Baram, Akah, Saribas, Sibu; Alfred 1963; Kapuas). 

Puntius rhomboocellatus Koumans, 1950 

Puntius rhomboocellatus Koumans. 1950:189 (type locality about 15 km from 
Bandjermasin in a canal along the highway from Oelin to Bandjermasin), 

Barhus telrazona Bleeker. 1857(7:14 (nee Bleeker 1855; type locality River Ka- 
hajan. Borneo), 

Punlius (Barbodes) telrazona Bleeker. 1863-64, 

Barbus kahajani Hoedeman. 1956:288 (rcplaccmem name for Barbus letrazoiia 
Bleeker. 1857). 

For discussion of synonymy see Alfred (1964: 1 39-140). 

Material ExAMiMED— Western Borneo: Kapuas 1976-4.2: 29. 6-3 1.7 mm (MZB 
3302. ZMA I 16.524); Kapuas 1976-5. 6: 17.0-32.0 mm (CAS 49268. MZB 3303. 
USNM 230217): Kapuas 1976-17. 21 7 mm (MZB 3304). Kapuas 1976-32. 21,9 
mm (MZB 3305), 

A Puntius very similar and apparently closely related to P. 
pentazona but with the predorsal, dorsal, and anal bars ex- 
panded to form "ocellate rhombi," the dorsal and sometimes 
predorsal bars not extending ventrally so far as in P. pentazona. 
Color pattern of the Kapuas specimens agrees well with that of 
the holotype off. rhomboocellatus (Alfred 1964, fig. le), al- 
though they have the central portion of rhombi less pronounced- 
ly pale. A midventral dark spot between pelvic-fin bases (not 
extending onto pelvic fins) (absent in P. pentazona). Parietal 
fontanel open. Lateral line complete, lateral line scales 24-25; 
predorsal scales 9-10; transdorsal scales 9; circumpeduncular 
scales 12, Complete scale rows between dorsal-fin origin and 
lateral line 4 (5 in P. pentazona). Total gill rakers on first arch 
10. 

Distribution. — Borneo (Kapuas, Barito. Kahajan), See Alfred 
(1964), 



66 



CALIFORNIA ACADEMY OF SCIENCES 



Puntius schwanenfeldii (Bleeker. 1853) 

Rarhm SchKancnkidii Bleeker. 1853/;517 (lypc locality Sumatra). 

Srslonms [Barlwdi":) Sclnvamicldi Bleeker. l860u;3:.V 

Punlnis iBdihodcs) Sih»'uihii-ldi Bleeker. 186.3-64:94 (unwarranted change of 

spelling). 
Barhus Schn-uncfcldi var. nihn: Vaillant. I902:'^5 (type locality Mahakam River 

at Tepoe). 
Punlnis schnunefeldi Weber and de Beaufort. 1416:178. 
Punnus schwanenfeldii Smith. 1945: li^O. 

Material Examined. — Malay Peninsula: Singapore. 25: 23.9-69.2 mm (CAS- 
Sll 34717) 

A very deep-bodied and large (to 350 mm lota! length) Pun- 
tius: body at all sizes without marks; dorsal fin with a large black 
blotch anteriorly, upper and lower lobes of caudal fin with dark 
submarginal longitudinal bands. Large individuals silvery or 
golden yellow in life, with dorsal fin red and caudal fin orange 
or blood-red (Smith 1945:190). 

Specimens from Singapore have lateral line scales 33-34, 
transdorsal 14-15, predorsal 13-14. circumpeduncular 16-17, 
and total gill rakers on first arch 9, short. Hardenberg (1936) 
reported 18-20 circumpeduncular scales in Kapuas specimens. 

Although no specimens of this species were obtained during 
the Kapuas survey of 1976, a large individual was seen in a 
fisherman's catch on the Sungai Pinoh near its confluence with 
the Melawi; it has been reported from the Kapuas by Bleeker 
(1863), Vaillant (1893, 1902), Hardenberg (1936), and Imaki et 
al. (1978). 

Distribution. — Thailand (Mekong, Meklong, Tachin, Chao 
Phrya, Chantabun, Sikuk, Tapi, Tale Sap, Tale Noi, Patani). 
Malay Peninsula (Malacca). Sumatra (Padang, Solok, Lake Sing- 
karah, Padang Pandjang. Pajakombo, Lake Manindjari, Kwan- 
tan or Indragiri, Si Djungdjung, Taluk, Ringat, Kampar R., 
Lamatang ilir R., Musi, Kompeh, Pangabuang). Borneo (Baram, 
Kapuas, Mahakam). 

Rasbora Bleeker, 1859-60 

Riishora Bleeker. l859-60c:435 (type species C'vi'iiinn nishnni Hamilton-Bu- 
chanan, 1822. by absolute tautonymy). 

Parlueiosoma Howes. 1 980: 183. 1 94 (1\ pe species Leiiei\eus urgyrouienui Bleeker. 
1850. by original designation) 

Barbels absent; jaws small, oblique; lower jaw with an up- 
wardly projecting symphyseal knob fitting into a concavity in 
upper lip when mouth is closed; gill rakers 6-22; pharyngeal 
teeth in three rows, elongate, uncinate, typically 2, 4, 5, /5, 4, 2. 
Lateral scale rows 21-44, predorsal scales 9-17; dorsal fin usu- 
ally with 8 (rarely 9) branched rays, its origin over or slightly 
posterior to a vertical through pelvic-fin origin; anal-fin branched 
rays usually 5 (rarely 6). 

As currently understood, Rashora comprises some 55 species 
and IS distributed throughout the Indian subcontinent, southern 
China, and continental southeast Asia (including Sumatra, Bor- 
neo, and Java). Somewhat less than half of the species are known 
from western Borneo (many of them recorded herein for the 
first time). It is likely that the species thus grouped together are 
polyphyletic; they probably could and perhaps should be placed 
in two or more separate genera. Howes (1980:194) proposed a 
new genus Parlueiosoma. and referred to it the southeast Asian 
species R. arg)'rotaenia. cephalotaenia. dusonciisis and volzi. as 
well as the Indian R. daniconius. It is evident that, \i Parlueio- 



soma is recognized, it will include additional southeast Asian 
species such as R. myersi. etc. LIntil the generic assignment of 
a number of species has been more thoroughly investigated, it 
seems best to refer to all of them as Rasbora. For further in- 
formation see the revision of Rashora by Brittan (1954, 1972). 

The best guide to identification of species of Rashora is Brittan 
(1954, 1972). The reprint edition of 1972 includes a series of 
color photographic plates with some new information, but the 
basic text is identical with that in the original work. The main 
change of concern here (incorporated in the figure legends of 
the added plates) is that the species earlier known as R. taemata 
Ahl, 1922 IS identified as R. agilis. The original description of 
R. taeniala is based on a poorly preserved immature specimen 
that is now lost, and I agree with Brittan that it should be 
regarded as a nomen dubium. A number of species have been 
described subsequently to 1972, notably R. a.xelrodi and R. 
bnttaiu but are readily identifiable. Brittan did not include in- 
formation on gill raker counts which I find helpful in distin- 
guishing certain species. Counts for all species I have been able 
to examine are presented in Table 2. Morphologically similar 
species with different gill raker counts include the western Bor- 
nean R. agilis and R. pauciperforata; R. hankanensis and R. 
emiealepis; and R. suhtilis and R. trilineata. 

Rashora hcauforti. known only from the Kumai River in 
southwest Borneo, may also have to be treated as a nomen 
dubium unless type specimens can be located. I have not been 
able to locate the types of any of the four new species including 
this Rashora described by Hardenberg (1938). They are not 
deposited in Amsterdam or Leiden, and my attempts to locate 
them elsewhere have been unsuccessful. Rashora horneensis, 
placed as a synonym of R. argyrotaenia by Weber and de Beau- 
fort (1916) and not recognized subsequently, is a very distinct 
species here reported from the Kapuas basin for the first time. 
So far as I have been able to determine, reports of R. leptosoma 
Bleeker, 1855 trom the Kapuas are based on misidentified R. 
argyrotaenia or other species; the only specimens of this dis- 
tinctive species known to me are Bleeker's types from Lahat, 
Sumatra. 

The Kapuas survey of 1976 obtained 22 species of Rashora. 
of which R. ennealepis and R. suhtilis are described as new. 
Further exploration in western Borneo undoubtedly will un- 
cover additional species. 

Rasbora agilis Ahl, 1937 

Rashora ai;dis Ahl. 1937:1 13 (type locality "Sumatra": described from aquarium 

specimen) 
Rashora laeniaui Knuan. 1949, 1954 

Material Examined. — Mala\ Peninsula: Trengganu, Merchang, 10: 24.2-37.1 
mm (CAS-Sll 47237), Western Borneo: Kapuas 1976-5. 10:26.5-34.1 mm (BMNH 
1982.3.29 42-44. FMNH 94215. MZB 3306); Kapuas 1976-7, 8: 24,2-31.0 mm 
(CAS 49269. MZB 3307); Kapuas 1976-8. 16: 13.4-33.4 mm (FMNH 94216. 
IRSNB 19735. MZB 3308); Kapuas 1976-10. 7: 16.7-25.8 mm (MCZ 58346. 
.MZB 3309); Kapuas 1976-16, 12: 2 1 .2-25.2 mm (MNHN 1982-680. MZB 33 10, 
RMNH 28842); Kapuas 1976-17. 3: 19,2-27.8 mm (MZB 3311. ROM 38624); 
Kapuas 1 976-2 1 . 1 7,6 mm (MZB 3992); Kapuas 1 976-43, 2:29,4-35.5 mm (MZB 
3312. flMMZ 209909); Kapuas 1976-47. 212 mm (MZB 3993); Kapuas 1976- 
51.4: 23.7-26.2 mm (MZB 3313. HSNM 230218) 

A slender Rashora with a very high, pointed dorsal fin, elon- 
gate caudal peduncle and deeply forked caudal fin. Live speci- 
mens are largely translucent and often colorless except for the 



ROBERTS-WESTERN BORNEO FISHES 



67 




Figure 47. Rasbora a.xclrodi Rapuas l'575-5. Above. 18.9 mm male (MZB 33201; below, 19.0 mm female (MZB 3320). 



opaque white abdomen and continuous black lateral stnpe run- 
ning just below horizontal septum. Lateral line incomplete, with 
only 2-6 pored scales anteriorly. Lateral line scales 29-33, pre- 
dorsal 12-14, circumpeduncular 12. 

According to M. Brittan (pers. comm.), this species was in- 
correctly identified in Brittan (1949, 1954) as R. laeniata Ahl, 
1922, based on an aquarium specimen supposedly from Su- 
matra. The identity ofR. taeniata is unknown; its holotype, like 
that of ^. agilis. was deposited in the Berlin Museum, and may 
have been destroyed. For further discussion see Brittan (1949). 

Distribution. — Malay Peninsula (Johore). Sumatra? (records 
based on aquarium specimens). Borneo (Kapuas). Billiton. 

Rasbora argyrotaenia (Bleeker, 1850) 

Lcuciscus argywlaema Bleeker. 1850:21 (t>pe locality Java). 
Rasbora argyrolaenm Bleeker, 1860a:448. 

Material Examined. — Western Borneo: Kapuas 1976-16. 8: 31.7-50.8 mm 
(,AMNH 48928. CAS 49270, MZB 3314); Kapuas 1976-17, 26: 29,4-49.8 mm 
(BMNH 1 982.3.29.45^9, FMNH 942 1 7. IRSNB 1 9736, MZC 58347, MZB 3315); 
Kapuas 1976-42, 6: 49.7-52.6 mm (MZB 3316, NIFI uncal.. RMNH 28843), 
Kapuas 1976-43, 4: 39.0-46,7 mm (MZB 3317, ROM 38621); Kapuas 1976-46, 
9: 32,2-52.4 mm (MZB 3318, UMMZ 209921); Kapuas 1976-47, 36,7 mm (MZB 



3994); Kapuas 1976-51, 1 1: 40.5-59.2 mm (MZB 3319, USNM 2.30219, ZMA 
116.325), 

For information about this common, widely distributed, and 
variable species, of which several subspecies and races have been 
described, see Brittan (1954, 1972). 

Rasbora axelrodi Brittan, 1976 

(Figures 47, 48) 

Rasbora axelrodi Brittan, 1976:94 (type locality Sumatra'^; descnbed from aquar- 
ium specimens). 

Material Examined, -Sumatra'': 3: 14,8-18,2 mm (CAS-SU 36685-6, holo- 
tvpe and paratypes); western Borneo; Kapuas 1976-5, 77: 7,3-19,3 mm (BMNH 
1982,3,29,50-54,CAS49271, FMNH 94218, IRSNB 19737, MCZ 58348, MNHN 
1982-681, MZB 3320, RMNH 28844, UMMZ 209853, USNM 230220), 

Diagnosis, — A very small Rasbora-Uke cyprinid, possibly re- 
lated to Danioninae, differing from all other Rasbora in its sexual 
dimorphism and sexual dichromatism. Largest specimen 19.3 
mm. Head relatively blunt, body moderately deep, caudal pe- 
duncle slender. Lateral line absent (i.e., without tube- or pore- 
bearing scales); lateral scales 32, predorsal 1 2, circumpeduncular 
12. Dorsal fin with 6 branched rays, anal with 5. In life, large 



68 



CALIFORNIA ACADEMY OF SCIENCES 



Table 2. Frequencies of Gill Rak£R Counts in Rasbora 



R. agilis 
Malay P.. Trengganu (CAS-SU 47237) 
Kapuas 1976-7 

R. agyrolaenia 

Kapuas IQ76-I6, 42 
R axclrodi 

Kapuas 1976-5 
R haukancns!^ 

Kapuas 1976-9 
Kapuas 1976-27 
Kapuas 1976-29 

R borapelcnsis iSnwXh. 1934) 

Bangkok (CAS-SU 44396) 
R bonwcnsis 

Kapuas 1976-14 
R hnnani 

Malay P. (CAS-SU 35615, paratypcs) 

Kapuas 1976-10 

R caudunaculala 
Malay P.. Johorc (CAS-SU 34644) 
Kapuas 1976-6 

R (YmT»(Jcrdon. 1849) 

Sri Lanka (CAS-SU 16505, 16507) 
R ccphalolaenia 

Kapuas 1976-8 

R (/fl/i;io«/(K (Hamilton-Buchanan, 1822) 
Nepal, Chitawan Valley (CAS 50344) 
Sn Lanka (CAS 52417) 
Burma, Irrawaddy (CAS 44957) 

R dor^hxcllata 
Malay P.. Pahang (CAS-SU 4728 1 ) 
Kapuas 1976-51 

R duSOIICHMS 

Malay P., Perak (CAS-SU 15344) 
Kapuas 1976-1. 14 

R cinllunvnii 

Kapuas 1976-1 
R (7i//w (Haniillon-Buchanan. 1822) 

Calculia (CAS-SU 4723^)) 
R rh'i;aii\ 

Kapuas 1976-24, 25 

Bungaran Isd. (CAS-SU 15328) 

R. cnncalcpfi 
Kapuas 1976-25 (paralypes) 
Kapuas 1976-26 (paralypes) 

R d. i'linciiU'pi-i 

Kapuas 1976-27 
R hclcrt^niorpha Duncker, 1904 

Malay P,, Johorc (CAS-SU 31203) 
R hiihhsi Briltan, 1954 

N Borneo (CAS-SU I 747X, paratypcs) 
R ;i;(ii/is()«/ Weber and dc Bcaulort, I'JHi 

Sumatra, Harau (CAS-SU 15331) 
R kalochrotna 

Kapuas 1976-23 



2-3+11-14=14(4). 15(3), 16(1) 
1-2+12-13=14(4), 15(1) 

2-4-1-10-12=12(1), 1.3(2), 15(1), 16(1) 

0-1-1-6-7=6(1), 7(1), 8(1) 

1-3 + 8-10=10(2), 1 1(7), 12-'(1) 
1-2 + 8-10 = 9(2), 10(7), 11(4), 12(3) 
2 + 9=11(1) 

2-3+10-11 = 12(2), 13(2), 14(1) 

2-3 + 9-10=11(1). 12(3), 1,3(2) 



1-3 + 7-9 = 9(3), 10(1), 11(1), 12(1) 
1+8 = 9(1) 



2-3 + 9-11 = 12(1), 13(2) 
3 + 8=11(1) 

3-5+14-17=17(3), IS(2). 19(1), 20(4). 21(1) 

2 + 9=11(1) 



2-4 + 8-12=11(1). 13(2). 14(1). 15(1) 

2-3 + 8-9=11(3) 

2-3 + 9-12=11(1), 12(2), 13(3), 15(1) 



0-3+6-9=9(3). 10(4), 12(1) 
2-3 + 8-10=11(2). 13?(l) 



3+10=13(2) 

2-3+10-11 = 12(3), 13(1), 14(1) 



1-2 + 8-9 = 9(1), 10(1). 1 1(1) 

2+10=12(1) 

:_3 + g-10=12(3). 13(1) 
2+10-11 = 12(1), 13(1) 

1+7-8=8(6), 9(3) 
1-2 + 7-8=8(3). 9(2) 

0-1+7-8 = 8(8), 9(2), 10(1) 

1-2 + 7-8 = 8(1), 9(3), 19(1) 

2-3 + 8-10=10(5), 12(2) 

2 + 7 = 9(1) 

2 + 8=10(2) 



ROBERTS- WESTERN BORNEO FISHES 



&9 



Table 2. CoNTirouED. 



R macti/ala Duncker, 1904 

Malay P., Johore (CAS-SU 39386) 
R meinkem de Beaufort, 1931 

Sumatra (CAS-SU 13333) 
R. niyersi 

Kapuas (CAS-SU 17345, paratypcs) 

Kapuas 1976-14, 24 

R. pauaperforata 
Sumatra, G. Sahilan (CAS-SU 15335, paratypcs) 
Kapuas 1976-4 

R. phihppina Gunthcr, 1 880 
Mindanao, Lanao Prov. (CAS-SU 38108) 

R. rai/iora (Hamilton-Buchanan, 1822) 
India, Uttarbagh (CAS-SU 34622) 

R. reticulata Weber and de Beaufon, 1915 

Nias Isd. (CAS-SU 15336, paratypc) 
R. rutteni Weber and de Beaufort, 1916 

Borneo, Sungei Wain (CAS-SU 15338, paratypc) 
R- sarawakensis 

Sarawak, Kuching (CAS-SU 33567, paratypcs) 

Kapuas 1976-6, 30 

R. seinilineala Wcbcr and de Beaufort, 1916 

N. Borneo (CAS-SU 47259) 
R slicnen Nichols and Pope, 1927 

Canton (CAS-SU 28748) 

Hong Kong (CAS-SU 31630) 

Hainan (CAS-SU 31799) 

R suhlilis 

Kapuas 1976-32, 40 (types) 
R Ia»arensi5 Weber and de Beaufort, 1916 

Sumatra. L. tawar (CAS-SU 15340, paratypc) 
R. trilincata 

Kapuas 1976-16, 37 

Malay P-. Johore (CAS-SU 47235) 

Thailand, Rayong R. (CAS 44434) 

R tuhbi Bnttan, 1954 

Brunei (CAS-SU 48406) 
R mophlhalma Ahl, 1922 

Saigon'' (CAS-SU 17357) 
R vd/er(/Z()n5 Deraniyagala. 1930 

Sn Lanka (CAS-SU 150(34, CAS 52414) 
R volzi 

Sarawak, Kuching (CAS-SU 33592) 

Kapuas 1976-24 



0-H-6-8 = 7-'(2), 8(1), 9(1) 

3-1-13=16(1) 

3-1-12=15(2) 
2^4-10-12=13(1), 14(3) 

1-2-1-9-10=11(2), 12(2) 
1-2-1-9-10=10(1), 11(2) 

3-4+11-12=14(2), 15(1), 16(2) 

3 + 9-10=12(1), 13(2) 

2 + 8 = 10 

1+7 = 8 

1-2 + 7-8 = 8(1), 9(2) 

1-2 + 7-10=8(4). 9(5). 10(1), 11(1), 12(1) 

1-2 + 9-10=11(6), 12(4) 



2-3 + 8-9=11(2). 12(2) 
2-3+9-11 = 12(2), 13(3) 
3+10=13(1) 



4-5+15-18=19''(l). 20(1), 21(4), 22(1) 

5+15 = 20(1) 

2-5 + 9-13=12(2), 13(3), 14(4), 15(1), 16(2) 
2-3+10=12(2), 13(1) 
3^+11-12=14(3). 15(5) 

1-3 + 8-10=10(4), 11(4). 12(8) 

1+8=9?(1) 

1-2 + 8-9=9(1). 11(1) 

2 + 9-10=11(3). 12(1) 
2+10=12(1) 



Kapuas males with a pale blue longitudinal stripe extending on 
dorsolateral portion of body, abdomen wine-red. Preserved 
males e.xhibit a dark longitudmal stripe in place of the blue 
coloration of live specimens. This stripe is absent in females, 
which are less colorful in life than males. The anterior margin 
of the anal fin may be milky-white in large males. The precise 
differences of live coloration in Kapuas males and females is 
not known, since they were not scxed while alive, and many of 
both sexes were juveniles without fully developed coloration. 
Brittan (1976:92) provides a color photograph of a live speci- 
men, presumably male, with a green or emerald green longi- 



tudinal stripe dorsolaterally and the abdomen and fins of a more 
nearly carmine-red color. 

My largest female, 19.0, apparently gravid (perhaps partially 
spent?), contained 33 eggs of about 0.7 mm. I suspect that re- 
production is year round or continuous unless interrupted by 
unfavorable conditions. 

Remarks on Relationships. — /?cK/)ora axelwdi differs strik- 
ingly from all other species assigned to Rasbora in having "per- 
manent" sexual dichromatism and extensive development of 
large, specialized tubercles in the male. So far as I am aware, 
no other Rasbora exhibit permanent or even temporary sexual 



70 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 48. Specialized tubercles on ventral surface of mandible. Above, Rasbora axelwdi. Kapuas 1976-5. 17.4 mm male (CAS 49271); below, Bmchydamo 
irniK Nepal, Reu River, 29,2 mm graMd female (CAS 50284). 



dichromatism or tuberculation comparable to thai of R. a.xel- 
loili. As sexual maturity is attained, males ofR. axelivdi develop 
increasingly bright coloration and heavy tuberculation, highly 
unusual for such a small species. Males develop relatively large 
tubercles on head, cheeks and shoulder region, those on opercle 
and overlying "pseudotympanum" in vertical rows (Fig. 47), 
The largest and most striking tubercles, however, occur on ven- 
tral surface of mandibles, on the lateral margins of two pairs of 
peculiar tubercle-bearing platforms (Fig. 48); comparable man- 
dibular tuberculiferous structures have not been observed in 
any other Rashora but are characteristic of both sexes in some 
of the smaller species of the c\prinid subfamiK Danioinae, 
including Bmchydamo nvio (Hamilton-Buchanan. 1822) (Fig. 
48). This observation suggests that Rasbora axelwdi may belong 
to Danioinae. a subfamily otherwise unknown from Borneo. 

Distribution. — K.nown with certainty only from the Mem- 
pawah basin of western Borneo (Kapuas IQ76-5). 

Rasbora bankanensis (Bleeker. 185.1) 

(Figure 44) 

Lctuiwus hiiiikiiiicnsi.s Bleeker, 185.11/192 (type localit\ Banka). 
Rd\hi>ni ItankanciiMi Bleeker. IX6(li(:454. 



Material Examined. -Western Borneo: Kjpuas 1976-7, 3: 12.9-14.0 mm (MZB 
i?<2\): Kapuas 1976-8.49; 8.0-27.3 mm (AMNH 48929. BMNH 1982.3.29.55- 
64. CAS 49272. MZB 3322); Kapuas 1976-9. 46: 30.7-45.1 mm (CAS 49273, 
FMNH 94219. IRSNB 19738, MCZ 58349, MZB 3323); Kapuas 1976-10, 58: 
10.7-49.0 mm (MNHN 1982-682, MZB 3324. RMNH 28845, ROM 38618, 
UMMZ 209854); Kapuas 1976-13, 13: 1 1.8-24.0 mm (MZB 3325. USNM 230221); 
Kapuas 1976-17. 15.2 mm (MZB 3995); Kapuas 1976-22, 6; 36. .3-18. 9 mm (MZB 
3326. I'SNM 230222); Kapuas 1976-24, 13: 24.1-48.4 mm (MZB 3327, ZMA 
116.526); Kapuas 1976-25, 23.7 mm (MZB 3328); Kapuas 1976-27, 27: 24.0- 
614 mm (CAS 49274, MZB 3329); Kapuas 1976-28, 21.0 mm (MZB 3330); 
Kapuas 1976-29, 2: 46.3-52.6 mm (CAS 49275. MZB 3331); Kapuas 1976-31, 
2: 30.8-39.0 (MZB 3332); Kapuas 1976-33, 27.9 mm (MZB 3333); Kapuas 1976- 
35.2: 31.9-34,2 mm (MZB 3334, RMNH 28846); Kapuas 1976-36,6: 26.9-42.8 
mm (AMNH 48930, MZB 3335); Kapuas 1976-37,9: 14.1-20.9 mm (CAS 49276, 
MZB 3336); Kapuas 1976-39, 11: 16.5-20.1 mm (MZB 3337, UMMZ 209899); 
Kapuas 1976-40, 2: 28.0-33.8 mm (MZB 3338); Kapuas 1976-42, 4; 14.5-26.2 
mm (MZB 3339. ROM 386051; Kapuas 1976-46, 2: 23.7-25.3 mm (MZB 33401; 
Kapuas 1976-47, 3: 16,0-19.7 mm (MZB 3.^1): Kapuas 1976-51, 24: 12.0-20.0 
mm (CAS 49277. MZB 3342). Kapuas 1976-55, 17: 18.1-23 5 mm (MZB 3343, 
USNM 230223) 

This IS the most ubiciuitous fish species in the collections 
obtained by the Kapuas survey of 1976. It is represented in 24 
of the 55 collections. A large proportion of the samples include 
small or very small juveniles, .luveniles as well as adults of this 
bluish-silvery species are distinguished by usually having black 



ROBERTS- WESTERN BORNEO FISHES 



71 




Figure 49. Rasbora bankanensis. Above, Kapuas 1976-10, 48.4 mm (MZB 3324); below, Kapuas 1976-29. 52.6 mm (CAS 49275). 



pigment on the distal end of the anteriormost enlarged anal-fin 
rays. Lateral line complete, lateral scales series 25-28; trans- 
dorsal scales 9, predorsal 10-12, usually 1 1, circumpeduncular 
12. 

Distribution. — Malay Peninsula (Perak. Johore, Muar R., 
Singapore). Sumatra (Moesi). Borneo (Kapuas). Banka (Mara- 
wang). 

Rasbora beauforti Hardenberg, 1937, species inquirenda 

Rasbora beauforti Hardenberg, 1937:10 (type locality Kumai River, southwest 

Borneo). 
Rasbora beauforti Bnttan, 1954:173 (summarued from Hardenberg 1938). 

M.^TERiAL Examined —None. 

This species is known only from Hardenberg's account; he 
did not indicate the disposition of type specimens and I have 
not been able to find them despite considerable correspondence. 
It seems clear that they are not in Holland (H. Nijssen. M. 
Boeseman, M, van Oijen. pers. comm., 1982-83) or in Bogor 
(Soetikno. pers, comm,, 1983). I have not been able to identify 
positively any specimens as R. beauforti and, if type specimens 
cannot be located, would be inclined to regard it as a nomen 
dubium. The original description (no figures) reads as follows; 

D. 1.8; A. 2. 5.: P. 1.1 2; V.1.7; L.r. 28-29; L.l. mcomplete, consistmg ot^ 
10 scales only; L.v. (before ventrals) 4-1/2-1-2-1/2. 
Oblong. Height about 4 in length. 5 in length with caudal. Head 



about one m height. Eye 3 in head, about equal to snout. Cleft of 
mouth rather strongly descendmg. not reaching vertical through front- 
border of eye. Origin of dorsal behind the middle between end of snout 
and origin of caudal, opposite to end of incomplete lateral line, 1 2 
scales from occiput. Dorsal nearer to ventrals than to anal, its height 
somewhat shorter than head. Pectorals as long as head without snout, 
ventrals somewhat shorter. Longest ray of anal as long as postorbital 
part of head and half eye. Longest ray of caudal about as long as head. 
Caudal peduncle surrounded by 12 scales. Colouration of formolspe- 
cies [sic] dark, brownish above, much lighter below. A conspicuous 
dark band along the sides, beginning on tip of snout and ending on 
caudal, running through the eye. This band is narrowest on the head 
and on the caudal fin. The black band is separated from the brownish 
back by a light streak. The first 12-13 scales in this streak have a 
blackish hindborder. Fms more or less pigmented, especially the dorsal 
and the caudal. Some specimens have the tip of the ventrals and of 
the anal blackish. 

Many specimens from the Kumai-nver. south-west Borneo. May 
1931. Longest specimen 44 mm. Named in honour of Prof. Dr. L. F. 
de Beaufort from Amsterdam. 

Rasbora borneensis Bleeker, 1 860 

(Figure 50) 

Rashora borneensis Bleeker. 1860(j:450 (type locality Bandjermassin in fluviis). 
Genus and species undet. Kottelat, 1982:434. fig. 56 (Mentaya basin). 

Material Examined —Southern Borneo: Bandjermassin. 48.9 mm (BMNH 
1866.5.2.1 55. syntypc. 62.5 mm total length) and RMNH 7497. 56.4 mm (RMNH 
7497. syntype. 74.0 mm total length); Kuala Kuaya. Sampitarea, Mentaya basin, 
72.6 mm (CAS 53054). Western Borneo: Kapuas 1976-14, 21: 13.0-53.5 mm 
(BMNH 1982.3.29.65-68. CAS 49278, MZB 3344); Kapuas 1976-16, 52.2 mm 



72 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 50. Rashora horneensis- Kapuas 1976-31. 50.3 mm (MZB 3346). 



(MZB 3345), Kapuas 1976-31. 50.3 mm (MZB 3346); Kapuas 1976-33, 72.8 mm 
(MZB 3347); Kapuas 1976-40. 25.4 mm (MZB 3348); Kapuas 1976-45, 7; 29.7- 
71.0 mm (MZB 3349. RMNH 28847, USNM 230224); Kapuas 1976-50. 40.5 
mm (MZB 3350); Kapuas 1976-55, 6: 1 7.0-23.6 mm (MZB 335 1. UMMZ 209930), 

This very distinctive species, known until now only from two 
syntypes obtained at Bandjermasin, has not been recognized 
by any ichthyologist subsequent to Bleeker. Weber and de Beau- 
fort (1916:61) placed it as a synonym of Rashnni argyrot acuta 
without comment, as did Brittan (1954:108) but with a question 
mark. It is apparently related to R. argyrotacnia and R. lepto- 
souia but differs from them and their other relatives in having 
a relatively small head and distinctive coloration. 

Head length 4.2-4.5 (vs. 3,4 in R. argyroiaciua). Lateral line 
complete, scales in lateral series 30-33. Predorsal scales 15-16, 
transdorsal 9, curcumpeduncular 14. Pale straw-colored in life. 
Narrow middorsal stripe from occiput to caudal fin (wider in 
R. aygyrotacuia). Thin, straight longitudinal dark streak over- 
lying or just above horizontal septum; broad longitudinal stripe 
faint, most evident on posterior half of body. Scales on dorsal 
and dorsolateral positions of body densely covered with ex- 
tremely fine, numerous melanophores except distally. where 
there is a submarginal clear area followed by a marginal band 
of large melanophores (as in R. arg)'wtae)ua). Large melano- 
phores on gill cover and on ventrolateral portion of anterior 
half of body form the most noticeable feature of coloration in 
these areas. Distal margin of caudal tin dusky; fins otherwise 
colorless. Some specimens (males?) with scattered small tuber- 
cles extensively distributed on dorsolateral portion of body pos- 
terior to dorsal fin. Total gill rakers on first arch 12-13. 

A description and photograph of the 72.6 mm specimen from 
Mentaya is given by Kottelat (1982). This specimen is similar 
to Kapuas specimens in most respects, but melanophores on its 
gill cover and ventrolateral portion of anterior half of body are 
much finer, and broad longitudinal stripe, although faint, is 
better developed and more evident on anterior half of body. 

Distribution. — Known only from Borneo (Kapuas, Men- 
taya, Barito). 

Rasbora brittani Axelrod, 1976 

Rashora bnttani Axelrod. 1976:94 (lype localily Johorc R)vcr). 

Material Examined. — Malay Penmsula; Johorc River, 7: 31-36 mm (CAS 
35615, paralypcs). Western Borneo: Kapuas 1976-7, 11; 9.9-14,7 mm (BMNH 
1982.3.29.69-75. MZB 3352); Kapuas 1976-10, 26: 1 1 .7-15.3 mm (CAS 49279, 
FMNH 94220, MZB 3353. USNM 230225); Kapuas 1976-37. 23 2 mm (MZB 



3354); Kapuas 1976-39. 2: 16.5-17.6 mm (MZB 3355. RMNH 28848); Kapuas 
1976-42. 16.1 mm (MZB 3996); Kapuas 1976-43.5: 16.6-22.4 mm (MZB 3356. 
IIMMZ 209910). 

A relatively small Rasbora. attaining 50 mm but most spec- 
imens smaller, rather similar to R. argyrotaenia in body shape 
and fin conformation, with pointed head, moderately slender 
body and caudal peduncle. Lateral line incomplete, with about 
nine pored scales anteriorly; lateral scales 31-34, predorsal 15, 
circumpeduncular 12. No lateral stripe. A small round spot m 
middle of caudal peduncle (absent in all other Kapuas Rasbora). 
In live specimens this spot may have bright orange or reddish- 
orange areas above and below it (Axelrod 1976:96-98); such 
coloration was not observed in Kapuas specimens, most of which 
came from siltv water and may have been blanched. 



Rasbora caudimaculata Volz, 1903 

Rasbora caudiniaviilala Vol/. 1903:559 (lype locality Scmangus or upper Moesi 

River, Palcmbang Residency. Sumatra). 
Rasbora doi-iimaculala Herrc. 1940:9 (type locality hrook 16 miles easi Kuching, 

Sarawak). 
For additional synonymy sec Brillan (1954. 1972). 

Material Examined. — Malay Peninsula: Johorc, Mawai Dist.. 5; 33.4-85.8 
mm (CAS-SLI 34644); Sarawak: brook 16 km E of Kuching. 27.2 mm (CAS SU 
33021, hololypc of R dor'^inuiciilalay. western Borneo: Kapuas 1976-6, 101 mm 
(MZB 3357), 

Some juveniles at about 30 mm exhibit a black mark at the 
tip of the longest dorsal-fin rays not seen in large specimens. 

Distribution.— Thailand? (see Brittan 1954, 1972). Malay 
Peninsula (Johore). Sumatra (Moesi). Borneo (Sarawak, Ka- 
puas). 

Rasbora cephalotaenia (Bleeker, 1852) 

LcKCisciisccphalolacnia Bleeker. 1852c/:97 (type locality Tjirutjup River, Billiton). 
Rasbora cephalotaenia Bleeker. 18606(:438. 

Rasbora lornien Ahl. 1922:32 (type localitN central Sumatra). Sec Brittan (1954, 
1972) 

Material Examined, — Western Borneo: Kapuas 1976-1, 60 6 mm (MZB 3358); 
Kapuas 1976-8, 2: 63, .3-67. 2 mm (CKS 49280, MZB 3359) 

A moderately elongate, heavy-bodied Rasbora with complete 
lateral line, lateral scale series 30-32, predorsal scales 12-13, 
total scales above lateral line 7, circumpeduncular scales 12. 
Attains at least 93 mm. Distinguished from all other species by 
its very distinctive color pattern of four punctuated longitudinal 



ROBERTS-WESTERN BORNEO FISHES 



73 




Figure 51. Rashora cnncatepis- Kapuas 1976-26, 38.0 mm {MZB 3997, hololype). 



Stripes, two dark and two light, forming by spots on scales. The 
two dark stripes, nearly midlateral, are continued anteriorly and 
posteriorly as a single solid stripe extending from snout-tip 
through eye and across gill cover, and on middle caudal-fin rays 
to end of caudal fin. A vertically elongate oval peduncular spot 
straddles the dark stripes where they unite at the caudal-fin base. 
The upper punctuated light stripe extends from the upper comer 
of the gill cover to the base of the uppermost principal caudal- 
fin rays. The lower light stripe extends from pectoral-fin base 
to base of last anal-fin ray. 

Distribution. — Malay Peninsula (Singapore, Johore). Su- 
matra (Gunung Sahilan, Taluk, Semangus R., Palembang, Moe- 
si. Borneo (Baram, Kapuas). Banka. Billiton. 

Rasbora dorsiocellata Duncker, 1 904 

Rasbora liorsiocellaui Duncker. 1904:183, pi. 1. fig. 2 (type locality Malay Pen- 
insula). 

Material Examined. — Malay Peninsula: Pahang. Tasek Bera. 11: 28.5-36.7 
mm (CKS-SU 47281 ); western Borneo: Kapuas 1976-16, 2: 21.6-24.4 mm (AMNH 
48931. MZB 33601; Kapuas 1976-32. 24: 14.3-21.3 mm (BMNH 1982.3.29.76- 
78. FMNH 94221. ZMH 6475. MNHN 1982-683, MZB 3361, USNM 230226); 
Kapuas 1976-42, 8: 17.1-22.4 mm (MZB 3362, UMMZ 209906); Kapuas 1976- 
43, 25.2 mm (MZB 3363); Kapuas 1976-46, 5: 1 7.8-24.6 mm (MZB 3364, RMNH 
28849); Kapuas 1976-47, 2: 21.2-22.7 mm (MZB 3365. ROM 38620); Kapuas 
1976-51. 11: 18.1-28.8 mm (CAS 49281, MZB 3366). 

A small Rasbora distinguished by a dark round spot in the 
middle of its dorsal fin. Lateral line highly variable, from about 
six perforated scales only to complete (Brittan 1954. 1972). 
Lateral line scales 25-30, transdorsal 7. predorsal 12, circum- 
peduncular 12. A .10.8 mm presumed male from Tasek Bera is 
markedly tuberculate. with numerous fine tubercles on posterior 
half of opercle and on ventral surface of branchiostegal mem- 
branes. 

Distribution, — Malay Peninsula (Pahang, Muar, Negri Sem- 
bilan, Johore). Borneo (Kapuas). 

Rasbora dusonensis Bleeker, 1851 

Rashnra dusonensis Bleeker, 1851((;14 (type locality Duson or Barito River at 
Bandjcrmassing). 

Material Examined. —Sarawak: Rcjang basin. SNHM, numerous uncata- 
logued specimens recently collected by Anthony Leiek; western Borneo: Kapuas 



1976-1. 10: .30.1-50.2 mm (BMNH 1982.3.29.79-80, CAS 49282, MZB 3367, 
ROM 38606); Kapuas 1976-2. 19,3 mm (MZB 3368); Kapuas 1976-3. 4: 16.9- 
29.5 mm (FMNH 94222. MZB 3369); Kapuas 1976-14, 6: 56.6-86.8 mm (CAS 
49283, MNHN 1982-684, MZB 3370, RMNH 28850, UMMZ 209856); Kapuas 
1 976- 15,71.9mm (MZB 3367 1 ); Kapuas 1976-51,3: 72.4-88.6 mm (MZB 3372, 
USNM 230227). 

A large Rasbora with subtle but distinctive coloration per- 
mitting identification of juveniles as well as adults: thin midaxial 
streak very distinct on posterior half of body; a sharply delim- 
ited dusky midlateral longitudinal stripe of fine melanophores 
extending uninterruptedly from posterior margin of eye and 
across gill cover to caudal-fin base; a sharply delimited pale 
longitudinal stripe of about equal width immediately dorsal to 
dusky midlateral stripe. Fins clear or dusky, without discrete 
marks. Lateral line complete, lateral scale series 33-35; trans- 
dorsal scales 9. predorsal 13-15. circumpeduncular 14. 

Distribution. — Malay Peninsula (Perak). Sumatra (Moesi). 
Borneo (Rejang. Kapuas, Barito). Thailand? (see Brittan 1954, 
1972:122). 

Rasbora einthovenii (Bleeker. 1851) 

Leuctscus einlhovenn Bleeker, 1851/:434 (type locality Sambas). 
Rashora einthovenii Bleeker. 1859t7:154. 

Material E.xamined. — Western Borneo: Kapuas 1976-1, 13: 15.9^0.1 mm 
(CAS 49284. MZB 3373, ROM 38609); Kapuas 1976-11,9:3 1.34-42.3 mm (CAS 
49285. MZB 3374, RMNH 28851, USNM 2.30228). 

Distinguished from all other species of Rasbora by a solid 
dark longitudinal stripe from snout-tip to end of middle caudal- 
fin rays, with portion on body and especially on caudal peduncle 
lying just below midline of body. 

Distribution. — Malay Peninsula (Johore. Singapore). Su- 
matra (Bagan Api Api, Gunung Sahilan, Siboga). Borneo (Sam- 
bas, Kapuas, Bungaran, Brunei, Labuan, Sandakan). 

Rasbora elegans Volz, 1 903 

Rashora elegans Volz. 1903:558 (type locality Palembang. Sumatra). 
Rashora lalerislnala Weber and de Beaufort. 1916:78. 

Material Examined. —Western Borneo: Kapuas 1976-24, 2: 67.2-77.7 mm 
(CAS 49286, MZB 3375); Kapuas 1 976-25. 10: 3 1 .6-85.6 mm (CAS 49287, MHNG 
2087.3-5, MZB 3376); Kapuas 1976-26, 4: 37.2-76.3 mm (MZB 3377, USNM 
230229). 



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CALIFORNIA ACADEMY OF SCIENCES 



Rasbora ennealepis new species 

(Figure 51) 

HoLOTVPE-MZB 3''Q7, 38.0 mm. Sungai Sekumpai. a small forest iributan 
oflhe Sungai Pinoh 23 km S of Nangapinoh (Kapuas 1976-26). 

P.^RATYPES.-CAS 49288 and MZB 3378, 5: 26.9-38.2 mm, same data as ho- 
lotype, BMNH 1982,3.29,81-82, CAS 49289, MZB 3379, RMNH 28852, USNM 
230230, 1 4: 26.2-44.9 mm. Sungai Tebelian, small forest tributar\ of Sungai Pinoh 
19 km S of Nangapinoh (Kapuas 1976-25) 

This species is similar to R. bankanensis. irom which it differs 
in having only 8-9 instead of 9-12 gill rakers (Table 2) and in 
details of coloration and squamation. Lateral line complete, 
lateral scale rows 24-25 (vs. 25-28 in R. bankanensis); trans- 
dorsal scales 9, predorsal usually 9, rarely 10 (vs. 10-12, usually 
11), circumpeduncular 10 (vs. 12); scales with striae few and 
broadly divergent (vs. scales with striae relatively numerous and 
nearly parallel). Sides of body with strongly reticulated color 
pattern (absent or weakly evident in R. bankanensis); anal fin 
without a noticeable concentration of dark pigment near distal 
end of enlarged anterior fin ra\s (usually present in R. banka- 
nensis): and striae of scales pigmented (striae without pigmen- 
tation in R. bankanensis). 

Rasbora ennealepis is slightly less elongate than R. banka- 
nensis. with a slightly narrower caudal peduncle and perhaps a 
slightly larger eye, although no attempt has been made to quan- 
tify these diflerences. The two species are also similar in color 
pattern except for the differences pointed out above. Like R. 
bankanensis. R. ennealepis has a complete midaxial streak, a 
broad midlateral stripe better defined on posterior half of body, 
a well defined, complete dorsomedial streak or stripe, a su- 
praanal streak continuous with a midventral peduncular streak, 
and melanophores along cleithrum somewhat concentrated to 
form a poorly defined humeral mark underhing posterior mar- 
gin of gill cover. 

In addition to the type specimens listed above, one lot of 13 
specimens 31.9-47.2 mm from Kapuas 1976-26 (CAS 49290, 
MZB 3380) apparently belongs to R. ennealepis. These tend to 
have 1 or 1 1 predorsal scales and lighter coloration but in other 
respects are similar to the type specimens. Consideration has 
been given to the possibility that the difl'erences seen between 
R. ennealepis and R bankanensis merely reflect ecophenotypic 
variation of R. bankanensis. This hypothesis is weakened by 
the observation that R bankanensis from mountain tributaries 
of the Sungai Pinoh do seem to differ in intensity of coloration 
and in subtle alterations of body form in ways that may indeed 
reflect ecophenotypic variation but are readily distinguishable 
from all of the specimens of R. ennealepis which is known only 
from mountain tributaries of the Sungai Pinoh. 

Etymology. — The name ennealepis (Greek) refers to the 
characteristic number of predorsal scales seen in this species. 

Rasbora kalochroma (Bleeker, 1850) 

U-iici^ciis kalochrinua Bleeker. 1850-272 (l\pe loealih Bandjermassing). 
Rushoyu kiilnchmina Bleeker. I859i(:154. 

Material Examined. — Western Borneo; Kapuas 1976-23, 5: 28 1-39,5 mm 
(C AS 49291, MZB 3381, USNM 230231) 

Two large dark blotches on side of body, one above pectoral 
fin and one centered over anal-fin origin, distinguish this species 
from all other Rasbora. Dorsal-fin origin slightly posterior to a 
vertical through anal-fin origin. Lateral line complete, lateral 



line scale series 29-32; predorsal scales 11-12; circumpedun- 
cular scales 12. Largest known specimen 59 mm. 

Distribution. — Malay Peninsula (Johore, Trengganu). Su- 
matra (Gunung Sahilan, rivers Kamperkiri, Kwantan, Rokan). 
Borneo (Kuching. Sadong, Sambas, Kapuas). Banka. Billiton. 

Rasbora myersi Brittan, 1954 

(Figure 52) 

Ra^hnra nnvrsi Briltan, 1954:117, fig, 25 (type locality Kapuas River at Potoes 

Sibaii), 
For synonymy see Briltan (1954, 1972:1 17-1 18), 

Material Examined, — Western Borneo: Kapuas River at Poctoes Sibau, CAS- 
SU 17342, 71 mm (holotype) and CAS-SU 17343. 3: 66-70 mm (paratypes); 
Kapuas 1976-6. 85.0 mm (MZB 3998); Kapuas 1976-14,4: 5 1,6-74,3 mm (CAS 
49292, MZB 3382); Kapuas 1976-15, 11: 34.5-71.5 mm (AMNH 48932. BMNH 
1982.3.29.83-86. MZB 3383); Kapuas 1976-24. 4: 75.4-81.8 mm (CAS 49293, 
MZB 3384, ROM 38623); Kapuas 1976-31, 8: 50,3-72,3 mm (FMNH 94223, 
IRSNB 19739. MZB 3385); Kapuas 1976-32. 2; 49.7-58.9 mm (CAS 49294. MZB 
33S6); Kapuas 1976-33. 6: 66.5-72.2 mm (MCZ 58350. MZB 3387): Kapuas 
I976-.34. 54.2 mm (MZB 3388); Kapuas 1976-36. 4: 39.6-65.5 mm (MZB 3389. 
MNHN 1982-685): Kapuas 1976-37. 15: 41.9-83.5 mm (KUMF 2850. MZB 

3390, RMNH 28853, ZMA 116,527): Kapuas 1976-39, 11: 50.0-79.5 mm (MZB 

3391, L'MMZ 209900, USNM 230232). 

Rasbora cf myersi Brittan, 1954 (immature) 

Material Examined. — Western Borneo: Kapuas 1976-15. 233: 8.7-16.9 mm 
(CAS 49309, MZB 3425); Kapuas 1976-33, 16; 7.8-19.7 mm (CAS 49310, MZB 
3426): Kapuas 1976-55, 7; 13.4-13.9 mm (MZB 3427, USNM 230239), 

Material obtained during the Kapuas survey of 1976 includes 
three lots of small, immature Rasbora. apparently conspecific, 
which might be R. myersi. The largest of these specimens, 19.7 
mm, apparently has a complete lateral line, and about 14-15 
predorsal scales; its color pattern is generalized and rather plain, 
with midlateral stripe and axial streak complete, scales on dorsal 
portion of body with melanophores on exposed portions form- 
ing a reticulate pattern, and no distinctive or unusual markings. 
Kapuas species most likely to be the adults include R. argyro- 
tacnia. caiuliinaculata. dusonensis. elegans. myersi. and volzi. I 
tentatively identify them as R. myersi. of which the smallest 
specimen otherwise examined is 34.5 mm, because of close 
similarity in overall morphology and color pattern. 

Rasbora pauciperforata Weber and de Beaufort, 1916 

Riiihiiiv paucipcrliirala Weber and de Beaufort, 1916:78, lig. 28 (type locality 
Sumatra. Gunung Sahilan and Deli) 

Material Examined —Sumatra: Gunung Sahilan. 4: 25.4-27.6 mm (CAS-SU 
15335. paratypes). Western Borneo: Kapuas 1976-1. 29: 25.5-32.5 mm (.AMNH 
48933, BMNH 1982,3,29,87-97. MZB 3392); Kapuas 1976-6. 19: 14.1-34,1 mm 
(CAS 49295, MZB 3393); Kapuas 1976-5, 29: 12.0-34,2 mm (FMNH 94224, 
IRSNB 1 9740, KIIMF 2851, MZB 3394); Kapuas 1976-7, 23: 8,6-22.5 mm (CAS 
49296, MZB 3395): Kapuas 1976-10, 24.4 mm (MZB 3396); Kapuas 1976-11. 
20: 20.6-34.0 mm (MZB 3397. MNHN 1982-686. RMNH 28854. ROM 38613); 
Kapuas 1976-23. 26.6 mm (MZB 3398); Kapuas 1976-32, 65; 15 3-22,9 mm 
(CAS 49297, MZB 3399, UMMZ 209880, USNM 230233, ZMA 1 16.528). 

A small, slender, colorful Rasbora with a sharp snout, narrow 
caudal peduncle, and two well marked longitudinal stripes. Lat- 
eral line incomplete, with 5-10 pored scales. Lateral scale series 
30-34; transdorsal scales 7, predorsal 13-14, circumpeduncular 
12. Gill rakers 10-12. Largest specimen 34.1 mm. 

Distribution. — Malay Peninsula (.lohore). Sumatra (Gunung 
Sahilan, Deli). Western Borneo (Kapuas). Billiton. 



ROBERTS-WESTERN BORNEO FISHES 



75 




Figure 52. Rasbora myersi. Kapuas 1976-37, 58.7 mm {MZB 3390). 



Rasbora sarawakensis, Brittan. 1951 

(Figure 53) 

Rmhoia saranakensis Bntlan. 1951:1 (lype locality brook lb miles east of Ku- 
ching). 

Material Examined. — Sarawak: brook lb miles east ol Kuching, 10: 18-38 
mm (CAS-SU 15375. 33567. holotype and paratypes): Rejang River, 3: 34-38 
mm (CAS-SU 33593, paratypes). Western Borneo: Kapuas 1976-6, 37: 17.7-42.4 
mm (AMNH 48934, BMNH 1982.3.29,98-101, CAS 49298, FMNH 94225, IRSNB 
19741, MZB 3400); Kapuas 1 976- 12, 7: 12.6-38.4 mm (MZB 3401, MNHN 1982- 
687): Kapuas 1976-13, 3: 14.0-19.7 mm (CAS 49299, MZB 3402): Kapuas 1976- 
24, 43.8 mm (MZB 3403); Kapuas 1976-25, 6: 26,2-36.0 mm (MZB 3404, RMNH 
28855): Kapuas 1976-26, 7: 19,0-42.5 mm (MZB 3405, ROM 38600, UMMZ 
209868); Kapuas 1976-30, 28: 19,0-47,5 mm (CAS 49300, MZB 3406, UMMZ 
209873, USNM 230234. ZMA 1 16.529). 

A relatively stout-bodie(i Rasbora with a large, pointed head, 
distinguished from all other species by its distinctive very in- 
tense lateral longitudinal stripe, basally darkened dorsal fin, and 
scale markings. It differs from all other species except R. rutteni 
Weber and de Beaufort, 1916 in having numerous close-set 
multicellular homy tubercles (so-called breeding tubercles) ex- 
lending in a narrow dorsomedial band on the posterior portion 
of caudal peduncle and dorsal margins of upper procurrent and 
uppermost principal caudal-fin rays. These tubercles are present 
in all of the material examined, although they are best developed 
in the largest specimens (female as well as male). Lateral line 
complete, lateral line scales 24-26; predorsal 1 1; circumpedun- 
cular 12, 

It IS apparently very closely related to R. nttteni Weber and 
de Beaufort, 1916, which differs from it in having a relatively 
rounded snout, 26-29 lateral line scales, and the dark lateral 
stripe markedly lighter anteriorly (rather than of equal intensity 
for its entire length) (Brittan 1951:3), 

Distribution. — Sarawak (Rejang basin; Kuching), Western 
Borneo (Kapuas). 

Rasbora subtilis new species 

(Figure 54) 

Holotype, — MZB 3407, 39.9 mm, ripening female, Danau Piam near Ketun- 
gau, 38 km NNE ol'Sintang, 29 July 1976 (Kapuas 1976-32). 

Paratypes. -CAS 49301, 36.0 mm, collected with holotype; CAS 49302, MZB 
3408, LISNM 230235, 5: 27,9-34.3 mm, Kapuas mainstream near Telokabik, 29 
km W of Putussibau. 1 1 August 1976 (Kapuas 1976-40). 



Diagnosis.— A slender, compressed Rasbora with incomplete 
lateral line and a diagonal black bar near tip of each caudal-fin 
lobe; gill rakers and scales exceptionally numerous. Largest 
known specimen 39.9 mm. Gill rakers 1 9-22 (more than in any 
other Rasbora except a few relatively large, heavy-bodied species; 
Table 2). Scales in lateral series about 34-36. pored scales 8- 
10, predorsal 14-17. transdorsal 13 (9-11 in nearly all other 
Rasbora), circumpeduncular 14-16 (fewer in nearly all other 
Rasbora). Pharyngeal teeth elongate, slender, nearly straight, 
and conical or slightly uncinate, apparently 2,3,5/5,3,2. 

The following observations are based on the 39.9 mm ho- 
lotype and 36.0 mm paratype from Danau Piam (paratypes from 
Kapuas mainstream in poor condition, apparently emaciated): 
head length 3.7-3.8, width at middle of gill covers 8.2-8.6; snout 
13,8-14,2; eye 12,0-12,1; bony interorbital width 12,5-12,9; 
body depth at dorsal-fin origin 5.9-6.1, width 1 1.2-1 1.4; dorsal- 
fin origin nearer caudal-fin base than snout-tip. predorsal length 
1.9-2.0; pelvic-fin origin somewhat anterior to vertical through 
dorsal-fin origin, prepelvic length 2,1; height of last simple dor- 
sal-fin ray 4.6-4,8; pectoral fin failing to reach pelvic-fin origin 
by a distance about equal to one-half eye diameter, pectoral fin 
length 4.9-5,3; pelvic fin failing to reach anal-fin origin by about 
same distance or somewhat less, pelvic fin length 5,7-6,2; caudal 
fin deeply forked, length upper lobe 3.1, 

Color in life, observed at Danau Piam: "body bluish green, 
caudal fin yellow with a diagonal black bar near tip of each 
lobe." Preserved specimens have body lightly pigmented, with 
a narrow midlateral stripe and nearly coextensive axial streak 
on posteriormost one-half or two-thirds; midlateral stripe ter- 
minating in a small, horizontally elongate, basicaudal spot, 
sometimes very poorly defined; scales on dorsal half of body 
with fine melanophores forming a faint reticulate pattern; dor- 
somedian longitudinal stripe absent or nearly absent, perhaps 
represented by a few moderately large and irregularly spaced 
melanophores; middle of abdomen with three or four longitu- 
dinal rows of scattered, moderately large melanophores; base of 
anal fin with large melanophores forming a nearly coextensive 
anal bar, continuous with ventromedian longitudinal stripe on 
caudal peduncle; all fins except caudal colorless, 

Rasbora subtilis resembles R. irilineata and they presumably 
are closely related. R. Irilineata is somewhat heavier-bodied, 



76 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 5.V Rashira idraHakciisis Kapuas 1976-24. 43.8 mm (MZB .W04). 



with fewer gill rakers and scales, and a well developed dorso- 
median stripe. 

Distribution. — Known only from the Kapuas. 

Etymology.— The name subtilis refers to the slender form 
and delicate coloration of this species. 

Rasbora trilineata Steindachner, 1870 

Rinhi'ici liihncaia Steindachner. 1870:67.3 (type lucality Pengulon Pane. Johorc; 
note pi. .1. tig. .3) 

Rashora slii;maliira Fowler. l'>34> 341. Iig, 5 (t\pe locality Krat. southeast Thai- 
land). 

Material Examined. — Thailand: Ra\ong River. 24: 1 y( 1-32.4 mm (CAS 44434). 
Malay Peninsula: .lohore, Kota Tinggi. 3: 46.8-53 8 mm (CAS-SU 47235); Johore. 
A\er Hitam. 36,0 mm (CAS-SU 15343); .lohore. Pengulon Patic (NMW 51458- 
61. including synlypcs oi R Irtlincaia. see discussion below). Western Borneo: 
Kapuas l'»76-Q. 13: 43.2-57.8 mm (AMNH 48935, BMNH 1982.3.29.102-106, 
MZB 34 1 I ); Kapuas 1976-16. 5: 30.7-39.7 mm (CAS 49304. MZB 34 1 2); Kapuas 
1976-31. 3: 46.1-58.8 mm (FMNH 94226. MZB 3413); Kapuas 1976-37, 33: 
34.0-55.5 mm (CAS 49305, IRSNB 19742. KUMF 2852. MNHN 1982-688, 
MZB 3414); Kapuas 1976-39, II: 35.5-58,7 mm (MZB 3415, RMNH 28856, 
ROM 38604); Kapuas 1976-42, 4: 36.-3-56.5 mm (MZB 3416, UMMZ 209907); 
Kapuas 1976-46. 7: 33.7-45.9 mm (MZB 3417. USNM 230236); Kapuas 1976- 
47. 2: 35,5-40.6 mm (MZB 3418. ZMA I 16.530) 

A relatively light-bodied.sicndcr /?a.s7'(ira with matching dark 



diagonal submarginal bars toward end of upper and lower cau- 
dal-fin lobes; in life these bars often accentuated by milk-white 
pigmentation. Supra-anal pigment heavy elongate streak lying 
along entire anal-fin base with scale and fin-ray counts similar 
to heavier-bodied species such as R. caudimaculata. Lateral line 
complete. Lateral scale series 29-32, transdorsal scales 9, pre- 
dorsal 12-13, circumpeduncular 12, Gill rakers on first arch 
3-1-12=15. Largest known specimens 60 mm, 

Syntypes of/?, tnlincata are not based upon the same species. 
The description seems to be based mainly or entirely on R. 
trilincala auctorum. but the figure is of a color variety of R. 
suniatrana (similar to the variety from Sungei Gunong Goa. 
Perak; Brittan 1954, fig. If). The syntypes have not been crit- 
ically re-examined since their original description, and this has 
led to doubts as to the identity of the species described by 
Steindachner (Brittan 1954:83), The NMW possesses four lots 
(NMW 51458-61) labeled as syntypes oi R. trilineata. These 
lots were located by Dr. R. Hacker and Mr. H, Ahnelt for me 
to examine during a visit to Vienna in September 1982, My 
identifications are as follows: 

NMW 51458, 12: 24,3-35.5 mm (all in good condi- 
tion )=0.vi'g(?i7(7'. 

NMW 51459. 35: 15,0-33.5 mm, separated into two sublets: 




Figure 54. Rasbora suhlilis Kapuas 1976-32, 39,9 mm ripe female (MZB 34U7, holotype). 



ROBERTS-WESTERN BORNEO FISHES 




Figure 55. Rasbora voizi. Kapuas 1976-24, 75.6 mm (CAS 49306). 



a) 24: 15.0-31.3 mm (in a vial, all badly dried, twisted and stuck 
together, apparently never sorted or studied)=several Thynni- 
chthys. four R. sitinatrana. four R. trilineata. one Aplocheilus. 
and b) 1 1; 20.0-33.5 mm (mounted on a glass plate, but now 
all badly dried). About six are R. trilineata. some still showing 
black submarginal bars with milky-white on the caudal fin. These 
specimens must have been in good condition orginally or they 
would not have been mounted so carefully; Steindachner's de- 
scription of the diagnostic caudal-fin coloration presumably was 
based on these specimens. Of the five other specimens of this 
mounted series, at least one has a well marked peduncular spot 
and seems to be R. siimatrana. 

NMW 51460, 7: 38.8-46.5 mm=R. suinatrana and 1: 42.2 
mm=/?. trilineata. All of the R. siimatrana are relatively deep- 
bodied with a peduncular spot. The 42.2 mm specimen oi R. 
trilineata has the slender narrow body typical of R. trilineata 
and lacks the peduncular spot. It is in good condition except 
that the distal two-thirds of its caudal fin is broken off. and thus 
the caudal-fin coloration diagnostic for the species cannot be 
observed. 

NMW 5 1 46 1 , 3: 49.8-70. 1 mm=R. siimatrana. All with round 
peduncular spot; largest with black coloration extending to tip 
of caudal-fin lobes, obviously not the caudal-fin coloration de- 
scribed by Steindachner. 

In conclusion, of the four presumed lots of syntypes of R. 
trilineata. only NMW 5 1459-60 include R. trilineata auctorum. 
NMW 51458, containing only Oxygaster. and NMW 5 1 46 1 , «. 
suinatrana. should no longer be considered syntypes of R. tri- 
lineata. A lectotype could be designated from NMW 51459 or 
51460, but due to the poor condition of the specimens there 
seems to be little advantage in choosing one of them for this 
purpose. Should future studies indicate a need for type selection, 
the wisest course probably would be to designate a neotype from 
a well preserved series of fresh specimens collected at or near 
the type locality. For the time being I can only affirm that Stein- 
dachner's syntypes (NMW 51459-60) include specimens of/?. 
trilineata sensu Brittan, 1954. Under the present ICZN, how- 
ever, the correct procedure would be to select a lectotype from 
the type series. 

Rasbora volzl Popta, 1905 

(Figure 55) 

Rasbora rolzi Popla, 1905: 1 75 (type locality Bongan and Howong rivers, Kapuas 
and Mahakam basms). 



Rashora mlzivav. fascial a Popta. 1 905: 176 (type locality Kajan River). Sec Brittan 
(1954, 1972:61-73). 

Material Examined. —.Sarawak: 16 miles E of Kuching, 12: 37,0-72.0 mm 
(CAS-SU 33592). Western Borneo: Kapuas 1976-24, 2: 70,3-74.2 mm (CAS 49306, 
MZB .3419); Kapuas 1976-25, 2: 49,7-50.3 mm (MZB 3420, USNM 230237); 
Kapuas 1976-30, 3: 25.6-88,7 mm (CAS 49308, MZB 3421). 

The two smallest Kapuas specimens. Kapuas 1976-30, 25.6- 
25.7 mm, differ from other Kapuas specimens examined in 
having midlateral stripe and bar on anal-fin base very dark and 
well defined; midlateral stripe extending uninterrupted full length 
of body, notably expanded ventrally in anterior half. Such pat- 
tern is observed in adults as well as juveniles of/?, volzi (or R. 
voIzi fasciata) from Kajan River, east Borneo (Popta 1906, pl. 
9, fig. 34) and Sarawak (present observations; Brittan 1954, 
1972:61-63, fig. 8), although midlateral stripe tends to be in- 
terrupted in posterior half of body. In larger Kapuas specimens 
examined midlateral stripe and anal bar very faint; at least in 
74.2 mm specimen from Kapuas 1976-24 (CAS 49306), lateral 
stripe uninterrupted, and slightly but distinctly expanded ven- 
trally in anterior half of body. 

Rasbora species undetermined 

Rasbora leplosoma {nee Blcckerl Vaillanl, 1 902:89 (Kapuas, in part, MNHN 1891- 
376 only). 

MaterialExamined. — Western Borneo: Kapuas. Sebroeang, 46,5 mm (MNHN 
1891-376, onginally identified as R Icplosoitia. subsequently reidentificd as R. 
pauiipcrforata). 

As indicated above, the true R. leptosoma is known only from 
Bleeker's type specimens from Lahat, Sumatra. Most of the 
Kapuas specimens identified as /?. leptosoma by Vaillant have 
been identified as other species of Rashora. but the present 
specimen is an exception. It may be briefly described as follows: 
lateral line incomplete, each side of body with about nine pore- 
bearing scales; lateral scale series about 34, predorsal scales 1 5, 
transdorsal 1 1, circumpeduncular 12; head length 3.5; eye di- 
ameter 1 1.6, body depth 4.7, caudal peduncle depth 9.5 (much 
deeper than in R. paiiciperforata): dark lateral stripe (badly fad- 
ed) continuous from opercle to caudal-fin base; axial streak 
extending from middle of body to end of hypural plate. 

Rasborichthys Bleeker. 1859 

/?aji/)(im7;(/;r,sBleeker. 1 859(i: 155 (type species /-<'i/a5a(ir//f7//7c/;/; Bleeker, 1857. 
by monotypy). 



78 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 56. Rashorulilln's hii/rhiui Kapuas 1976-41, 80.8 mm (MZB 3429). 



A Sundaic cyprinid genus with a narrow, sharp-snouted head; 
elongate subcyUndrical or fusiform body, with a median keel 
extending between pelvic and anal fins; gill rakers numerous 
and very elongate or lanceolate, 9-10 + 32-35=41-45. Dorsal- 
fin branched rays 7. anal 17-19, Hyaline eyelid very well de- 
veloped, extending anteriorly from snout to in front of nostrils 
and posteriorly almost onto gill cover. Jaws terminal, lips thin. 
Pharyngeal teeth in three rows 1,3,5/5,3,1. Lateral line com- 
plete, slightly decurved anteriorly but otherwise nearly straight. 
Lateral scale series about 60, predorsal scales 25, transdorsal 
18, circumpeduncular about 28, scales between lateral line and 
pelvic-fin origin 5-1/2. Vertebrae 23-24+19-20=42(3), 43(3). 
The genus comprises a single species. 

Rasborichthys helfrichii (Bleeker. 1857) 

I Figure >6) 

lA'ucncii', HeUnchn Bleckcr. IS57i;;l.S (t\pe loLalil\ kahajan River). 
Rashoruhlhv^ Hclfiichii Blocker. 18.59,/: 1 5.S; 186l.l<;:4.56. 

Material Examined —Weslern Borneo; Kapuas 1976-3.3, 21: 77. 1-93, s mm 
(BMNH 1982.3.29.109-10, CAS 49311, MNHN 1982-689, MZB 3428, RMNH 
28857, IIMMZ 209889); Kapuas 1976-41, 13; 76.7-92.8 mm (AMNH 48936, 
FMNH 94227. MZB 3429, IISNM 230240, ZMA 1 16.532). 

Distribution. — Sumatra (BatangHari). Borneo (Kapuas, Ka- 
hajan). 



Rohteichthys Bleckcr, I860 

Rohlcuhlhv: B\ceker. 186l)(;395 (tvpe species llaihus micnilcpi-. Bleckcr, 1850. 
by monotypy). 

Body moderately deep, strongly compressed. Dorsal profile 
strongly sloped, head concave above eyes. Abdomen anterior 
to pelvic fins flattened, posterior to pelvic fins strongly com- 
pressed or cultrate, with a well developed midventral membra- 
nous keel. Last simple dorsal-fin ray stout and heavily serrate; 
dorsal-fin branched rays 8, anal-fin branched rays 5. Lateral line 
complete, nearly straight, midlateral. Scales without radii; lat- 
eral scales 68-72, transdorsal 27, predorsal about 35, circum- 
peduncular 33. Vertebrae 18+15=33 (2, Kapuas). Body and 
fins plain except for a large dark midpeduncular spot. 

Mouth terminal, moderately upturned. Jaws narrow, mod- 
erately elongate. Barbels absent. Rostral cap thin, deeply incised, 
its anterior margin straight. Lips thin. No discrete horny jaw 



sheaths. Mandibular symphyseal knob weakly developed. Sub- 
lacrimal groove deeply incised. Adipose eyelid weakly devel- 
oped. Gill rakers moderately long, bony, 2-3+11 = 13-14 on 
first arch. Pharyngeal teeth uncinate, in three rows, 2,3,5/5,3,2. 
Relationships of Ruliieichlliys have not been discussed since 
Bleckcr (1860:397) compared it to Rohtee (now Osieobrama) 
vigors/, to which it is perhaps closely related. It also seems to 
be a close relative of Cyclocheilichtlivs. The numerous special- 
ized rows of hypertrophied cephalic cutaneous papillae, not pre- 
viously noted in Ruhtcichlhys. are virtually identical in the two 
genera. Rohteidilhys has numerous transverse rows of papillae 
on the dorsal surface of the head from snout-tip to occiput, 
transverse rows on the isthmus between the mandibular rami, 
vertical and longitudinal rows on the cheek and mandibular 
regions, vertical rows on the opcrcle, and oblique rows on the 
subopercle. all as in Cyclocheilichthys. The intermandibular pad 
is relatively elongate in Rohteichthys and not so sharply incised 
but is otherwise similar to the papillose intermandibular pad of 
Cyclocheilichthys. Other unusual or specialized characters shared 
with Cyclocheilichthys include the unusually straight lateral line 
and overall similarity of head and body form (including concave 
head, and abdomen flat anteriorly but strongly compressed pos- 
teriorly). In overall morphology Rohteichthys is remarkably 
similar to deep-bodied species of Cyclocheilichthys such as C 
repasson. The morphology of the pharyngeal jaws and teeth of 
Rohteichthys (Weber and de Beaufort, 1916; fig. 42) is virtually 
identical to that of Cyclocheilichthys (op. cil.. fig. 66). It differs 
from Cyclocheilichthys in having the snout more pointed and 
the jaws upturned and terminal (rather than subtcrminal) and 
the cultrate portion of the abdomen with a well developed mem- 
branous keel (entirely absent in all species of Cyclocheilichthys). 
Only one species of Rohteichthys is known. 

Rohteichthys microlepis (Bleeker, 1850) 

(Figure 571 

Innl'iis microlepis Bleeker, 1850:12 (type loealiU Banjcrmassing, in lluviis). 
SvM, mills munilcpis Bleckcr, 1851./;160, 
Koliiir iiiii'iKlcpii. Bleckcr, 186ftj;396, 

Material Examined. — Western Borneo: Kapuas 1976-33, 4: 90.2-97.2 mm 
(C AS 443 1 2, MZB 3430, RMNH 28858); Kapuas 1976-44, 3: 107-146 mm (MZB 
3431, USNM 230241). 

Distribution. — Sumatra (Palembang, Lahat, Djambi). Bor- 
neo (Kapuas, Barito). 



ROBERTS-WESTERN BORNEO FISHES 



79 




Figure 57. Rohleichlhys nucrolcpis. Sumatra (after Weber and dc Beaufort 1916. fig. 41). 



Schismatorhynchos Bleeker, 1855 

Schismalorhynchos Bleeker. I855/i:258. 269 (type species Lohochcilos hclcro- 
rhynchus Bleeker. 18.'>3. by monotypy). 

Juveniles and adults of both sexes with a large, deeply un- 
dercut and heavily tuberculate secondary rostrum. Rostra! cap 
and upper lip similar to those of Lohocheilps: rostral cap thick, 
its margin entire, with a deep subrostral groove; upper lip almost 
as thick as rostral cap, its margin entire; no separate upper homy 
jaw sheath. Homy sheath of lower jaw enormous, but not so 
thickened or as strongly projecting dorsally as in Lobocheilos. 
the margin strongly curved and with a very thin, flexible cor- 
nified cutting or scraping edge. Lower lip fleshy, but not thick 
or deeply incised, as in Lobocheilos, and its free anterior margin 
weakly fimbriate. Palate with delicate, ventrally projecting, lon- 
gitudinally crcnulate lobes. Short rostral and maxillary barbels 
present. 

A well developed secondary rostrum occurs m only two other 
cyprinoid genera, both Asian. It is characteristic of several species 
of Garra (including G. gotyla) and one of Gastromvzon {G. 
borneensis). It is lacking in the Gastroniyzon and Garra known 
from western Bomeo. In all three of these Asian genera the 
secondary rostrum generally is evident in juveniles long before 
sexual maturity, and tends to be well developed in both sexes; 
the relationship of sex to development of secondary rostrum is 
not well understood; sometimes it seems to be somewhat larger 
in males, but in other instances it may be larger in females (e.g., 
some Garra). The secondary rostrum and portion of the snout 
underlying and in front or to the side of it tend to be heavily 
tuberculate. The function of the tuberculate snout in these fishes 
is not well understood, but it is probably involved with inter- 
as well as intra-specific territorial behavior, non-sexual as well 
as sexual. 

Schismatorhynchos comprises two species, 5. heterorhynchiis 
and S. nukta (sometimes placed in a separate genus, Nukta): 
the latter occurs in southem India. 



Schismatorhynchos heterorhynchos (Bleeker, 1853) 

(Figure 58) 

Lobocheilos heterorhynchos Bleeker, 185.V;524 (type locality Solok, in fluviis). 

Schismatorhynchos lohocheltoides Bleeker, 1855i:259, 260 (unwarranted substi- 
tution of species name), 

Schismatorhynchos hetcrorhrnchos Bleeker. 1860i);131 (unwarranted spelling 
emendation). 

Tylognalhiis heterorhynchiis Guntlier. 1868ii:67. 

Material Examined. — Sumatra; BatuSanghar, 136 mm (CAS-SU 8077). West- 
ern Bomeo: Kapuas 1976-24, 2: 62.6-87.9 mm (MZB 3432. RMNH 28859); 
Kapuas 1976-27, 6: 83.9-105 mm (CAS 49313. MZB 3433, USNM 230242); 
Kapuas 1976-29. 88.0 mm (MZB 3434), 

Distribution. — Sumatra (Solok, Lahat, Batu Sanghar). Bor- 
neo (Kapuas, Mahakam, Kinabatangan). 



Thryssocypris Roberts and Kottelat, 1984 

Thrrssocypns Roberts and Kottelat, 1984:142 (type species Thryssocypris sma- 
rciiitlinit\ Roberts and Kottelat. 1984, by original designation). 

Small, insectivorous, anchovy-like cyprinids with strongly 
compressed head and body, beaklike snout, no barbels. Differs 
from most other cyprinid genera in western Bomeo in having 
pharyngeal teeth in two rows. For other diagnostic features and 
osteological description see Roberts and Kottelat (1984). 

Relationships of Thryssocypris to other cyprinids are un- 
known. Two species, one in the Kapuas and one in the Mekong. 



Thryssocypris smaragdinus Roberts and Kottelat, 1984 

(Figure 59) 

Tliryssocvpris smaragdinus Roberts and Kottelat, 1 984: 1 46 (type locality Kapuas). 

Material Examined. —Western Borneo; Kapuas 1976-36. MZB 3435. 49.4 
mm holotype) and CAS 49314. USNM 230243. 4; 46,8-54,0 mm (paratypes). 

This species is known only from the type specimens collected 
in the mainstream of the upper Kapuas River. 



80 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 58. Schisinukirlniulios Iwlcrorhynchos Kapuas 1976-29, 88.0 mm (MZB 34.M). 



Thynnichthys Bleeker, 1859 

Thvnnuhlhvi Bleeker. 1859u:153 (type species Lcutisciu ihyim-iidcy Bleeker, 1852. 
by subsequeni designation of Bleeker. 186.t/);20l). 

Body somewhat heavy-set, more or less finely scaled. Lateral 
line complete. Dorsal fin with 8-10 branched rays, anal with 5. 
Head broad with large gill covers. Mouth terminal, broad. Pre- 
maxillae, maxillae, and lower jaw very thinly covered with skm; 
no discrete rostral cap. lips, or horny jaw sheaths. Premaxillae 
moderately protrusible. Barbels absent. Gill rakers confined to 
uppermost portions of trailing edge of first gill arch and of lead- 
ing and trailing edges of arches 2-4; leading edge of first gill arch 
rakerless. Pharyngeal pad anteriorly with broad longitudmal 
sulci opposed to long rakerless portions of lower limbs of gill 
arches, posteriorly with complex epithelial folds or convolutions 
opposed to raker-bearing portions of arches. Pharyngeal teeth 
with a flat, oblong crown, in three rows wedged together. 4-5,3- 
4.2-3/2-3.3-4.4-5 (Weber and de Beaufort 1916:121). 

Thvnnicliihys comprises several species in southeast Asia and 
India. Two species are known from western Borneo, and one 
from eastern Borneo. They arc most readily distinguished by 
differences in scale counts. 



Thynnichthys polylepis Bleeker. 1860 

(Figure 611) 

lln-iimchlhvs pohicpis Bleeker. 1859(;.154 (nonien nudum); 1860:407 (type lo- 
ealily Palembang, Sumatra, and Ponlianak. Borneo), 

Materlm. Examined, — Wesiem Borneo: Kapuas 1976-15. 105 mm (MZB 3436); 
Rapuas 1976-20. 7: 72.9-96,0 mm (CAS 49315, MZB 3437); Rapuas 1976-33, 
V 69,7-1 15 mm (MZB 3438. I'SNM 230244); Kapuas 1976-41. 2: 102-103 mm 
(,MZB 3439, RMNH 28860): Kapuas 1976-49. 62.3 mm (MZB 3440), 

Distribution. — Sumatra (Palembang; Danau Sialong lo- 
tong), Borneo (Kapuas). 

Thynnichthys thynnoides (Bleeker, 1852) 

I.ciuiHii\ ihyniwidc.s Bleeker, 1852i':599 (type localuv Palembang, in lluviis). 
Thynnichthys thynnoidei Bleeker, 1859i(:153, 1860i;:408, 

Material Examined. — Western Borneo: Kapuas 1976-33, 7: 1 19-158 mm (CAS 
49316, MZB 3441, RMNH 28861, USNM 230245). 

Distribution. — Thailand (Mekong, Chao Phrya, Tale Sap, 
Tale Noi, Patani R,), Malay Peninsula (Perak. Pahang). Sumatra 
(Palembang. Djambi. Kampar R,. Gunung Sahilan. Kwantan 
R., Danau Sialong lotong), Borneo (Kapuas). 



Key to Thyumchlhys of Borneo 

la Lateral line scales 47; scales between lateral line and dorsal 
fin 1 1 T. valllaiili Weber and de Beaufort, 1916 

lb Lateral line scales 56-61; scales between lateral line and 
dorsal fin 12-13 T. thynnoides 

Ic Lateral line scales 65-75; scales between lateral line and 
dorsal fin 16-17 T. polylepis 



Tor Gray, 1833 

ro) Gray, 1833:96 (type species Cypnniis lor Hamillon-Buchanan, 1822, by ab- 
solute lautonymyl. 

Tor cf tambra (Valenciennes. 1842) 

Hcirhus laiiihra Valenciennes in Cuvier and Valenciennes. 1842:190 (type locality 
"cn\ irons dc Builen/org," ,lava). 



^S^ 










mm 


jd**^ 


s^i^SSHfc 


Ss~^^^ 







■ -^^^^l^^ 




iPPPiPW^ 


iwp* 


^ 


'A 




.^•i 




iivCP**^ 


1 


■^^^^ 


Wtt^im^ 


li 


^ 


f. 


■■■^ 





Figure 59, I hryssoLVpns Miuircinciiniis Kapuas 1976-36, 49.4 mm (MZB 3435, hololype). 



ROBERTS-WESTERN BORNEO FISHES 



SI 




Figure 60. TInnmchlhys polylepis^ Kapuas 1976-41, 102 mm (MZB 3439). 



Lahcobarbus lainbra Bleeker. 1857/);355. 

"'.Punliusslreeien Myers. 1 927: 1 (type locality Baloi River, Rejang basin. Sarawak). 

Material Examined.— Western Borneo: Ivapuas 1976-30, 105 mm (MZB 3442). 

The single specimen of this species obtained by the Kapuas 
survey of 1976 has been examined by Walter Rainboth. He has 
tentatively identified it with Puiitius strecteh. a probable syn- 
onym of Tor tamhra. 

Tor tambroides Bleeker, 1854 

(Figure 61) 

Lahcobarbus tambrotdes Bleeker. 1 854a:92 (type locality numerous places in west- 
ern Sumatra and western Java). 

Material Examined,— Western Borneo: Kapuas 1976-24. 8: 37.1-84.6 mm 
(CAS 49317. MZB 3443. USNM 230246); Kapuas 1976-25. 4: 65.0-90.5 mm 
(RMNH 28862, MZB 3444); Kapuas 1976-29, 8: 15,9-29,4 mm (CAS 49318, 
MZB 3445). 



Gyrinocheilidae 

The extraordinarily specialized southeast Asian mountain 
stream fishes assigned to Gyrinocheilus. the sole genus com- 
prising the family Gyrinocheilidae, are very poorly known mor- 
phologically and systematically. Their unique specialization is 
a spiracle-like opening at the upper limit of the gill cover by 
means of which the gills are ventilated during long periods while 
the heavily treaded sucker-like lips are expanded and attached 
to the rocky or stony substrate thus occluding the oral respiratory 
current. The algae-feeding Gyrinocheilus are also the only 
cyprinoids in which the pharyngeal teeth are absent (young spec- 
imens should be examined to see if they also lack the teeth). 

Gyrinocheilus superficially at least resembles the cyprinid 
subfamily Labeoinae, but no osteological or other studies have 
been done to determine its relationships. The thick, expandable 
lips are covered with numerous parallel rows of large, close-set. 
square or rectangular papillae. Similar labial papillae do not 
occur in any African, Indian, or southeast Asian cyprinids but 
do occur in Semilabeo Peters, 1880 (with Pseudogyrinocheilus 
Fang, 1933, a junior synonym), the species of which are endemic 
to southern China (where Gyrinocheilus is unknown). Semilabeo 
lacks the spiracle-like specialization but otherwise closely re- 
sembles Gyrinocheilus. They probably are closely related. 



Gyrinocheilus VaiUant, 1902 

Gvnnoclieiliis Vaillant, 1902:107 (type species Gyrinocheilus piisliilosiis Vaillant. 
1902, hy monotypy). 

Gyrinocheilus includes several poorly known species in Thai- 
land (especially the Mekong) and Sumatra badly in need of 
systematic study, A single species is known from western Bor- 
neo. 

Gyrinocheilus pustulosus Vaillant, 1902 

(Figure 62) 

Ginm)f/(c;/»5/>»5(»/(«iH Vaillant. 1902:1 1 1 (type locality "'embouchure du Raoen. 
haut-Sihau," Kapuas). 

Material Examined— Western Borneo: Kapuas 1976-27, 299 mm gravid fe- 
male (MZB 3446), 

Gyrinocheilus pustulosus seems to be the largest gyrinocheilid. 
The very numerous almost parallel rows of close-set breeding 
tubercles covering almost the entire side of the head has been 
observed only in this species. My 299 mm gravid female is very 
similar in all respects to Vaillant's illustration of one of the type 
specimens (reproduced here as Fig. 62). 

Distribution. — Known only from the Kapuas. 

Homalopteridae 

Homalopteridae, as currently understood, constitutes a large 
and diverse family of loach-like fishes widely distributed from 
India to China and in the main islands of Sundaland. The main 
concentrations of species, however, occur in southern China and 
on Borneo. Nearly all of the species are adapted to living on the 
bottom of swift-flowing mountain streams or in riffles of low- 
land streams, and some are among the most highly specialized 
of cyprinoids. All are characterized by more or less enlarged 
and horizontally flattened paired fins, and there is a strong ten- 
dency for increased numbers of pectoral- and pelvic-fin rays. In 
several of the most specialized taxa of China and Borneo the 
greatly enlarged pelvic fins are united to each other posteriorly, 
a condition otherwise unknown in lower teleosls. 

The more specialized homalopterids are poor swimmers but 
have evolved new modes of creeping and crawling locomotion 
(Wickler 1971). In all homalopterids the ventral surface of the 



CALIFORNIA ACADEMY OF SCIENCES 




FiciiRtM, Tor lamhiKiJc Rapuas 147(i-24. .S4 h mm (( AS 443 I 7) 



anteriormost rays in the pectoral and pelvic fins bear large ad- 
hesive or Irictional pads; these pads are unculiferous, as ob- 
served in numerous Chinese species by Chang (1945) and in 
several Bornean species by me (Roberts 1982(7). In at least one 
Chinese species. Pseudogastromyzon niycrsi Herre. 1932. the 
lips bear specialized unculiferous ridges employed in feeding 
upon filamentous algae and associated food items (Roberts, un- 
published observation). 



Higher classification of Homalopteridae is in an unsatisfac- 
tory state. There is evidence that the two subfamilies currently 
recognized. Homalopterinae and Gastromyzontinae. are un- 
natural assemblages, and that it may well be that Homalopter- 
idae itself IS polyphyletic. Current concepts of the subfamilies 
originated with Hora (1932). who recognized Homalopterinae 
chiefly on the basis of having two or more simple rays in the 
pectoral and pelvic fins, while Gastromyzontinae have only one 




Fic.iiRt fi2 GYrtihh-hcilus pusdilmm Kapuas (alter Vaillant I902;1 12). 



ROBERTS- WESTERN BORNEO FISHES 



83 



simple ray in each fin. Indeed, for the Bomean taxa. this simple 
distinction does not seem to result in a natural classification 
into two subfamilies. Thus, Bomean Homalopterinae comprises 
only species of Homaloptera and of its close relative, the mono- 
typic Neohomaloptera (Homaloptcra is by far the most speciose 
homalopterid genus, with about 40 species). Bomean Gastro- 
myzontinae, although morphologically diverse, also appear to 
represent a monophyletic group. Problems arise, however, in 
attempts to integrate the numerous Chinese genera and species 
into this classification, as has been done by Hora (1932), Chang 
(1945), Silas (1953). and Chen (1980). In my opinion none of 
the Chinese species is congeneric with Bomean species, and the 
species from the two areas probably do not even belong in the 
same subfamily. 

One of the most striking differences between Bomean Hom- 
alopterinae and Gastromyzontinae and the Chinese taxa cur- 
rently assigned to these subfamilies concems the enlarged rib 
which is ligamentously attached to the pelvic-fin girdle in the 
vicinity of the origin of the anteriormost pelvic-fin ray. This 
enlarged rib supposedly distinguishes Homalopteridae from 
Cobitidae. in which it is lacking. In Bomean Homalopterinae, 
so far as known, the enlarged rib is born on the 10th vertebra, 
while in Bomean Gastromyzontinae it is bom on the 1 2th ver- 
tebra (Table 3). This provides an additional distinction between 
Bomean members of the two subfamilies and suggests that we 
are in fact dealing with two monophyletic lines in Bomeo. It is 
of interest that the relatively generalized Glaniopsis, some of 
the species of which superficially at least resemble the cobitid 
subfamily Nemacheilinae. appears on this basis to be part of 
the Bomean Gastromyzontinae. An apparently unique character 
shared by several Bornean Gastromyzontinae involves the na- 
ture of the tuberculation in the males. In all or almost all Bor- 
nean Homalopteridae. in mature males the dorsal surface of the 
anteriormost pectoral-fin rays bears large pads of fine, close-set 
multicellular homy tubercles. In addition, males of Bornean 
Gastromyzontinae may have uniserial rows of fine, multicellular 
tubercles on the snout, cheek, or gill cover, or on the body 
immediately above the base of the pectoral fin. Such "comb- 
like" or "ctenoid" tuberculation occurs in Gastromyzon cteno- 
cephalus Roberts, 1982, Neogastromyzon paucimdiatus Inger 
and Chin, 1961, Pwtomyzon griswoldi, and an undescribed 
species of Hypergaslromyzon from the Lupar basin, Sarawak 
(pers. obs.) but has not been found in any Chinese species. 

The constant position of the enlarged rib and distinctive na- 
ture of the male tuberculation set Bomean Gastromyzontinae 
apart from Chinese species. Some of the more generalized Chinese 
homaloplerids lack the enlarged rib (Crossostoma. Formosama. 
Vanneniania): in others it is present on the 10th or on the 1 Ith 
vertebra, but has not been observed on the 12th (Table 3). 
Another character suggesting that Bomean and Chinese Gas- 
tromyzontinae are not so closely related involves the suprapelvic 
flap in those forms with greatly enlarged pelvic fins. In the 
Bomean forms (Gastromyzon. Hypergaslromyzon. Neogastro- 
myzon) the suprapelvic flap is continuous anteriorly with a flat 
lateral extension of the body wall which forms the ventrolateral 
abdominal margin. In Chinese taxa the flap arises directly from 
the dorsal surface of the pelvic fin. 

The Kapuas survey of 1976 resulted in discovery of several 
new Gastromyzontinae. Most of these belong in Gastromyzon 
and have been described already (Roberts 1982fl'); in addition. 




Figure 63. Mouthparts of Bomean Gastromyzontinae. a, Hypergaslromyzon; 
b, Gaslroittv:on. c, Neogaslromyron. 



an undescribed genus and species is reported herein [Hyper- 
gaslromyzon hiimilis). These discoveries bring the total known 
homalopterid genera to about 43 and the species to over 160. 
A synopsis of homaloplerids is provided by Silas (1953). 

Key to Homalopteridae of westem Bomeo 

la Pectoral fins with 22-31 rays, a single unbranched ray 
anteriorly; pelvic fins united posteriorly, each with 18- 
24 rays (Gastromyzontinae) 2 

lb Pectoral fin with 11-17 rays, with at least two un- 
branched rays anteriorly; pelvic fins separate, each with 
7-11 rays (Homalopterinae) 7 

2a Mouth broad, almost as wide as head; cutting edge of 
horny jaw sheaths straight (Fig. 63b. c) 3 

2b Mouth narrow, less than one-third as wide as head; cut- 
ting edge of homy jaw sheaths curved (Fig. 63a) 

Hypogastromyzon htimilis 

3a Head width at least 5; sublacrimal groove absent; rostral 
and maxillary barbels project from posterior margin of 
rostral cap (Fig. 63c); adhesive pad of anteriormost pec- 
toral-fin ray with anterior margin continuous with skin 
of abdomen Neogastromyzon nieuwenhuisi 



84 CALIFORNIA ACADEMY OF SCIENCES 

Table 3. Vertebral Characters of Homalopteridae sensu lato. For definition of vertebral counts see p. 22 



Vertebral counts Vertebra beanng enlarged nb 



Homaloptennae 
HomalopWra nebtilosa 20+ 1 1 =3 1(4) 10(5) 

Kapuas 1976-6 (CAS 49323) 21 + 10=31(1) 

Homaloplera ophiolcpis 
Kapuas 1976-27 (CAS 49325) 22+14 = 36(2) 10(2) 

Homaloplera orlhogonmla 
Kapuas 1976-30 (CAS 49326) 21 + 10=31(1) 10(1) 

Homaloplera slephensoni 
Kapuas 1976-32 (MZB 3479) 21 + 11 = 32(1) 10(1) 

Mahakam (RMNH 7633, holotypc) 21 + 11 = 32(1) 10(11 

Homaloplera Iweediei 
Kapuas 1976-13 (CAS 49327) 19+12 = 31(1) 10(3) 

20+11=31(2) 

Ilomali ipiera :i <lh iigeri 
Kapuas 1976-29 (CAS 49331) 22+11=33(1) 10(1) 

Neohomaloplera johorensis 
Malay Peninsula (CAS-SU 66426) 21 + 1 1(2) 10(2) 

Gastromyzontinae 
Gaslromyzon horneensis 
BaramR. (CAS 53177) 24 + 9=33(1) 12(3) 

24+10=34(1) 
25+10=35(1) 

Gaslromyzon conlraelm 
Kapuas 1976-24 (CAS 44186) 22 + 9 = 31(2) 12(3) 

22+10=32(2) 

Gaslromyzon faseialiis 
BaramR. (CAS 53175) 23+10=33(2) 12(2) 

Gaslromyzon rtdens 
Kapuas 1976-24 (CAS 49322) 24 + 9 = 33(2) 12(2) 

Glaniopsis haniischi Boulenger, 1 899 
Ml Kinabalu (CAS 53185) 25+11 = 36(6) 12(6) 

Glaniopsis nuilliradiala Roberts. 1982 
Baram basin (CAS 53179, paratypes) 24+12=36(1) 12(15) 

25+11=36(7) 
26+10=36(1) 
25+12 = 37(6) 

Hypergaslromyzon Inimihs 
Kapuas 1976-30 (CAS 49333, MZB 3480. types) 24 + 8 = 32(2) 12(2) 

25 + 8 = 33(1) 

"Hvperiiaslromvzon" sp. 

BatangAi, Lupar basin (uncat.) 22 + 9 = 31(3) 12(3) 

Neogasln^myzon nieioyenliuisi 
BatangAi. Lupar basin (CAS 53173) 22 + 8 = 30(1) 12(3) 

22 + 9 = 31(2) 

I'arhoinalopiera murosloma (Boulenger, 1899) 
Rcjang basin (FMNH 68134) 26+10=36(2) 12(5) 

27 + 9=36(3) 

Proiomyzon aphelocheilns Inger and Chin, 1962 
Sungei Kaingeran, N. Borneo (FMNH 68167) 25+10=35(1) 12(6) 

25+11=36(1) 
26+10=36(3) 
27 + 9 = 36(1) 

PrtHomvzon hinneensis Hora and Jayaram. 1951 

Sungci Kaingeran. N. Borneo (FMNH 68186) 24 + 9 = 33(4) 12(5) 

25 + 8 = 33(1) 



ROBERTS-WESTERN BORNEO FISHES 



Table 3. CoNTihorED. 



Vertebral counts 



Vertebra beanng enlarged nb 



Pwlomyzon griswoldi (Hora and Jayaram, 1951) 
Keningau. N. Borneo (FMNH 68133) 

Prolomyzon »hileheadi (VaiWanl. 1893) 
Ranau Distnct, N. Borneo (FMNH 68139) 

Chinese 
Beaufonia lewreni {Nicho\s and Pope, 1927) 

Hainan (CAS-SU 32570) 
Formosania lacustre 

Formosa (CAS-SU 18255) 

Hemimyzon formosanus (Boulenger, 1894) 

Formosa (CAS-SU 49364) 
Liniparhomaloplera dispans (Lin, 1934) 

Hong Kong (CAS-SU 32589) 

Pseudogaslromyzon fasciams (Sauvage, 1878) 
Kiangsi (CAS-SU 39644) 



Sinogaslromyzon sanhocnsis Fang, 1931 

China (CAS-SU 27502) 
Stnohomaloptera hojfmanm (Herre, 1938) 

Hainan (CAS-SU 33002, 33003, types) 
Smohomaloplera vaolanemis Fang. 1931 

China (CAS-SU 27503) 



244-9=33(6) 

234-10=33(1) 
244-9=33(5) 



24-1-9=33(1) 

24-H3 = 37(l) 
25 -H 3=38(1) 

254-12=37(1) 



244-11=35(1) 
254-10=35(2) 

274-12=39(1) 
284-12=40(1) 
294-11=40(1) 



23+13=36(1) 
21 + 12=33(2) 
23+13=36(1) 



12(6) 
12(6) 

11(1) 

none(2) 

10(1) 
none(3) 

11(3) 

10(1) 
10(2) 
10(1) 



3b Head width at most 4; sublacrimal groove present; ros- 
tral and maxillary barbels project from anterior margin 
of rostral cap (Fig. 63b); adhesive pad of anteriormost 
pectoral-fin ray with anterior margin free from abdomen 

{Gastroniyzon) 4 

4a Sublacrimal groove extending dorsally onto side of head 

G. ridens 
4b Sublacrimal groove confined to ventral surface of head 

5 

5a Postoral pouch present G. fasciatm 

5b Postoral pouch absent 6 

6a Subopercular groove present; abdomen scaieless 

G. contractus 
6b Subopercular groove absent; specimens over 40 mm with 
abdomen at least partially covered with scales 

G. lepidogaster 
7a A pair of small maxillary barbels at each comer of mouth; 
pectoral-fin rays 12-13; pelvic-fin rays 7 

Neohomaloptem johorensis 
7b A single small maxillary barbel at each comer of mouth; 
pectoral-fin rays 13-17; pelvic-fin rays 8-10 (Honia- 

lopicra) 8 

8a Pectoral fin falls considerably short of pelvic fin 9 

8b Pectoral fin reaches or overlaps pelvic fin 12 

9a Body compressed (deeper than wide), scales 56-67 10 
9b Body cylindrical or depressed, scales 45-48 1 1 



10a Scales 56-61. not keeled; eye 4,5-7 in head H. ogihici 
lOb Scales 63-67. keeled; eye 10-12 in head 

H. orthogoniata 

I la Body cylindrical, body and fins uniformly dark brown 

or blackish H. zollingeri 

I I b Body depressed, dorsal surface of body with faint round 

marks, paired fins strongly barred H. ophiolepis 

12a Pectoral fin reaching pelvic fin; scales 46-51, carinate 

H. cf stephensoni 

12b Pectoral fin overlapping pelvic fin; scales 32-40, non- 

carinate 1 3 

13a Lateral line scales 30-32; transverse scales 4-1/2-5-1/2/ 

4-1/2-5-1/2 H. tweediei 

13b Lateral line scales 36-40; transverse scales 6-6-1/2/6-6- 

1/2 H. ncbidosa 

Gastromyzon Giinther, 1874 

Gaslromyzon Giinther. 1874 454 (t\pc species (iuHioniyzoii horiwcnsis Giinther, 

1874, b\ nionotypy) 
Lcpidoglanis Vaillant, 1889:81 (type species Lcpulogluim monlicnh Vaillani, 1889. 

by monotypy), 

DiKGViosis. — Gastromyzon differs from all other Gastromy- 
zontinae in having a sublacrimal groove between lacrimal bone 
and rostral cap, and rostral and maxillary barbels projecting 
from anterior (rather than posterior) margin of rostral cap. Pec- 



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Figure 64. Uuslioniy: 



Kapuas 197b-25, 34.4 mm (MZB .1447, holot\pc). 



toral fin with 23-31 rays. Pelvic fins, united posteriorly, with 
18-25 rays. 

In my revision of Gastromyzcni the number of species was 
mcreased from four to nine, including three new species dis- 
covered m the Kapuas basin (Roberts l'>82(/). Descriptions and 
figures of all of the species are given in the revision. The genus 
is restricted to Borneo. 

Gastromyzon contractus Roberts, 1982 

(FigLiri.' 64) 

(iastniinvznn ci'iilracdi', Roberts. 1982i/;5U4, tig. b (type locality Sungai Tebelian, 
a tributary ofSungai Pmoh, 19 km upstream from Nangapmoh. Rapuas basm). 

Material Examined. — Western Borneo. Kapuas basin (holotype and para- 
types); Kapuas 1976-24. 17; 29,l-.'t6,9 mm (BMNH 1982.3.29.111-112, CAS 
44 1 86, MNHN 1 982-690. MZB 3448, RMNH 28863, USNM 230247, paratypes); 
Kapuas 1976-25. 30.2-34.4 mm (MZB 3447. holotype. CAS 49319. paratype); 
Kapuas 1976-27, 2 2S 9-29 8 mm (CAS 49320. MZB 3449. paratypes). 

Apparently the smallest Gasimniyzon (largest specimen 36.9 
mm), differing from all other species in having pelvic fin and 
dorsal fin (when depressed) reaching to or beyond level of anal- 
fin origin (depressed dorsal fin not reaching level of anal-fin 
origin, and pelvic fin reaching at most to anal fin in all other 



Gastromyzon). Sexually mature males develop dense patches of 
minute breeding tubercles on dorsal surface of first 8- 1 branched 
pectoral-fin rays (Roberts \9S2(i. fig. 2). 
Known only from Kapuas basin. 

Gastromyzon fasciatus Inger and Chin, 1961 

(Figure 6S) 

(iasinirnvzi'ii fascialiis Inger and Chin, 1 96 1 ; 1 73 (type locality Sungai Dapu, Baleh 
Ri\er, Rajang basin, Sarawak). 

Material Examined, -Sarawak; Baleh R,, 6: 48.1-65,9 mm (BMNH 
1 960, 1 2, 30, 1. FMNH 681 1 5 and 68 120; paratypes); Truson R„ Lawas, 70,4 mm 
(FMNH 45852); Pa Brayong, Truson R,. 2; 64,9-70,5 mm (FMNH 45853); Akah 
R,, Mehgong, 3: 44,1-51,0 mm (FMNH 68583); Sadong R„ 45.1 mm (CAS-SU 
32378); Arur Dalam. Padapur R,, Baram basin, 6: 38,6-69,3 mm (IRSNB 19725), 
North Borneo; Sungai Kamgeran, Tamhunan, 5: 20,6—46,5 mm (FMNH 68128); 
Parutan R,. Tambunan. 4; 1 7,2-32,8 mm (FMNH uncat), Kapuas basin; Kapuas 
1976-24. 59,0 mm (MZB 3450); Kapuas 1976-25, 56,0 mm (MZB 3451 ); Kapuas 
1976-26, 44,7 mm (MZB 3452); Kapuas 1976-27, 24,5 mm (MZB 3453); Kapuas 
1976-30,3: 39,5-57,5 mm (CAS 49321, MZB 3454, IISNM 230248); Bongan. 4: 
22,5-54,1 mm (RMNH 7636), Mahakam basin; Bo, 2; 53,2-56,8 mm (RMNH 
7638); Bluu or Bloeoc, 2; 39,2-54,3 mm (RMNH 7794), 

In life the nearly black or bluish-black coloration of the body 
and fins with thin, pale or almost white bars or stripes made 



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87 




Figure 65. Gaslroinyzon fascialus. Kapuas 1976-24. 59.0 mm (MZB 3450). 



this species stand out from all the others of the genus Gaslro- 
myzon encountered in the Kapuas. Predorsal scales 40-55; gill 
opening angular and subopercular groove present (gill opening 
vertical and subopercular groove absent in most other Gastro- 
myzon). 

Distribution (Fig. 66). — Northern Borneo (Kapuas, Maha- 
kam). 

Gastromyzon lepidogaster Roberts, 1982 

Gastromyzon lepidogaster Roberts. 1982</:509, fig. 9. 10 (type locality sources of 
Mcngalong River, Beaufort Distnct. north Borneo). 

Material Examined. — Western Borneo: Bongan. 72.3 mm (RMNH uncat.). 
North Borneo: Kajan basin. Mahakam basin. See Roberts (1982(/:509), 

Remark.. — This species is characterized by having the ab- 
domen of specimens over 40 mm more or less extensively cov- 
ered by minute scales (Roberts 1982^, fig. 9). The abdomen is 
scaleless in all other Gastromyzon except G. megalepis Roberts, 
1982 (from the Rajang basin), some specimens of which have 
a few relatively large scales on the posterior part of the abdomen 
between the pelvic fins. In the holotype and some other speci- 
mens of G. lepidogaster virtually the entire abdomen is covered 
with fine scales. In specimens from the Mahakam and Kapuas 
basins scales are confined to about the posterior half of the 
abdomen. Color in life unknown. 

Distribution (Fig. 66).— Northern Borneo (Kapuas, Maha- 
kam). 



110' 



112° 



• G borneensis 

♦ G contractus 

T G ctenocephaius 

A G lasciatus 

* G lepidogaster 
O G megalepis 



114° 

it G monticola 
o G punclulatus 
a G ridens 




Figure 66. Gastromyzon Geographical distribution. 



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Figure 67. Uiislrot>i\:iiii ruh-iis Kapuas l'*76-24, 50.0 mm {MZB 3455, hololype). 



Gastromyzon ridens Roberts. 1982 

(Figure h7) 

(iaslnmirziin rulcii-. Roberts. 1982i/:515, fig. 14 (type locality Sungai Plnoh, Ka- 
puas basin). 

Matkreal Examined. — Western Borneo: Kapuas 1476-24. '): 24.3-50.11 mm 
(UMNH iq82.3.2<).l 13. CAS 49322. FMNH 44228. MNHN 1982-691 . RMNH 
28864. USNM 230249; holotype and paratypcs). Sarawak: Senah, 6: 35.4-48.6 
mm (BMNH 1893.3.6.270-275, paratypes), .Sadong River, 9: 21.2-36.5 mm (CAS- 
SU 32378). In addition. I recently identified numerous specimens of this species 
from the Batang Ai. a high gradient tributary of the Balang Lupar in Sarawak, 
collected by .loan Cramphorn (BMNH). 

Gaslroinyzoii rulcns dilfers from all other members of its 
genus in having the sublacrimal groove extending well onto the 
side of the head, which gives it a very distinctive appearance (a 
laughing expression, hence the name ridens). This character is 
well marked even in the smallest specimens examined. 

DisTRiBinioN (Fig. 66), — Sarawak and western Borneo. 



Honialoptera Hocven. 1833 

Unmali'plcra van Hasselt i)i Kuhl and van Hasselt. 1823:133 (nonicn nudum). 
Ilnmalitptcia Hocven. 1833:21 1 (type species lloiiiuloplcra occllala Hoeven. 1833. 

b\ monotypy). Original publication not seen; see discussion by Hora (1932: 

274-276). 



Uclxui Vinciguerra. 1890:330 (type species llnnicilnpu-ni hihiicala Blyth. 1860. 

b\ monotyp\ ). 
l/omaloplcroiiics Fowler. 1905:476 (type species HuDialnpicra nassinkn Bleeker. 

1853. by original designation and nionolyp\). 
Chopram Prashad and Mukerji. 1929; 188 (type species C/io/i/nd; /Hp/io/a Prashad 

and Mukerji. 1929. by original designation and monotypy). 
Ilnnidldplcrula Fowler. 1940:379 (type species lloinalnplcnda riplcvi Fowler. 1940, 

b\ original designation and monotypy). See Silas ( 1953:188). 

Homaloptera nebulosa Alfred. 1969 

(Figure 68) 

Hoiihih'pifiu iichiilnvi .Aillred. 1969:227. pi I. fig. 304 (t>pe locality "River Sok. 
Kanipong Sok. Kelantan." Malay Peninsula). 

Material Examined. — Malay Peninsula: River Sok, KampongSok. Kelantan. 
21.8 mm (CAS-SH 66428. paratype). Western Borneo: Kapuas 1976-6, 5: 19.7- 

33 5 mm (CAS 49323. MZB 3456): Kapuas 1976-8. 25. 1 mm (MZB 3457); Kapuas 
1976-25. 3: 29.9-31.8 mm (BMNH 1982.3.29.114-115, MZB 3458); Kapuas 
1 976-26. 7: 28.2-33.7 mm (FMNH 94229. MNHN 1 982-692. MZB 3459. RMNH 
28865); Kapuas 1976-27. 3: 30.5-32.3 mm (MZB 3460. HMMZ 209871 ); Kapuas 
1976-29. 4 34 9-35 9 mm (CAS 49324. MZB 3461); Kapuas 1976-30, 8: 26.8- 

34 9 mm (MZB 3462. IISNM 230250. ZMA 1 16.533). 

This species has been known until now only from the holotype 
and four paratypes, 22.3-24.5 mm. Thus the Kapuas specimens 
represent a substantial increase in maximum known standard 
length (to 35.9 mm) as well as the first record for Borneo. 



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89 




Figure 68. Homalopicra nchulosa Kapuas 1976-6, 31.8 mm (CAS 49323). 



Homaloptera ogilviei Alfred, 1967 

Homaloplera ogilviei Alfred, 1967:587, fig. 1,2 (ivpc localiiv Tclai River, Negri 
Sembilan, Malay Peninsula), 

Material Examined. — Malay Peninsula: River Jelai, Negri Sembilan, Malay 
Penmsula. 2: 26,4-28.5 (CAS-SU 66421; non-typesl. Western Borneo: Kapuas 
1976-10, 26.6 mm (MZB 3463). 

Body compressed (deeper than wide), eye diameter 4.5-7 in 
head, scales carinate, 56-6 1 , caudal peduncle slender (depth 1 5- 
16), body with a dark vertical bar continuing onto anterior 
portion of dorsal fin; paired fins (especially pectoral) almost 
vertically oriented (versus almost horizontally in all or most 
other Homaloplera). 

The peculiar orientation of the paired fins may be related to 
the tendency of this species to stay in vegetation (Alfred 1969) 
rather than on rock or gravelly bottom like most other Hom- 
aloptera. 

Distribution. — Known only from the Malay Peninsula (Ne- 
gri Sembilan, Trcngganu, Johore, Pahang, Perak [Alfred 1969: 
220] and the Kapuas). 

Homaloptera ophiolepis Bleeker, 1853 

(Figure 69) 

Homaloplera ophiolepis Bleeker, 1853^:157. 160 (type locality ■'Bandong, in flu- 
viis"). 

Material Examined. —Western Borneo: Kapuas 1976-27, 4: 62.8-111 mm 
(CAS 49325. MZB 3464, RMNH 28866): Kapuas 1976-50, 2: 36.9-39.0 mm 
(MZB 3465, USNM 230251), 



Distribution. — Sumatra (Lahat). Borneo (Kapuas. Maha- 
kam), Java (Bandung, Parongkalong). 

Homaloptera orthogoniata Vaillant, 1902 

Homaloplera bilmeala Boulcnger, 1894:251 (notofBlyth; Baram River. Sarawak). 
Sec Weber and de Beaufort (1916:15-17), 

Homaloplera orthogonialu Vaillant, 1902: 122, fig. 33-35 (type locality "embou- 
chure du Raocn" [Kapuas]). 

Material Examined.— Malay Peninsula: Perak. Tapah Fisheries Station, 2: 
83.8-97.2 mm, CAS-SU 39390; River Tahan, King George V National Park, 
Chegar Sireh. 56.4 mm, CAS-SU 66423. Western Borneo: Kapuas 1976-30. 2: 
52,1-54.7 mm (CAS 49326. MZB 3466). 

Perhaps the most ornate species of Homalopterinae. The thin, 
dark longitudinal stripe through the eye and a similar vertical 
stripe extending below it are present in all specimens examined. 
In some, stripes are emphasized by a thin pale area surrounding 
them. In larger specimens the three saddle-like marks on the 
dorsal surface of the body may break up or be augmented by 
smaller ventrolateral marks. 

In life the darker marks on the fins and body of the Kapuas 
specimens were a rich reddish brown, the lighter areas a rich 
tan. This species is perhaps most closely related to Homaloptera 
erythrorhina Valenciennes in Cuvier and Valenciennes, 1 846, 
which has similarly compressed body, fine carinate scales, and 
small eyes but coloration consisting of a series of five round 
spots confined on its dorsal surface and a well defined bar across 
the middle of the dorsal fin. H. erythrorhina is known from 
Sumatra and Java (Weber and de Beaufort 1916). 



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FioirRE 6>^. llonuili>i>lcra ophiolcpis Kapuas 1976-27, 111 mm (CAS 49325). 



Distribution. — Malay Peninsula (Perak, Pahang, Negri Sem- 
bilan, Johore; Alfred 1969). Sarawak (Baram). Borneo (Kapuas, 
Mahakam). 



Homaloptera cf. stephensoni Hora. 1932 

(Figua- 70) 

Honmloptera sicphcnsoni Hora. 1932:281 (type local)t\ upper Mahakam River. 
Borneo). 

Material Examined. — Western Borneo: Kapuas 1976-27. 17: 37.0-53,9 mm 
(BMNH 1482 3.29.1 19-1 10, CAS 49332. MNHN 19S2-h94. MZB 3477. RMNH 
28869); Kapuas 1976-29, 6: 38.6-57.2 mm (MZB 3478. USNM 230254); Kapuas 
1976-32. 57.5 mm (MZB 3479). Eastern Borneo upper Mahakam River. 48.2 
mm (RMNH 633. holotype o( H \liphcnsoni) 

Diagnosis. — A Homaloptera with head shorl; snout moder- 
ately pointed; head and anterior portion of body feebly de- 
pressed, ventral surface broad and flattened; scales on dorsum 
and sides moderately carinate; lateral line pored scales 44-52; 
pectoral fins, not overlapping pelvic fins when body is straight, 
with 5 simple and 1 1-1 2 branched rays; pelvic fins with 2 simple 
and 8 branched rays; dorsal-fin origin about equidistant between 
snout-tip and caudal-fin base or slightly closer to snout-tip; cau- 
dal peduncle elongate; caudal fin narrow, with pointed lobes, 
lower lobe longer than upper; dorsal surface of body with 5 
round spots, sides with 7-8 elongate blotches (sometimes par- 
tially fused, especially anteriorly); total vertebrae 32. 

The following description is based on the 48.2 mm holotype, 
a gravid female, and the 1 1 largest specimens from the Rapuas. 
46.7-57.5 mm (information on Kapuas specimens in parenthe- 
ses): head length 4.2 (3.9-4.4); snout moderately elongated and 
pointed, length 8.7 (8.3-9.2); horizontal diameter of eye about 
equal to interorbital width, 18.9 (18.3-21.3); body depth 6.5 
(7.2-9.7); length 7.4 (6.3-7.5). depth 16.1 (15.1-17.6); lateral 
line scales 44 (46-52); predorsal 17(1 9-22); circumpeduncular 
about 10 (12-14); dorsal-fin origin about midway between snout- 
tip and base of caudal fin, distinctly posterior to a vertical line 
through pelvic-fin origin; dorsal-fin branched rays 7; pectoral- 



fin simple rays 5 and branched rays 11-12; dorsum with 5 dark 
oval blotches, second centered on dorsal-fin origin, and sides 
with 7-8 dark elongate blotches centered on lateral line; ver- 
tebrae 21 + 1 1=32 in single Kapuas specimen radiographed. 

The holotype of//, stcphcmoni has a deeper body and slightly 
fewer scales than the Kapuas specimens; also its snout appears 
to be slightly shorter, more rounded, and downtumed (Hora 
1932. pi. 11. fig. 1). These differences lead me to suppose that 
they might not be the same species. The deep body of the ho- 
lotype of //. stephensoni is at least partly due to its being a gravid 
female; a gravid 57.2 mm female from the Kapuas has the body 
depth distinctly less (7.2 vs. 6.5). In both of these specimens 
the maximum size ofthe eggs is about 0.5 mm. In other respects, 
however, the Kapuas specimens are very similar to the Maha- 
kam holotype and I tentatively conclude that they are conspe- 
cific. 

Distribution. — Borneo (Kapuas, Mahakam). 

Homaloptera tweediei Herre, 1940 

ll.inuilnptcra Im'cdu'i Herre. l940i;:7-8, pi. 1 (type locality "a shallow rapid creek 
m the Mawai district, .lohore. about 40 miles north of Singapore"). 

Material Examined —Malay Peninsula: Mawai district, .lohore. 3: 23.0-26.0 
mm (C.'^S-SU 33012. 33013; holotype and paratypes) Western Borneo: Kapuas 
1976-10. 2: 17.0-17 7 mm (BMNH 1982.3.29.116. MZB 3467); Kapuas 1976- 
13.4: 17.7-20.7 mm (CAS 49327. MZB 3468): Kapuas 1976-14. 21.9 mm (MZB 
3469); Kapuas 1976-16. 11,3 mm (MZB 3470): Kapuas 1976-37. 2: 16.0-16.3 
mm (CAS 49328. MZB 347 1 ); Kapuas 1 976-39. 23: 1 2.5-1 7. 5 mm (MNHN 1 982- 
693. MZB 3472. RMNH 28867. UMMZ 209903. USNM 230252. ZMA I 16.534); 
Kapuas IQ76-47. 2: 2(1 1-20.3 mm (CAS 49329. MZB 3473). 

This IS probably the smallest species of Homaloptera. Alfred 
(1969:225-227) reported 82 specimens from nine localities in 
the Malay Peninsula of 14.4-30.0 mm; the species evidently 
attains sexual maturity at standard lengths of 20 mm or less. 
The Kapuas material includes 35 specimens from seven local- 
ities; the largest of these is only 21.9 mm. and a large majority 
are under 20 mm. 

Distribution. — Malay Peninsula. Borneo (Kapuas). 



ROBERTS-WESTERN BORNEO FISHES 



91 




Figure 70. fluinnlttplcra cf. slepht-nsom. Kapuas 1976-32, 57.5 mm (MZB 3479). 



Homaloptera Zollinger! Bleeker, 1853 

(Figure 71) 

Homaloplera jaraitica van Hasselt in Kuhl and van Hassclt. 1823:133 (nomen 

nudum). 
Hoinatoplem Zollingen Bleeker. 1853i;:159 (type locality "Balavia. Bandong. in 

fluviis"). 
'] Homaloptera nigra Alfred, 1969;217 (type localit> Tahan River. Pahiang. Malay 

Peninsula). 

Material Examined. — Malay Peninsula: River Tahan, Pahang, 2: 45.6-58.3 
mm (CAS-SU 66424, paratypes H nigra). Western Borneo: Kapuas 1976-22. 6: 
35.6-40.5 mm (CAS 49330. MZB 3474): Kapuas 1976-27, 8: 56.0-79.5 mm 
(BMNH 1982.3.29.117-118, CAS 44185, MZB 3475, RMNH 28868. USNM 
230253): Kapuas 1976-29, 2: 59.4-68.5 mm (MZB 3476, CAS 49331). 

The differences between H. nigra anti H. zollingen indicated 
by Alfred are rather slight; with the limited material at hand I 
am unable to decide whether they are distinct species. 

Distribution. — Thailand (Bangpakong). Malay Peninsula 
(Perak. Pahang, Johore). Sumatra (Lahat). Borneo (Kapuas). 
Java (Bandung, Batavia). 



Hypergastromyzon new genus 

Type %?i.<:\iS. — Hypcrgasiromyzon hiiiinlii neu species. 

Diagnosis.— A gastromyzontin with head and body moder- 
ately broad and very strongly depressed; pectoral and pelvic fins 
greatly enlarged; pectoral-fin origin below eye; pelvic fins com- 
pletely joined posteriorly; mouth small, about one-third as wide 
as head, with horny jaw sheaths strongly curved; three pairs of 
very short barbels, two rostral and one maxillary; rostral cap 
largely fused to or indistinguishable from upper lip except around 
base of rostral barbels; lower lip with medial groove well de- 



veloped posterolaterally but interrupted anteromedially (Fig, 
63a); snout broadly rounded in dorsal view, strongly acuminate 
viewed from the side; gill opening vertical, very small, above 
middle of pectoral-fin base; subopercular groove absent; base 
of first pectoral-fin ray separated from head by a deep groove 
or skin fold parallel to anterior margin of ray on ventral surface 
of head and continuing dorsally on head to behind base of ray 
(Fig. 63a); suprapelvic flap originating immediately behind pec- 
toral-fin base and continuing as a free flap above anterior third 
of pelvic-fin base, but entirely free from pelvic fin itself; ab- 
domen entirely scaleless. Pectoral fin separated from head by a 
deep groove extending ventrally to pectoral fin parallel to an- 
terior margin of first pectoral-fin ray and dorsally to pelvic fin 
to origin of first branched pectoral-fin ray. Dorsal surface of 
fleshy pectoral-fin base scaleless. 

The only other described Bomean gastromyzontins with greatly 
expanded pectoral and pelvic fins and completely joined pelvic 
fins are Gastn>niy:on and .\'eogastronn-:on: the other genera 
have pectoral and pelvic fins much smaller, with far fewer rays, 
and pelvic fins widely separated. Gastromyzon and Neogastro- 
myzon agree with each other and differ from Hypergastromyzon 
in having a broad mouth nearly as wide as the head, with homy 
jaw sheaths nearly straight transversely and a broad rostral cap 
separated from upper lip b> a deep rostral groove. The shape 
of the head is very different and body depth much greater in 
these genera than in Hypergastromyzon. Hypergastromyzon su- 
perficially resembles the Chinese gastromyzontin genus Beati- 
fortia Hora, 1932. Beaufortia has a narrow mouth as in Hyper- 
gastromyzon but differs in having origin of first pectoral-fin ray 
flush with surface of head dorsally and ventrally (rather than 
separated from head by a deep groove or fold of skin), supra- 



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CALIFORNIA ACADEMY OF SCIENCES 




Figure 71. Il,>niali>plcra :i<llingcrL Rapuas 1976-27, ^\^ mm (MZB 3475). 



pelvic flap originating near middle of pelvic fin and strongly 
attached to pelvic fin (rather than originating just behind pec- 
toral-fin base and entirely free from pelvic fin), pelvic fins in- 
completely joined posteriorly, and free margin of rostral cap 
lobosc. These characters suggest that Bcaufortia is more closely 
related to other Chinese genera than to Hypcrgasuoinyzon or 
any of the Bornean genera. 

Hypcrgastromyzou is known only from two specimens of an 
undescribed species obtained by the Kapuas survey of 1976. 

EjxMOi^OGW — Hypergastroniyzon (masculine), from Greek 
hyper and Gastwmyzon. 

Hypergastromyzon humilis new species 

(Figures (i.la. 11) 

HoLOTYPE. — MZB 3480, 35.7 mm (probably subadult female), Sungei Tamang. 
small forest stream with rocky bottom flowing mto Sungai Pinoh opposite mouth 
of Sungei Relawai (Kapuas 1976-30). 

Paratype. — CAS 49333, 37.5 mm (probably suliadult lemale), collected with 
holotype. 

Head length 4.0-4.2, width (measured just in front of pectoral- 
fin origin) 4,1-4.4; snout 8.7-8.8; eye 15.9-18.8; interorbital 
width 13.7-1 4.2; gill opening small, about equal to eye diameter, 
16.3-18.3; mouth width 15.6-16.6; length of all barbels one- 
fourth or less of eye diameter; greatest depth body 7.5-9.3; least 
depth caudal peduncle 13.2-14.2; pectoral fin originating below 
middle of eye, length 2.5; pelvic fin originating well in advance 
of vertical through dorsal-fin origin, length 3.4; anal fin small, 
widely separated from pelvic fin, length 7.9-8.5; distance be- 
tween postenor margin of pelvic fin and anal-fin origin 9.9-14.2. 

Pored scales in lateral series about 56; scale rows between 
dorsal fin and lateral line 1 1-12, between lateral line and pelvic 
fin 6; predorsal scales (very irregular) approximately 30^2; 
circumpeduncular 18-20. Anterior third of body with scale row 



immediately above lateral line and all scale rows below it con- 
siderably larger than scales more dorsally situated; this differ- 
ence in scale size diminishes toward mid-body, and in posterior 
half of body all scales are nearly equal in size. Dorsal body 
surface entirely scaled except fleshy pectoral-fin base below gill 
opening and adjacent to pectoral fin and dorsolateral margin 
and free portion of suprapelvic flap which are scaleless. Ab- 
domen entirely scaleless. 

Dorsal-fin rays iii7 (last ray simple); anal-fin rays ii4 (last ray 
simple). Pectoral-fin rays i28-30 (last 3-4 rays entirely divided 
to base, last ray sometimes simple). Pelvic-fin rays i 18-1 9. Pel- 
vic-fin membranes entirely united posteriorly. Posteriormost 
part of pectoral-fin membrane with a single straight ray which 
appears to be median but actually originates from left or right 
pelvic fin (pelvic fin contributing this ray with one more ray 
than pelvic fin of opposite side). Ventral surface of anteriormost 
11-12 pectoral- and 8 pelvic-fin rays with thick adhesive pads. 

Dorsal surface of head and most of body above pectoral fins 
with small, widely spaced and almost uniformly distributed 
tubercles; posterior portion of body with a few very small scat- 
tered tubercles. Dorsal surface of pectoral fin with a single row 
of 1-6 tubercles superficially similar to those on head and an- 
terior portion of body including pectoral-fin base; first and gen- 
erally largest tubercle lying at or near base of ray, others di- 
minishing in size and confined to basal half of ray. Dorsal surface 
of some pelvic-fin rays with single row of very small tubercles. 
The general nature and distribution of tubercles in the holotype 
and paratype suggests they are females; males of most Bornean 
gastromyzontins have densely packed rows of numerous tuber- 
cles on the pectoral fin, and some have peculiar comb-shaped 
or ctenoid tubercles on the cheek or pectoral-fin base, neither 
of which occur in these specimens. 

Dorsal surface of head with numerous small dark round spots. 



ROBERTS- WESTERN BORNEO FISHES 



93 




Figure 72. Hypergaslromyzon hiiniilis Kapuas 1976-30. 35.7 mm (MZB 3480, holotype). 



Dorsal surface of body with nine regularly spaced, broad trans- 
verse dark bars alternating with pale intervals, superficially sim- 
ilar to coloration of Homaloptera and very different from that 
in other Bornean gastromyzontins. Pectoral, pelvic, and anal 
fins with faint broad bands; caudal fin with two widely spaced 
dark vertical marks separated by a pale interval with fine mel- 
anophores; dorsal-fin rays uniformly covered with fine mela- 
nophores but interradial membranes without melanophores and 
dorsal fin relatively lacking color pattern. 

Distribution. — Kapuas. 

Etymology.— The name hiiniilis (Latin, on the ground, low) 
refers to the depressed body form of this species. 



Neogastromyzon Popta, 1905 

Neogastwmyzon Popta, 1905:180 {type species Neogaslromyzon mcimenhiitsn 
Popta. 1905, by monotypy). 

Diagnosis.— Gastromyzontins with mouth opening broad, 
almost as wide as head; pelvic fins united posteriorly, each with 
1 5-22 rays. Superficially similar to Gastromyzon but differing 
from that genus in having no sublacrimal groove; rostral and 
maxillary barbels projecting from posterior margin of rostral 
cap (the primitive condition) rather than from its anterior mar- 
gin; and a narrower, more elongate body. 

Neogastwmyzon comprises two species known mainly from 
northern Borneo. A single specimen has been collected in the 
Kapuas basin. 



Neogastromyzon nieuwenhuisii Popta, 1905 

Gastromyzon borneensis Vaillanl. 1902:18 (nee Giinther; Haul-Sibau. Rapuas). 
Xeogaslromyzon meuwenhmsu Popta. 1905:181 (type locality Howong River, 

Mahakam hasin). 
Gaslroinyzon meuwenhmsi Weber and de Beaufort. 1916:4. 

Material Examined —Western Borneo: "Silau" [=Sibau, upper Kapuas], 20.3 
mm (RMNH 7795), Sarawak: Sadong R., 29.3 mm (CAS-SU 68827); Batang Ai, 
Balang Lupar basin, 6: 34.5-38.9 mm (CAS 53172-53173), 

The 20.3 mm specimen from Sibau here identified as A', nieu- 
wenhuisii represents the only known specimen of Neogastw- 
myzon from the Kapuas basin. It has 23 pectoral- and 20 pelvic- 
fin rays. Inger and Chin (1961) reported a large number of 
specimens from the Rajang basin, Sarawak. The specimens re- 
ported above from the Sadong and Lupar represent the first 
records for these basins. 

The only other species in the genus is A', pauciradiatus (Inger 
and Chin 1961), from the Rejang basin, characterized by only 
15-16 pelvic-fin rays (vs. 18-21 in A', nieuwenhuisii). 

Distribution. — Sarawak, Kapuas, Mahakam. 



Neohomaloptera Herre, 1944 

Neohomaloptcra Herre. 1944:51 (type species Homaloptera (.Neohomaloptera) 
johorensrs Herre, 1944, by original designation and monotypy; proposed as 
subgenus). 

Neohomaloptera, among the smallest Homalopteridae, is dis- 
tinguished from all other genera by having two maxillary barbels 



94 



CALIFORNIA ACADEMY OF SCIENCES 



instead of only one at each comer of mouth. Superficially it 
resembles some of the smaller species oi Homaloptera. Unlike 
Homaloptera. most of which live in mountain streams or in 
riffles in lowland streams. Neohomaloptera tends to be found 
in vegetation in slow-flowing lowland streams or swampy areas. 

Neohomaloptera johorensis (Herre, 1944) 

Hoiualoplera [Seohomalnpicra) lohnrcnsis Herre. 144-4:31 (type locality brook 

near Simpang Rengam. Johore, Malay Peninsula). 
St'ohomaloplera johorensis Silas. l'S53:20.V 

Material Examined. — Malay Peninsula: brook near Simpang Rengam. Johore, 
2: 18,0-20.5 mm (CAS-SU 39840- .3984 1, holotype and paratype); River Sarong 
Buaya. Johore. 2: 1 8. 7-20. 3 mm (CAS-SU 66426). Western Borneo: Kapuas 1976- 
4. 17,5 mm (MZB 3481) 

This is apparently the smallest species of Homalopterinae. 
Alfred (1969:223) reported 52 specimens from the Malay Pen- 
insula, the largest only 23.7 mm. It differs from all other mem- 
bers of the subfamily by its low pectoral-fin ray counts and in 
having two maxillary barbels on each side instead of only one. 
Its scale counts, 3 1-37, are lower than those of all other species 
except H. tweedwi. which has 32-36. 

Distribution. — Malay Peninsula (Johore). Western Borneo 
(Kapuas). 



Cobitidae 

This family has more species than any other Eurasian fresh- 
water fish family except Cyprinidae. Loaches are perhaps more 
diverse in the Kapuas than anywhere else. All four cobitid 
subfamilies currently recognized are present: Botiinae (one ge- 
nus and three species); Cobitinae (five genera and 12 species), 
Nemacheilinae (two genera and seven species), and Vaillantel- 
linae (one genus and two species). In addition, the strange cobitid 
EUopostoma n!e,^aloniycler. which does not seem to fit into any 
of these subfamilies, is known only from the Kapuas. 

A number of cobitid genera, especially in the subfamily Co- 
bitinae, are poorly defined and ditficult to distinguish. Most of 
the southeast Asian genera and species have not been studied 
osteologically and in a number of instances adequate generic 
diagnoses are unavailable. 

.Apparently all genera of Cobitinae are charactenzed by sexual 
dimorphism in which the pectoral fins of males are enlarged 
and otherwise modified. The pectoral-fin ray most highly mod- 
ified may be either the second (first branched) ray or the inner- 
most (medial) ray. One new species from the Kapuas has males 
with bony serrations on the second pectoral-fin ray, a special- 
ization previously unreported. Other males with modified pec- 
toral-fin rays bearing variously shaped bony projections or mul- 
ticellular keratinous tubercles (breeding tubercles). As this sexual 
dimorphism has not been well studied, an effort has been made 
to record it here and describe some of the interspecific differ- 
ences that occur. 

Blind loaches have been reported from caves in Iran, Iraq, 
and China, but the blind Lcpidocephalus spectrum described 
herein is the first known blind loach that lives in a normal 
riverine habitat. 

The number of species known in the family Cobitidae has 
grown enormously since Linnaeus (1758) published his account 
of the "genus" Cobitis with five species. Van Hasselt (1823) 
reported the discovery of five new species in Java, some of which 



occur also in Borneo, and made one of the earliest efforts to 
split the Linnacan genus Cohnis. Van Hasselt's names pose some 
serious nomenclatural problems which I have attempted to re- 
solve in the following account. Bleeker, Vaillant, Fowler, and 
others have described additional new genera and species from 
the East Indies or Indonesia and Borneo, and additional taxa 
undoubtedly remain to be discovered there. In the present ac- 
count one genus and six species are described as new. Another 
new species discovered during the Kapuas survey of 1976, Ne- 
Diachcilus kapiiascnsis, has been described by Kottelat (1984). 
It appears that information on vertebral number is helpful in 
distinguishing various genera and species of loaches. Data on 
vertebral counts of loaches from southeast Asia and elsewhere 
are presented in Table 4. Abdominal and caudal as well as total 
counts are given, which is much more informative than total 
counts only. The table shows that Lepidocephalichthys and Ne- 
macheilm have characteristic vertebral formulae exhibited by 
most or at least many of the species. The similarity of the ver- 
tebral counts or formulae oi Acanthopsoides gracilis and Acan- 
topsis choirorhynchus may indicate that they are in fact closely 
related, as suggested by their superficially similar appearance. 
Phyletic isolation of Barbucca. EUopostoma. Lcpidocephalus, 
and I aillantella is supported by their distinctive vertebral for- 
mulae. Barbucca. possibly the smallest nemacheilin loach, has 
the lowest vertebral counts known in Cobitidae. 

Acanthopsoides Fowler, 1934 

Acanthopsotdcs Fowler. l934/i;l03 (type species Acanthopsoides gracilis Fowler, 
1934. by original designation and monotypy), 

Di.AGNGSis. — Superficially similar to Acantopsis. but snout less 
elongate, suborbital spine only slightly anterior to eye, and dor- 
sal fin with only 8-9 rays, its origin posterior to a vertical through 
pelvic-fin origin. Vertebrae 40-42. 

A single known species. 



Acanthopsoides gracilis Fowler, 1934 

Aoanlhopsoides gracilis FowJer. |934/i:103. fig. 55 (type locality "Chien Mai, 
North Siam" [Mekong basin]). 

Material Examined, — Western Borneo: Kapuas 1976-10. 3: 27.6-44.8 mm 
« AS 49342. MZB 3506): Kapuas 1976-2 I. 26,9 mm (MZB 3507); Kapuas 1976- 
22, 7 34,8-16.0 mm (CAS 49343. MZB 3508); Kapuas 1976-25. 45.0 mm (MZB 
1509), Kapuas 1976-33. 28.3 mm (MZB 3510); Kapuas 1976-37. 42: 25.5-42.3 
mm (BMNH 1982,3.29,127-131. CAS 49512. FMNH 94232. IRSNB 19744. 
KL'MF 2853. MZB 3511); Kapuas 1976-39. 78: 21,6-44,1 mm (ANSP 152021. 
(_ AS 49344. MNHN 1982-697. MZB 3512. RMNH 28873. UMMZ 209902. 
USNM 230261. ZMA I 16.536); Kapuas 1976-43.2: 23.2-35.5 mm (CAS 49345, 
MZB 3513): Kapuas 1976-51. 2 26 1-27,8 mm (CAS 49346. MZB 3514). 

Acanthopsoides gracilis occurs sympatrically with Acantopsis, 
and has sometimes been mistaken for its young, although the 
two are readily distinguishable. The largest recorded .-1. gracilis 
IS only 54 mm. Acantopsis attain at least 225 mm. 

Sexual dimorphism has not been reported previously. The 
senesof42 specimens from Kapuas 1976-37 comprises 13 males 
25.5-28.3 mm and 29 females 33.0-42.3 mm. Males have en- 
larged, upcurved pectoral fins, with the second ray about three 
times thicker than the others. On the distal third of the second 
ray, the branched portions of the ray each bear a single irregular 
row of minute breeding tubercles. Such a clear-cut difference in 
male-female body size is unusual for Cobitinae, and may not 



ROBERTS-WESTERN BORNEO FISHES 



95 



be typical of this species. The sample from Kapuas 1976-25 
consists of a single much larger male (45.0 mm) with a similarly 
modified second pectoral-fin ray. 
Distribution.— Thailand (Mekong). Borneo (Kapuas). 

Acantophthalmus van Hasselt. 1823 

Acantophthalmus van Hasselt. 1 823: 1 .33 (type species Acaninphthalmus fascmlus 

van Hasselt, 1823=Coft///s kuhlii Valenciennes in Cuvier and Valenciennes, 

1846, by subsequent designation of Bleeker. 1863t':38). 
Acanlhophthalmus Bleeker. 1858-59(':304, 1860a:73 (unwarranted change in 

spelling). 
Pangio Blyth, 1860:169 (type species Cohilis paiigui Hamilton-Buchanan, 1822, 

by monotypy)- 
CoNlophis Myers, 1927:4 (type species Acanlhophthalmus angiullans Vaillant, 

1902, by original designation). 
lEucirrhichthys Perugia, 1892:1009 (type species Eiiciirhichthys donae Perugia, 

1892, by monotypy). See Remark under Acantnphlhalnius angiullans below. 

NoMENCLATURAL NoTE. — In attempting to determine the 
original authority, type species designation, and proper spelling 
of this genus one has to deal with Acantophthalmus van Hasselt 
(1823:133) which has been variously misquoted, misinterpret- 
ed, or overlooked by others. This represents one of the most 
difficult of the nomenclatural problems posed by the publication 
of van Hasselt (1823). It is perhaps well to recall here that his 
account was never intended for publication; it does not represent 
his efforts at formal description, but merely an attempt to ex- 
plain in a private letter to Temminck the extent and nature of 
the Javanese fish collections obtained by Kuhl and him. 

The paragraph in which Acantophthalmus is treated may be 
cited in full: 

Cobnis OaocinhusJah. nostra. InvroetZunda. Kitjaeraeheeft. met 
de Cohiiis taenia Linn., dat gemeen, dat hij onder het oog eenen be- 
weegbaren doom heeft: dit teeken komt mij voor een gewigtig te zijn, 
als vcrschiedenen karakteren in de genera der Cypriniden door Cuv. 
opgesteld, en zou alzoo als eenen afdelling van Cobitis, den naam 
Acantophthalmus verdienen. De zonderlin structuur der luchtblaas, is 
geheel die der Cobiten en hij leeft als deze in kleine beken; ik reken 
totdezen Acantophthalmusnog 2 specien, diczieh alleenonterschieden 
door eene meer afterwaarts geplaatse dorsaal-vin. Zij dragen in de 
teekeningdenamen foifw/HS en iuiu/»ci«Mihi Rambockassangbeide 
in het Zundasche. 

An English translation of this paragraph, prepared with the 
help of M. Boeseman, was published by Alfred (1961:86), and 
may also be quoted here: 

Cohitis Octocirrhus Tab. nostra. Irivroet in Sundancse. K.itjaerae 
has. in common with Cohilis taenia Linn, a moveable spine below the 
eye; this character seems to me to be as important as various characters 
in the genera of Cypnnes erected by Cuv., and as a section of Cobitis 
would consequently deserve the nami; Acantophthalmus. The unusual 
structure of the air-bladder is exactly as that of the Cobites and it lives 
just like these in small brooks. To this Acantophthalmus I reckon 
another 2 species which distinguish themselves solely by a more back- 
wardly placed dorsal fin. In the drawing they bear the names fasciatus 
and Javaniciis Mihi both Rambockassang in the Sunda region. 

Alfred did not analyze the nomenclatural significance of this 
paragraph, nor has anyone else to my knowledge. The name 
Acantophthalmus occurs in two other places in van Hasselt 
(1823), always with the same spelling, but without nomencla- 
turally significant remarks. The nomenclatural status of Acan- 
tophthalmus. A. fasciatus. A. javanicus, and Cobitis octocirrhus 
of van Hasselt (1823) must all rest solely on this one paragraph. 
No taxonomic character(s) are given to validate any of the three 



new species: Cobitis octocirrhus is not distinguished from Cobitis 
taenia Linnaeus, and Acantophthalmus fasciatus and .-1. java- 
nicus are not distinguished from each other. Had the figures 
referred to by van Hasselt been published, they undoubtedly 
would have sufficed for species recognition. 

The status of Acantophthalmus van Hasselt (1823) may now 
be considered. Van Hasselt undoubtedly intended it to include 
the three new species from Java as well as the European Cobitis 
taenia Linnaeus, 1758, but not the other species placed in Cobitis 
by Linnaeus (anableps. harbatulus, fossilis). The character of a 
subocular spine was in fact recorded by Linnaeus, but was used 
by him only to distinguish C. taenia ("Cobitis aculeo bifurco 
infra utrumque oculum") from the other four species of his 
Cobitis which all lack the spine. We know now that the spine 
occurs in the majority of species in the large subfamily Cobitinae 
and in all species of Botiinae. 

So far as I have been able to determine, a valid type species 
has never been designated for Acantophthalmus. Bleeker ( 1 863: 
38) designated Acantophthalmus fasciatus van Hasselt, 1 823 as 
type species, but this is a nomen nudum. It is believed that 
Cobitis kuhlii Valenciennes, 1846 in fact represents the original 
A. fasciatus. LInder ".icanthophthalmus [sic] van Hasselt" Jor- 
dan (1917:116) stated "type .-icanthophthalmus fasciatus van 
Hasselt is unavailable (Code, Article 12).'" There is no discussion 
of the nomenclatural history of the genus Acantophthalmus and 
no mention of a type species in the important paper on the 
species of the genus by Fraser-Brunner (1940). 

A strict interpretation of the Code would seem to require that 
Cobitis taenia Linnaeus be recognized as type species of Acan- 
tophthalmus. since it is the only one of the species included by 
van Hasselt which is nomenclaturally available. Since C taenia 
is the type species of Cobitis. .Acantophthalmus would become 
a junior objective synonym of Cobitis. 

The original spellings .Acantophthalmus and .Acantopsis by 
van Hasselt clearly were intentional. Both names appear re- 
peatedly with no change in spelling. It should be pointed out 
that numerous systematists have regarded "acanto" as a correct 
Latinization (e.g., Hypacantus Rafinesque, 1809; Acantophis 
Berthold, 1827; Acantoceras Pavlov, 1892). In numerous in- 
stances, in fact, earlier names spelled with "acantho" subse- 
quently were corrected to "acanto" (see Neave 1939:12). Thus 
while nearly all subsequent authors have used the spellings 
.■icanthophthalmus and .Acanthopsis for these genera, it seems 
that this is not permissible according to the present Code. Sim- 
ilar problems arise in names in which "opisto" and "opistho" 
have been used interchangeably. It seems that the best way to 
resolve such duality in spelling, at least in most instances, is to 
adhere to the original spelling, as is done here with .Acan- 
tophthalmus and .Acantopsis. 

.Acanlhophthalmus Bleeker, 1 859, automatically takes the same 
type species as .Acantophthalmus van Hasselt, 1823 (Code, Ar- 
ticle 67(i)(ii)), 

Two Bornean species placed until now in this genus. A. lo- 
rentzi Weber and de Beaufort, 1916, and .A. sandakanus Inger 
and Chin, 1962, are herein referred to Leptocephalichthys. 



Acantophthalmus anguiliaris Vaillant, 1902 

.■icanthophthahms angiullans Vaillant, 1902:151 (type locality Kapuas). 
.icanthophthalmis vermiculans Weber and de Beaufort. 1916:34 (type locality 



96 



CALIFORNIA ACADEMY OF SCIENCES 



Table 4. Frequencies of Vertebral Counts in Cobitidae. For definition of vertebral counts see p. 22. 



Aicinrhopsoidcs gnicilis 

Kapuas 1976-22 
Acanlophlhalimii angiiillarn 

Perak (paralypes C. perukt'iisn) 

Kapuas 1976-21 

Acanlophthalmus muraenilonuis dc Bcaulorl. 1933 
Singapore (CAS-SU 32602. paratype) 
Johore (CAS-SU 32601) 

AcaiUophlhalnnis nnvrsi Harry, 1949 

SE Thailand (CAS-SU 14888, paratype) 
Acunlcphlhalmus ohiongus 

Kapuas 1976-29 
Acanliiphllialmus pangia (H-B.. 1822) 

Nepal, Chitawan {CAS 50180) 
Aianliiphlhahtuis cf. pangia 

Burma. Mitkyina (CAS-SU 32618) 
Aiiinliiphlhalnuis scmiciihliis 

Johore (CAS-SU 32611) 

Kapuas 1976-17 

Acanlophlhalmus shelforJi 

Kapuas 1976-16 
AiatilDphlhahiiiii ^iipcrbiis 

Kapuas 1976-17, paratypes 
Acanlopsis L'ht>iriirhvnchos 

Kapuas 197b-35 
Aiaiilnpsis lachnosloma Rutter, 1897 

S, China, Swatow (CAS-SU 1812, holotype) 

Barbucca diabolua 
Kapuas 1976-37, paratypes 

Bona hymcnophysa 

Kapuas 1976-.36, 39 
Boua macracanlha 

Kapuas 1976-19 

Boua reversa 
Sumatra, paralypes 
Kapuas 1976-27, 29, paratypes 
Mahakam. paratypes 

EUopiKtoma megalomyclcr 

Kapuas 1976-14 
Lepidocephaluiilhvi gunica (H.-B,. 1822) 

Nepal (CAS 50532) 
LcpidiKcphalichrhys hassclli 

W, Java (CAS-SU 20499. 24094. LIMMZ 155673) 

l.cpicl( Kcphalichlhys lorcmzi 

Kapuas (ZMA 103.259, holotype, ZMA 116.551) 
Liputiiccphalichilns prislcs 

Kapuas 1976-16, 42, paralypes 
Lcpuloccphalichthys sandakanensn (Inger and C hin, 1962) 

N Borneo, Sandakan (CAS-SU 49967, paratypes) 
I.cpuJiiLCphaliLs macrnchir 

Kapuas (RMNH 7783, holotype L pallcns) 
Lcpuii Kcphahi^ spectruni 

Kapuas 1976-48, paratypes 
Ncinacheiliis lascialus (Val. m Cuv and Val., 1846) 



27-28-H2-14=40(2). 41(3). 42(1) 



50-H9-20=69(2). 70(1) 
50-52-1-19=69(1), 70(1), 71(1) 

33-H5=48''(l) 
34-35+14-16=49(1), 50(5) 



35 -fl 3=48(1) 

33-35+12 = 45(1), 46(8), 47(2) 

50(1) 

40+14-15 = 54(1), 55(1) 

34-37+12-15=46(1), 48(1). 49(4). 50(2). 51(1) 
33-35 + 1 3-14=46( 1 ), 47( 1 ), 48(9), 49(2) 

34-36+14-15=49(1). 49(3), 50(1) 

35-36+ 15-16 = ,50(1), 51(2), 52(1) 

28-29+13-14=42(2) 

29+15=44(1) 

19-20 + 8-9 = 28(7), 29(6) 

20-21 + 12-13 = 33(6) 

19-20+10-11=30(3) 



20-21 + 12-14 = 33(3), 34(1) 
21-22+12-13 = 33(5), 34(4) 
20-21 + 12-13 = 33(3) 



24-25 + 9 = 33(1), 34(1) 
24-25+11=35(1), 36(3) 
24-25+11-12 = 35(1). 36(4) 
25-26+1 1-12 = 36(1). 37(1). 38(1) 
23-24 + 9-10=33(4). 34(1) 
23-25+11-13 = 35(1), 36(4) 
28+15 = 43(1) 
28+14-15=42(1). 43(1) 



ROBERTS-WESTERN BORNEO FISHES 



97 



Table 4. Continued. 



Java, Buitzenzorg (CAS-SU 20498) 
Neinacheilus kapuasensis 

Kapuas 1976-29 
Nemacheilus laclogeneus 

Kapuas 1976-45, types 
Nemacheilus maculiceps 

Kapuas 1976-27. paratypes 
Nemacheilus saravacensis 

Kapuas 1976-8 
Nemacheilus selangoncus 

Kapuas 1976-39 

Nemacheilus cf. hmgipecloralis 

Kapuas 1976-45 
Vaillantella euepiptera 

Kapuas 1976-37 

Vaillantella maassi 
Pahang. R. Tahan (CAS-SU 50180, paratypes \' llavokisciala) 
Kapuas 1976-29, 50 



20-23-1-12-13=33(1), 34(10), 35(4) 

22-H 1-13 = 33(1), 34(6), 35(1) 

24-H4=38(2) 

20-22 + 11-13 = 33(5), 34(4) 

22+12 = 34(1) 

21-22+11-12=33(101,34(10) 

23+12=35(1) 

33-35+19-21=52(2), 53(2), 54(2) 



39-10 + 20=59(1), 60(1) 
38+19=57(2) 



"Sumatra, river Kampar kiri"). See Smilh (1945:302), Inger and Chin (1962: 

119-120), and below. 
''Eucirrhiclnhys doriae Perugia, 1892:1009 (type locality Sarawak). See Tweedie 

(1956), and below. 
Cohitpphis anguillans Myers, 1927:4. 
Cobilophis perakensis Herre, 1940n:8 (type locality lake above Chendcroh Dam, 

Perak, Malay Peninsula). See Mora (I941a:49-51), Tweedie (1956:57), and 

below. 

Material Examined. — Malay Peninsula: 4: 46.1-60.5 mm. lake above Chen- 
deroh Dam, Perak (CAS-SU 33004-5. holotype and paratypes o( Cobilophis pe- 
rakensis). Western Borneo: Kapuas 1976-21. 3: 70.2-76.8 mm (CAS 49334, MZB 
3482, USNM 230255). 

Diagnosis. — By far the most elongate Acantophthalmus. with 
body depth 14 or more and vertebrae 69-7 1 (versus body depth 
10 or less and vertebrae 45-55 in all other species). 

The three Kapuas specimens of this species are two males, 
71.0-76.8 mm, and a female, 70.2 mm. The males have large 
upturned pectoral fins, much larger, about twice as long as those 
of the female, and with the second fin ray greatly thickened. A 
faint, dusky midlateral stripe runs length of body, the upper 
portion of which is covered with fine, more or less uniformly 
distributed spots. In the female, anal-fin origin directly below 
base of last dorsal-fin ray; in the two males it is below the base 
of the second or third from last dorsal-fin ray. Principal caudal 
rays 8 + 8(2), 8 + 7(1). Female pectoral-fin rays 8; anal-fin rays 
1 ,6; pelvic-fin rays 6. Male pectoral-fin rays 7 or 8; anal-fin rays 
2,6; pelvic-fin rays 6. All four specimens in the type series of 
C. perakensis (SU 33004-5. 46.1-60.5 mm SL) appear to be 
females (or immature males) with small pectoral fins; the origin 
of the anal fin varies in position from below about the fourth 
to the second from last dorsal-fin ray. 

Myers (1927) distinguished Cobilophis (type species Acan- 
tophthalmus anguillans Vaillant) from Acantophthalmus on the 
basis of its having an extremely attenuated, anguilliform body, 
and some part of the anal fin under the dorsal fin. Myers included 
Acantophthalmus vcrmicularis Weber and de Beaufort (1916: 
34) from Kampar Kiri R., Sumatra, in Cobilophis. A third species, 
C perakensis Herre (1940:8) was described from Chenderoh 



Dam. Perak. Malay Peninsula. In these species the body depth 
is reported as 1 4.3- 1 8 in standard length, whereas in olher Acan- 
tophthalmus known at the time of Myers' description, it is about 
7-10. 

Conclusion.— Acantophthalmus anguillans and .4. pera- 
kensis are identical. Females of .4. anguillaris from the Kapuas 
agree closely with the entirely female type series of .-1. perakensis 
in relative position of dorsal and anal fins and in body depth, 
the only characters cited to differentiate the species by Herre. 
and I have not found any other significant difierences between 
them. .4. vcrmicularis. based on a single 70 mm specimen, pos- 
sibly a male, is probably also a junior synonym of.-l. anguillaris; 
It is described as having "last dorsal ray above first anal ray." 

Remark. — If the synonymy is correct, .1. anguillans includes 
individuals with and without more or less elongate barbels. The 
head or cheeks are usually scaleless, so far as observed or re- 
ported. Eucirrhichthys doriae Perugia, 1892:1009 (Weber and 
de Beaufort 1916:27, fig. 10), from Sarawak has a long nasal 
barbel and head extensively scaled, but is otherwise similar to 
.-I. anguillans. Since nasal barbels and head squamation occur 
repeatedly in Cobitinae they can be regarded as primitive char- 
acters for the group, and their presence or absence should not 
be used to define genera. It seems probable that E. doriae is 
congeneric and possible that it is conspecific with .-(. anguillaris. 

Distribution. — Malay Peninsula (Tahang, Kelantan, Perak) 
(Hora 1941). Sumatra (Kampar Kiri River). Western Borneo 
(Kapuas). Sarawak? North Borneo (Tawan district) (Inger and 
Chin 1962). Thailand (Fowler 1934). 



Acantophthalmus oblongus (Valenciennes, 1846) 
new combination 

(Figure 73) 

Acanlnphlhaliiiiis javainctis van Hasselt. 1823:1 33 (nomen nudum). 

Cohiiis ohlonga Valenciennes in Cuvier and Valenciennes, 1 846:76 (type locality 

Java; description based on van Hasselfs unpublished figure of .4. javanicus). 
Acanthophthahmis lanuiiciis Bleeker. 1860:75. 



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CALIFORNIA ACADEMY OF SCIENCES 




Figure 73. Acanlnplnhalnius oblotigns Rapuas 1976-29, 62.6 mm male (CAS 49335). 



'' \caulhophthalmus luanue Inger and Chm. 1962:1 18, tig. ?6 (Kinabatangan R. 
at Dcramakol) 

Material Examined. — Western Borneo: Kapuas 1976-29, 13: 49.4-62.6 mm 
(C.JiS 49335, MZB 3483, RMNH 28870, USNM 230256); Kapuas 1976-47, 39.2 
mm (MZB 34841 

Acanioplulialniiis oblongiis and the extremely elongate A. an- 
guillans are perhaps the only plain colored Acanlophthalmiis in 
the East Indies. Specimens from Borneo, Sumatra (Weber and 
de Beaufort 1 9 1 6:32) and Java should he compared to determine 
whether they are in fact conspecific, Weber and de Beaufort 
(1916:31) placed this species as synonym oi Acanthopbthalmus 
pangia. a plain colored species from Burma and India, hut they 
apparently are not identical. This is the first record of A. oblongus 
from Borneo. Most of the Kapuas specimens were uniformly 
dark brown in life; a few were pale tan or gray. 

In sexually mature males the pectoral fin is enlarged, due 
mainly to the elongation and widening of the second ray. The 
lepidotrichia of this ray are very numerous and broad, and the 
distal half of its dorsal surface is covered with a patch of hundreds 
of small, densely packed tubercles. 

Distribution. — If the synonymy given here is correct, the 
species is distributed in .lava, western Borneo, Sumatra, and 
probably, eastern Malay Peninsula (for the latter see A. pangia 
in Hora 1941:47). 

Acantophlhalmus semicinctus Fraser-Brunner, 1940 

ALUnlhiiphthalmus scinicinclus Fraser-Brunner, 1940 172 (type locality Mawai 
district, Johore, and Malacca, Malay Peninsula). 

Material Examined. — Malay Peninsula: Johore, Mawai district .A. W. Herre. 
27 Fehruarv 1937, 38: 24.7-53.8 mm (CAS-SU 3261 1; non-types, but probably 
part of the series from which holotype was drawn); Johore, 5 miles north of Rota 
Tinggi, 4: 35.3-49.8 mm (CAS-SU 32612); Johore, near Kulai. 3: 48,6-52.5 mm 
(CAS-SU 39392); Johore, Kota Tinggi, 8; 23.7-52.5 mm (CAS-SU 39391); Ma- 
lacca, Lake Chin Chin Jasin, 1 55 7 mm (CAS-SU 31078). Western Borneo: 
Kapuas 1976-16, 3; 41. .3-48.9 mm (BMNH 1982.3.29. 121-2, MZB 34851; Kapuas 
1976-17, 17; 33.3-51.0 mm (CAS 49336, FMNH 94230, MZB 3486. RMNH 
28871, UMMZ 209858); Kapuas 1976-39, 25.5 mm (MZB 3487); Kapuas 1976- 
47,4: 40, 1-48.0 mm (MNHN 1982-695, MZB 3488); Kapuas 1976-51. 7: 24.7- 
42.9 mm (MZB 3489, IISNM 230257). 

Diagnosis. — Until now this species has been known only from 
the southern Malay Peninsula. 

In A. semicinclus the pale marks on the dorsal surface of the 
body are usually entirely transverse (usually complete but some- 
times partial), or transverse alternating with small round spots 



(never longitudinal). When these spots are oval (i.e., elongated), 
they are oriented transversely. 

In two 52.5 mm males from Kota Tinggi the pectoral fin is 
greatly enlarged and upcurved. The second ray is thickened. The 
skin on the dorsal surface of the branches of the second pectoral- 
fin ray is greatly thickened, and in the depression thus created 
between the branches lies a series of about 15 transverse, ex- 
tremely regular, tightly seried rows of elongate breeding tuber- 
cles. 

None has a separate row of smaller marks ventral to the marks 
covering the dorsal surface of the body. 

Acantophlhalmus shelfordi Popta, 1903 

Acanlhophlhalnius shclfurdi Popta, 1903:231 (type locality Sarawak, river near 
Kuching). 

Material Examined— Western Borneo: MZB 3490, 45.2 mm, Kapuas 1976- 
6. MZB 3491. 31.8 mm, Kapuas 1976-7; BMNH 1982.3.29.123-126 and MZB 
3492 (1). 5: 31.4-52.3 mm; Kapuas 1976-8; CAS 49337 (6) and MZB 3493 (I), 
7: 30.2^6.2 mm, Kapuas 1976-12; MZB 3494. 33,3 mm. Kapuas 1976-13; FMNH 
94231 (4) and MZB 3495 (I), 5; 44,2-51,0 mm, Kapuas 1976-16; IRSNB 19743 
(3) and MZB 3496 (1). 4; 30.3-53.8 mm, Kapuas 1976-17; MZB 3497,26.8 mm, 
Kapuas 1 976-2 1 ; MZB 3502, 317 mm, Kapuas 1 976-37; CAS 49338 (8), MNHN 
1982-696 (4), MZB 3498 (10), RMNH 28872 (4). UMMZ 209901 (4), USNM 
230258 (4). and ZMA 1 16 535 (4). 38: 24,0-52.6 mm, Kapuas 1976-39. 

This species has an extremely variable color pattern. 
Distribution. — Malay Peninsula. Sarawak. Kapuas. 

Acantophthalmus superbus new species 

(Figure 74) 

Holotvpe. — MZB 3499, 51.5 mm. Kapuas basin, small forest stream about I 
km up Sungai Tajan from Tajan. 87 km E of Pontianak (Kapuas 1976-51). 

Parj\types.-CAS 49339, 3; 32.7-34.6 mm, collected with holotype, MZB 3500 
( I ) and LISNM 230259 (2), 3; 42.5-47. 1 mm. Sungai Tekam near Kapuas main- 
stream 5-6 km upstream from Sanggau (Kapuas 1976-16); CAS 49340 (3) and 
MZB 3501 (I). 4: 48.1-57.1 mm, several small forest streams near Kapuas main- 
stream within 10 km upstream of Sanggau (Kapuas 1976-17). 

Diagnosis, — Distinguished from all other Acantophthalmus 
by its distinctive coloration. 

All specimens have a row of about 10-11 dark saddlelike 
marks on head and body; in smaller specimens all of the marks 
tend to be discrete or but narrowly joined, while in larger spec- 
imens those on the body tend to be at least partly joined. In 
specimens of all sizes the dark marks are totally or partially 
separated at the dorsal midline by a pale area; in small specimens 



ROBERTS-WESTERN BORNEO FISHES 



99 






Figure 74. Acantophthalmus superbus. Kapuas 1976-51. a. 51,5 mm (MZB 3499, holotype); b. 34.2 mm (CAS 49339, paratype). 



the pale area tends to form a series of short, separate longitudinal 
stripes and transverse bars; in larger specimens it is more van- 
able, but generally more continuous, and may form a serpentine 
middorsal stripe (Fig. 74). In live specimens the dark areas are 
purple or magenta, and the pale areas, below as well as above 
the dark ones, are orange. All other species oi Acantoplnhalmus 
known from the Kapuas are relatively drab. A. superbus tends 
to have more vertebrae than other Acantophthalmus from the 
Kapuas except A. anguillans (Table 4). 

Etymology.— The name superbus (Latm) refers to the beau- 
tiful coloration of this species. 



Acantopsis van Hasselt. 1823 

Acantopsis van Hasselt, 1823:1 33 {lypc species .-l(-'a'7/cp5/5 dialuzona van Hassell. 

1823, by monotypy). 
Acanthopsis Bleaker 1858-59(':303 (unwarranted emendation). 
As pointed out by Alfred (1961:86), the description oi Acaniopsts given by van 

Hassell appears to be sufficient for its recognition. 
Note. — Concerning the spelling of .-ifaii/opiw see note under Actiiilophlhalmiis 

Diagnosis. — Distinguished from other Cobitinae by its great- 
ly elongated head and snout, dorsally situated eyes, and sub- 
orbital spine entirely far anterior to eye (instead of below eye, 
or rarely, slightly anterior to it). Dorsal fin with 10-13 rays, its 
origin anterior to vertical through pelvic-fin origin. Caudal fin 
forked or deeply emarginate. Vertebrae 40-44. 



Acantopsis choirorhynchos (Bleeker, 1854) 

Cohilis choirorhynchos Bleeker, 1 854u:95 (type locality Sumatra, Palembang "ubi 

confluunt flumina Lamantang el Emm"). 
Acanlhopsis choirorhynchos Bleeker, 1860u:66, 

Material Ex amtned. — Western Borneo: Kapuas 1976-9, 1 1 1 mm (MZB 3503), 
Kapuas 1 976-22. 93.2 mm (CAS 44 1 84): Kapuas 1976-35, 4: 84.2-1 1 3 mm (CAS 
49341. MZB 3504); Kapuas 1976-36, 2: 70.2-75.7 mm (MZB 3505, USNM 
230260). 

Following Smith (1945:296) and Alfred (1961), I tentatively 
treat A. choirorhynchos as distinct from A. diahizona. Further 
study of the species oi Acantopsis is needed. 

In sexually mature males of .-J, choirorhynchos the pectoral 
and pelvic fins are greatly enlarged; the second pectoral-fin ray 
IS greatly thickened, with hundreds of tubercles on its dorsal 
surface and thousands of unicellular homy projections or unculi 
on its ventral surface (Roberts 1982a:66, fig, 12e, 14), 

Chen (1981:22) placed Acantopsis lachnostoma Rutter, 1897 
as a junior synonym of.-!, choirorhynchos. thus extending the 
range of this species to southern China. I have examined the 
holotype of, -J. lachnostoma {C \S-SV 1812, 127 mm ripe female 
from Swatow, Kwangtung Prov,) and compared it directly with 
two Kapuas specimens of A. choirorhynchos 96-101 mm. The 
two laxa are congeneric but not conspecific. Acantopsis lach- 
nostoma has the large, elongate head, dorsally situated eyes, 
suborbital spine anterior to eye. and oral cavity lined with villus- 
like papillae diagnostic of the genus Acantopsis. On the other 
hand the head is somewhat shorter, snout less pointed, caudal 



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CALIFORNIA ACADEMY OF SCIENCES 




Figure 75. harhmca dtabolua Kapuas 1976-37. 23.2 mm npe female (MZB 3536. hololype). 



peduncle much deeper, dorsal-fin rays fewer, and vertebrae ap- 
parently more numerous than in A. choirorhyiichos or any other 
member ofAcamopsis. The hoiotype of .-1. lachnosloma has head 
4.7, caudal peduncle depth 13.1, dorsal-fin rays iii7-l/2. and 
vertebrae 29+ 15=44 (vs. head 3.8, caudal peduncle depth 22.3- 
23.0, dorsal-fin rays iiil 0-1/2 and vertebrae 28+14=42 in Ka- 
puas A. iiwirorhynchos). It seems that .1. lachiioslonia is a valid 
species endemic to southern China; it may be the most primitive 
species ot Acanlopsis. 

Distribution. — The systematics of Acanlopsis is too poorly 
known to make meaningful statements about species distribu- 
tions. 



Barbucca new genus 

Type species. — Barbucca dhih'lica new species 

Diagnosis. — Related to Nemacheihis but distinguished by its 
smaller size; foreshortened body and large, broadly overlapped 
paired fins; peculiarly truncate snout and downtumed jaws; la- 
bial in addition to rostral and maxillary barbels; fewer fin rays; 
fewer vertebrae; and unusual tubcrculation. 

Largest specimen, a gravid female (eggs to 0.25 mm diameter) 
only 23.2 mm (all other Nemacheilinae attaining more than 30 
mm?); lower lips with bulbous posterolateral paired thickenings 
giving rise to moderately elongate labial barbels posteromedial 
to maxillary barbels; homy jaw sheaths very large; fin-ray counts 
almost invariable, dorsal-fin rays usually iii-iv8 (not 8-1/2), and 
ii-iii5, pectoral ilO, and pelvic i6; caudal fin truncate, principal 
rays 9 + 9; pectoral- and pelvic-fin axial flaps absent; vertebrae 
28-29 (all other Cobitidae with 30 or more vertebrae, Ne- 
macheilus with 32-42; see Table 4). 

Breeding tubercles on caudal peduncle exceptionally large, 
present in both sexes, about 6-8 in an irregular row on lower 
half of body extending immediately in front of and above anal- 
fin base (peduncular tubercles, characteristic of most Sundaic 
Nemacheilus. usually present in male sex only, generally asso- 
ciated with dorsolateral portion of caudal peduncle nearer cau- 
dal than anal fin). Some specimens of both sexes with one or 
two moderately large tubercles on gill cover near posterior mar- 
gin. Mature males invariably with a very distinctive, slightly 



concave, or hook-shaped patch of numerous very small, close- 
set tubercles extending below and posterior to middle of eye 
(comparable tubercles unknown in any other cobitids). Sub- 
ocular fleshy papilla absent (present in males of most Sundaic 
Ncniachcilus). No other sexual dimorphism observed. 

Et\molog\.— Barbucca (feminine), from Latin barba (beard) 
and bucca (cheek), in reference to the tubcrculatc cheek patch 
in males. 

Barbucca diabolica new species 

(Figure 7 5) 

HoLOTYPE— MZB 3536. 23.2 mm. gravid female, small forested stream where 
it flows into Sungai Mandai 2-3 km upstream from confluence with Rapuas 
mainstream, 17 km WSW of Putussibau. 10 August 1976 (Kapuas 1976-37). 

Paratypes.-BMNH 1982.3.29.132-134, CAS 49354. FMNH 94233. MNHN 
1982.698, MZB 3537. 29: 17.5-22,3 mm. collected with holot\pe; MZB 3538, 
RMNH 28876. 4: 1 5.6-20.6 mm. Sungai Mandai Ketchil near its confluence with 
Kapuas mainstream. 18 km WSW ofPutussibau. 1 1 .August 1976 (Kapuas 1976- 
39); MZB 3539. USNM 230268, 9: 17.6-20.6 mm, Sungai Seriang, forested 
tributary ofSungai Palin. 37 km W ofPutussibau. 12 August 1976 (Kapuas 1976- 
42). MZB 3540, 22.2 mm. Sungai Gentu. near where it flows into Kapuas main- 
stream. 55 km NE of Sintang, 16 August 1976 (Kapuas 1976-46). MZB 3541, 
ZMA 1 16.537, 4: 14.8-17.9 mm, small forest stream flowing into Kapuas main- 
stream NE of Gunung Selunggul. 53 km NW of Sintang and 10 km NW of Silat, 
16 August 1976 (Kapuas 1976-47); CAS 52591, 21.2 mm. Malay Peninsula. 
.lohorc. J Vierke. ca, 1978 

The most distinctive characteristics of the species are given 
in the generic diagnosis. It may be further characterized as fol- 
lows (proportional measurements from hoiotype): head length 
3.6; eye large, diameter 13.2; interorbital space broad, about 
equal to eye. 12.9; snout 8.3; medial and lateral rostral barbels 
relatively thick but short, their length much less than eye di- 
ameter; maxillary barbel moderately elongate, extending pos- 
teriorly to below middle of eye, its length slightly greater than 
eye diameter; pectoral fin broadly overlapping pelvic fin, its 
length 3.8; pelvic fin length 4.3; origin of dorsal fin slightly nearer 
snout-tip than caudal-fin base, predorsal length 2. 1 ; body width 
(at pectoral-fin origin) 5.3, depth (at dorsal-fin origin) 5.9; depth 
caudal peduncle 9.3, length 8.9. Scales relatively large and thin, 
deeply embedded and difficult to count especially anteriorly and 
dorsally, more exposed and slightly larger on posteriormost part 
of body, approximately 45-50 in lateral series. 

Body with six broad dark vertical bands (rich brown in life) 



ROBERTS-WESTERN BORNEO FISHES 



101 




Figure 76. Bona. Above, B hyint-iwphysa. Kapuas I47b-3V, SU.5 mm (MZB 351(1); below. B rc\i-i\a. Kapuas 1976-27, 87.6 mm (MZB 3521. holot\pe). 



separated by narrow, slightly oblique pale vertical bands (golden 
in lite); dorsal and anal fins colorless; basal half or two-thirds 
of caudal fin with broad vertical bars; thin vertical uniserial rows 
of melanophores overlapping lepidotrichial joints of fin rays; 
pectoral and pelvic fins with dark basal marks on dorsal surface 
but otherwise colorless. In live specimens the eyes shined bright 
red, recalling the tapetum lucidum of nocturnal higher verte- 
brates, a feature looked for but not observed in any other Kapuas 
cobitids. 

Note.— The existence of this species first came to my atten- 
tion during a visit to the Raffles Museum in 1972, when Dr. 
Eric Alfred kindly showed me a single specimen he had collected 
in Johore (present disposition of this specimen is unknown, but 
a second specimen from Johore, CAS 52591, was collected by 
Dr. Jorg Vicrke). 

Etymology.— The name diaboiica alludes to the glowing red 
eyes and spiked tail characteristic of the species. 

Distribution. — Malay Peninsula (Johore). Western Borneo 
(Kapuas). 

BotiaGray, 1831 

Bulla Gray, 1831:8 (type species Bolia aliuurhac Gray, 1831, by monolypy). 

Ilviucnphysa McClelland. 1839:443 (type species not designated). 

Sviicrossiis Blyth, 1860:166 (type species .Syiicn>.\.siis hcnliimrci BIyth. 1860. by 

monolypy), 
Ilrmcnophvsa Bleeker. 1858-59c:303; Giinther, 1868(j:366 (unwarranted spelling 

emendation oi Hyinenphysa McClelland, 1839). 



Botia hymenophysa (Bleeker, 1852) 

(Figure 76) 

Cohitis hymenophysa Bleeker, 1852i/:602 (type locality Palembang, in lluviis), 
Ihnicnnphysa MacClellandi Bleeker. 1 858-59/:358 (unwarranted substitute name 

for Cohins hymcnophrsa). 
Bona hrmcnophysLi Bleeker, 186.3-64:6 

Material Examined. — Western Borneo: Kapuas 1976-36, 4: 95,3-148 mm 
(CAS 49347, MZB 3515, RMNH 28874): Kapuas 1976-39, 2: 80.5-84.8 mm 
(MZB 3516, USNM 230262). Southeastern Borneo: Moara-Teweh. Banto-Fluss. 
2: 85.0-126 mm (NMW 48528). 

Diagnosis, — A Botia with complex vertical bars consisting 
of alternating pale and dark bands separated by darker thin 
vertical streaks; as pale bands generally somewhat wider than 
dark ones, effect is 12-14 pale bands with very dark margins 
on a slightly dark background. Dorsal-fin branched rays 12- 
1/2 or 13-1/2. 

Fin rays of all fins without melanophores except where tra- 
versed by features of color pattern; dorsal fin with a solid dark 
submarginal blotch on uppermost portion of last unbranched 
and first three branched rays and their interradial membranes. 
Rest of dorsal fin with about six evenly spaced oblique dusky 
bands breaking up into spots on interradial membranes and fin 
rays; middle portion of caudal fin with 5-6 narrow vertical dusky 
bands of fine melanophores; pectoral, pelvic, and anal fins clear 
or with relatively few fine melanophores on interradial mem- 
branes. 



102 



CALIFORNIA ACADEMY OF SCIENCES 



70° 



80° 90° 



100° 110° 120° 130° 



-30° 




-20° 



-10° 



▲ hymenophysa 
• macracantha 
▼ reversa 



'^^^£5, 



■Oc^ CCS'' ^^ 

"^ U -10° 



I 



'III! 
Figure 77. Bona Geographical distribution ot the three species found in Borneo, 



Proportional measurements on two specimens 98.5-124 mm 
from Kapuas 1976-36 (CAS 49347) arc head 3.1; snout 5.4- 
5.7; eye 2 1 .9-23.8; body depth 4,2; body width 8. 1-10.05; depth 
of caudal peduncle 6.3-6.5; predorsal length 1.7. Kapuas spec- 
imens have dorsal branched rays 12-1/2(5) or 13-1/2(1); pec- 
toral rays 15(1). 16(1); pelvic rays 8(2). The 124 mm specimen, 
a male, has pectoral fins with dorsal surface of distal one-half 
of first 6-7 rays with 2-3 rows of about 6 tubercles each across 
each lepidotrich. No other sexual dimorphism observed. 

Distribution (Fig. 77). — Sumatra, Borneo, Java. Although 
frequently reported from the Mekong basin and other parts of 
Thailand and the Malay Peninsula, there are no confirmed rec- 
ords of this species from mainland Asia. At least some records 
of B. hymenophysa from Indochina and the Mekong basin are 
based on Botia helodes Sauvage. 1876. Specimens reported from 
the Malay Peninsula (Weber and de Beaufort 1916;24; Mohsin 
and Ambak 1983:1 17) should be reexamined to confirm their 
identity. Some Bornean and Sumatran records are based on B. 
reversa new species. 

Botia macracantha (Bleeker, 1852) 

Cohiiii macracanilnis Bleeker. 1 852i/:b03 (type locality "Palembang el in flumine 

Kwanlcn, Sumatrae occidcntahs"). 
Hymenophysa macracanilnis Bleeker, \ibOa:h2. 
Bona macracantha Bleeker, 186.1-64:5 

Diagnosis. — Distinguished from all other Botia by having 8 
barbels instead of 6 and enormously enlarged suborbital spines. 
Branched dorsal-fin rays 8. Total vertebrae 30. \'cllowish brown 
or orange with three broad vertical bars, the first bar passing 
obliquely through the eye and suborbital spine sheath. 

Distribution. — Malay Peninsula (Johore). Sumatra (Telok 
Belong; rivers Pangabuang, Musi, Kwanten; Batang Hari. Lake 



Manindjau), Borneo (Barito, Kahajan, Kapuas. Bangan, Ma- 
hakam). Unknown from Java and northern Borneo. 

Botia reversa new species 

(Figure 76) 

Hymenophysa hymenophysa Fowler, liJlM .SflO (record based entirely or partly on 
/; reversa from Balu Sanghar) 

HoLOTVPE. — MZB 3-521, 87.6 mm, Kapuas basin, rocky channel in mainstream 
olSungai Pmoh, ,37 km S of Nangapinoh (Kapuas 1976-27). 

Paratvpes.-CAS 49349, 4: 85,8-101 mm, collected with the holotype, MZB 
3522 ( I) and USNM 230264 (3). 4: 90.7-103 mm, Kapuas basin, rocky channel 
in mainstream of Sungai Pinoh at Nanga Saian, 45 km S of Nangapinoh (Kapuas 
1976-29); RMNH 7651, 2: 71. 0-75. 1 mm. Mahakam basin. Long Bluu; RMNH 
28042, 97.2 mm, upper Mahakam; CAS-SU 8008, 4: 62.6-75.0 mm, Batu San- 
ghar, Tanah Datar, Padangsche Bovenland (elevation 1 ,5(X)-3,000 feet), Sumatra 
(note: additional specimens from this colleclion should he at ANSP); UMMZ 
155675, 2: 65 5-95.7 mm, Tjisokan. ,Iava. 

Diagnosis. — A Bolia with complex vertical bars apparently 
most closely related to B. hymenophysa from which it differs in 
having a shorter snout, slightly more robust body, 9-1 1 instead 
of 12-13 branched dorsal-fin rays, a slightly deeper caudal pe- 
duncle and shorter caudal fin, and a strikingly diflferent color 
pattern which in at least some respects is the reverse of that in 
B hymenophysa. 

The holotype and 1 7 paratypes of B. reversa may be further 
characterized as follows (condition of paratypes in parentheses); 
head length 3.0 (2.9-3.4); snout 6.2 (5.7-6.7); eye 23.1 (21.8- 
25.8); suborbital spine extending posteriorly to below middle 
of eye; body depth 3.9 (3.8-4.6); body width 8.2 (6.9-9.6, great- 
est in gravid females); depth of caudal peduncle 5.6 (5.3-6.2); 
predorsal length 1.8(1.75-1.85). 

Frequencies of fin rays are as follows: dorsal-fin branched rays 
9(5). 10(12). 11(1); pectoral-fin rays 13(2), 14(10), 15(4), 16(2); 



ROBERTS-WESTERN BORNEO FISHES 



103 



pelvic-fin rays 8(18). A 103 mm Kapuas male has 1-2 rows of 
2-3 tubercles across each lepidotrich on middle portion of pec- 
toral-fin rays 3-5. No other sexual dimorphism observed. 

Body with about 12 complex vertical bars consisting of al- 
ternating pale and dark bands separated by darker thin vertical 
streaks; as dark bands generally somewhat wider than pale ones, 
effect is of 12-13 dark bands with very dark margins on a pale 
background (reverse of pattern seen in B. hymenophysa). 

Fin rays of all fins but especially of dorsal and pectoral fins 
more or less densely covered for their entire length or all but 
their tips by fine melanophores; most specimens with interradial 
membranes of dorsal, pectoral, pelvic, and anal fins largely or 
entirely clear except for fine melanophores on portions of in- 
terradial membrane immediately adjacent to fin rays; dorsal fin 
of some specimens with faint longitudinal stripes broken up into 
spots, faintly visible in middle of interradial membranes and 
somewhat darker where they overlie fin rays; only two speci- 
mens with a vertical black mark near tip of dorsal fin between 
last unbranched and first branched ray; basal half of caudal fin 
with four narrow vertical faint dark stripes (sometimes broken 
up into spots). 

While the number of complex vertical bars is about the same 
in B. revcrsa and B. hymenophysa. in B. rcvcrsa the impression 
gained is of somewhat wider dark bars separated by pale inter- 
spaces while in B. hymenophysa the reverse is true. The two 
species presumably are closely related and it is reasonable to 
hypothesize that one of them has undergone at least a partial 
color pattern reversal in its evolution. A curious feature of the 
reversal is that in B. reversa fine melanophores are concentrated 
in the segmental or lepidotrichial joints of virtually all fin rays, 
whereas in B. hymenophysa the joints are devoid of melano- 
phores or have very few of them. 

Distribution (Fig. 77). — Known only from the Kapuas and 
Mahakam rivers of Borneo (mountain stream tributaries), Java 
and Sumatra. A note with the specimens from Sumatra indi- 
cates they were taken at 1.500-3,000 feet. It apparently does 
not occur sympatrically with the lowland species B. hymeno- 
physa. 

Etymology.— The name reversa (Latin) refers to the "re- 
versed" color pattern of this species compared to that of B. 
hymenophysa. 

Ellopostoma Vaillant, 1902 

Elloposloma Vaillant, 1902:145 (type species .tpciiopnu inegalomycler Vaillant. 
1902. by monotypy). 

Diagnosis. — Distinguished from all other cobitids by its 
oblique, squared-off snout (shaped mainly by enormously ex- 
panded maxillae); mouth highly protrusible; a single pair of well 
developed maxillary barbels; ceratobranchial 5 with about 30 
conical teeth in a single row; suborbital spine absent; dorsal fin 
elongate, with 18-19 rays, its origin far in advance of a vertical 
through pelvic-fin origin; pectoral fins not sexually dimorphic; 
vertebrae 33-34. 

It can now be affirmed that El/opostoma is a member of the 
family Cobilidae. In addition to the single row of ceratobran- 
chial teeth, it has three branchiostegal rays and a Weberian 
apparatus with large lateral capsules of very spongy bone. The 
protrusible jaws, while highly modified, are basically similar to 
those of cyprinoids or cobitoids. Basihyal bone Y-shaped, as in 



many Cobitidae. On the other hand, relationships of Ellopos- 
toma to other cobitids are unclear, and it clearly cannot be 
readily assigned to any of the subfamilies of Cobitidae currently 
recognized. 

The genus was known only from the Kapuas basin, but re- 
cently has been found in the Tapi basin in peninsular Thailand 
(M. Kottelat, pers. comm.). Only one species is described. 

Ellopostoma megalomycter (Vaillant, 1902) 

(Figure 7X) 

.ipenoplus /)ic,?£j/o»irfto- Vaillanl. 1902:3145 (type locality "Kapuas"=Sintang?). 
Ellopostoma inegalomycler Weber and de Beaufort. 1916:237: Roberts. 1973 
(redescription of type specimens). 

Material Examined.- Western Borneo: Kapuas 1976-14. 17: 39.9^8.8 mm 
(BMNH 1 98 1.4. 1 3. 20-26. CAS 49350, MZB 3523. RMNH 28875): Kapuas 1976- 
15. 41.5 mm (MZB 35241: Kapuas 1976-22. 45.7 mm (MZB 3525): Kapuas 1976- 
37, 1 I: 38.5-45.8 mm (CAS 49351. MZB 3526). 

Lepidocephalichthys Bleeker, 1863 

Lcpidocephahchlhys Bleeker. 1863(:35. 38. 42 (type species Cohiiis liasselli (Va- 
lenciennes m Cuvier and Valenciennes. 1846. by original designation). 

Plalacanlhus Day. 1865:296 (type species Plalacanilnis agrensis Day, 1865. by 
monotypy). 

Diagnosis. — Lower lip deeply divided into posteriorly pro- 
jecting lobes, each lobe bearing two or three short barbel-like 
projections. Dorsal-fin rays 7-9, anal-fin rays 6-7, pectoral-fin 
rays 8-9, pelvic-fin rays 7-8. Pectoral fins of mature males highly 
modified. Vertebrae 34-38. 

This genus appears to be very widely distributed. The species, 
perhaps most numerous in India, are poorly known. Two Bor- 
nean species previously placed in .Acantophihalmus have been 
studied by me and are here referred to Lepidocephalichthys for 
the first time; L. lorenizi (Weber and de Beaufort, 1916), known 
only from the Kapuas, and L. sandakamis (Inger and Chin. 
1962), known only from the Kinabatangan basin in northern 
Borneo. L. sandakamis resembles .Acantophthalmus in having 
the dorsal-fin origin very far posterior, about midway between 
verticals through the pelvic- and anal-fin ongins (Inger and Chin 
1962, fig. 54e), and vertebral formula (Table 4). For L. lorenizi 
see below. 

Lepidocephalichthys hasseiti (Valenciennes. 1846) 

Colviis Haiselli Valenciennes in Cuvier and Valenciennes. 1846:74 (type locality 

Java). 
'ICobins harbaluloides Bleeker 1851/:435 (type locality '"Sambas, in lluvus"). 
Lepidocephalichthys Hasseltn Bleeker. 1860a:71. 
''Cohitichthys harhaiuloidcs BkekcT. 1860rf:21 ("Sambas. Poniianak") 
Lepidocephalichthys Hasseliu Bleeker. 1863i.':35. 
'^Misgiinms harbatnloides Bleeker. 186.3-64:13. 

This species, known from Java and Sumatra, has not been 
reported previously from Borneo. Its inclusion here is based on 
published accounts of the holotype and only known specimen 
of Cobitis barhatidoides which is apparently lost. Its generic 
identity is dubious. According to Bleeker the holotype was in 
such poor condition that he was unsure whether it had 8 or 10 
barbels. The published accounts, including a figure presumably 
based on the holotype (Bleeker 1863-64, pi. 103, fig. 1) suggest 
that C. barhatidoides belongs in Lepidocephalichthys. It differs 
from the species known from Borneo in having the dorsal-fin 
origin almost directly above that of the pelvic fin. In this and 



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Figure 78. Eliopostoma incKulomycler Kapuas I'iTb-N, 48 8 mm (CAS 49530). 



Other respects it agrees with L. hasselti. Its status presumably 
will remain unresolved until the holotype is found or more 
specimens are collected from Borneo. 

Alfred ( 1 96 1 :33) reported on a specimen presumed to be the 
holotype oiCobitis barbatuloides. RMNH 4960, supposedly col- 
lected at Sambas by J. Einthoven. I have examined this dried, 
badly damaged and completely discolored alcoholic specimen 
and conclude that it is too small to be the holotype. Much of 
the head is missing and all of the fins are reduced to stumps; 
the mouth and barbels are absent; a small detached piece of the 
head with an eye was found in the bottom of the jar. The stan- 
dard length is probably about 33.0 mm, and at most 33.5 mm. 
Making a generous allowance for the missing portions of its 
head and tail, the total length could not be more than 42.5 mm 
(the caudal fin of a 34. 8 mm L. hasselti is only 8.8 mm long). 
The total length of Bleeker's holotype was 46 mm. If specimens 
identifiable with Cobilis barhaltiloides become available and it 
proves to be a distinct species, it should be redescribed and a 
neotype should be designated. 

Lepidocephalichthys lorentzi (Weber and de Beaufort, 1916) 
new combination 

(Figure 79) 

Acanlhophlhahmis lorentzi Wchcr and de Beaufort. l916:32-.33, fig, 12 (type 
locality "Borneo, upper Kapuas"; "Poetes Sibau" or Putussibau accordmg to 
label in bottle). 

Material Examined —Western Borneo: upper Kapuas at Putussibau. 32,3 mm 
(ZMA 103.259. holotype) and 2: 31 6-32,3 mm (ZMA lib 551. non-types). 



This species apparently is known only from the female ho- 
lotype and two additional specimens (male and female non- 
types) collected together with the holotype but previously un- 
recorded. 

Diagnosis. — A Lepidocephalichthys with dorsal-fin origin 
distinctly posterior to vertical through pelvic-fin origin. Mature 
males with innermost pectoral-fin ray bladelike (enlarged and 
hardened); second ray of pectoral fin unmodified. 

Although the bottle label for the three specimens examined 
indicates they are syntypes, the original description appears to 
be based on a single specimen and ends with the statement 
"Length 40 mm [type in the Zoological Museum of Amster- 
dam]." The two largest specimens, a male and a female, are 
both 32.3 mm SL; the female is about 39.4 mm and the male 
39.7 mm TL. Although the female is only about 39.4 mm TL 
and the male about 39.7 mm TL and thus slightly closer to the 
40 mm indicated for the holotype, the figure accompanying the 
original description agrees closely with the female in the straight 
condition of the body, details of color pattern, and much shorter 
pectoral fin. I therefore conclude that the 32.3 mm SL female 
can be positively identified as the holotype and have indicated 
It as such above. 

The male is similar to the two females in all respects except 
pectoral-fin morphology. All three specimens have one simple 
and seven branched pectoral-fin rays (not 1,8 as stated in the 
original description), but in the male the pectoral fin is sub- 
stantially larger and its rays are modified. The outermost 
branched ray is markedly thicker than the other rays but does 




Figure 79. Li'piiioccphulichlhys lorcnl:i Kapuas. Putussibau. 32.3 mm female (ZMA 103.259, holotype, after Weber and de Beaufort 1916). 



ROBERTS- WESTERN BORNEO FISHES 



105 





Figure 80. Lcpidocephahchthys prislcs- 
holotype). 



Kapuas 1976-47, 30.9 mm female (USNM 230266. paratype); b-c. Kapuas 1976-18, 24.5 mm male (MZB 3527. 



not appear to be otherwise modified, while the innermost two 
pectoral rays are thickened for their basal half and bear the so- 
called "lamina circularis." consisting mainly of a flange-like 
bony projection from the middle of the dorsal surface of next 
to last ray. In all three specimens the dorsal-fin origin lies pos- 
terior to a vertical line through the base of the pelvic fins. 

Lepidocephalichthys pristes new species 

(Figure 80) 

HoLOTYPE. — MZB 3527, 24.5 mm male, small swampy stream about 30 km 
W ofSmtang on road from Sanggau to Sin tang (Kapuas 1976-18). 

Paratypes.-CAS 49352 (3). MZB 3528 (1). 4: 20.4-39.2 mm, Sungai Tekam, 
where it enters right side of Kapuas mainstream about 5-6 km upstream from 
Sunggau (Kapuas 1976-16); MZB 3529, 25.8 mm, small forest streams flowing 
into Kapuas mainstream within 10 km upstream of Sanggau (Kapuas 1976-17); 
MZB 3530 (1), USNM 230265 (1), 2: 27.6-32.7 mm, Danau Piam near Ketungau 



(Kapuas 1976-32); MZB 3531, 37.9 mm. Sungai Senang, 37 km W of Putussibau 
(Kapuas 1976-42); MZB 3532 (1), USNM 230266 (1), 2; 23.6-30.9 mm. small 
forest stream flowing into Kapuas mainstream near Gunung Setunggul (Kapuas 
1976-47); MCZ 56064. 2; 30.1-30.8 mm, Lundu (coll. H. W. Smith, no date or 
additional information). 

Diagnosis, — Dorsal fin usually with 7 branched rays (8 in 
one paratype from Lundu), its origin well in advance of a vertical 
through pelvic-fin origin; pectoral fin of sexually mature males 
with inner margin of enlarged second ray bearing 8-1 1 medially 
directed serrae (spinelike projections of enlarged lepidotrichia), 
caudal fin weakly rounded, truncate, or slightly concave (not 
forked). Total vertebrae 33-34, 
Distribution. — Western Borneo (Lundu, Kapuas), 
Etymology. — The name pristes. a noun in apposition (Greek, 
saw) refers to the serrate pectoral-fin ray observed in males of 
this species. 



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CALIFORNIA ACADEMY OF SCIENCES 




Figure 81- LcpidiKcphalu!, spectrum Kapuas 1976-50, 74 7 mm (MZB 3533, holotype). 



Lepidocephalus Bleeker, 1858-59 

Lepuloccphalus Bleeker, 1858-5''i':303 (type species Cnhiiis macrochir Bleeker, 
1854, b\ suhscqucnt designation of Bleeker, 1863( 35) 

Diagnosis. — Superficially similar to Acantophihalinus but with 
a much deeper body, depth 5-6 in standard length; vertebrae 
42-43. 

Only two species known, the type species with eyes, and a 
closely related undescribed species without eyes discovered dur- 
ing the Kapuas survey of 1976. 

Lepidocephalus macrochir (Bleeker, 1854) 

C'uhilii mucwclur Bleeker, 1854i;:97 (type locality Sumatra orientalis. provinc. 

Palembang, ubi contluunt Hum. Lamatang ct Emm, Surakarta. Javae centralis. 

in llumine Pepeh). 
Lepuloccphalus macrochir W?eV.ei . 1860ij:7U. 

Lcpidocephalichlhys pallcns Vaillant, 1902:154 (type localits Kapuas). 
Lepuloccphalus pallciis Weber and de Beaufort. 1916:28. 
Acanlhophlhalinus pahangensis Ae Beaufort. 1933:31 (type locality Mentakab. 

Pahang Ri\er. Malay Peninsula). See Hora (1941a:55-56)- 

Matekial Examined— Western Borneo: Kapuas. 43 mm (RMNH 7783. ho- 
lot\pe /- pallcn\)- Bleeker collection, localit\ unknown, 76.5 mm (BMNH 
I8(i6,5,2 55, ssnt\pe L niucroclur). 

The only Kapuas specimen of L. macrochir (RMNH 7783, 
holotype of L. pollens) has dorsal-fin rays iiiS, anal ii6, pectoral 
ilO, pelvics 16; about 34 scale rows between dorsal-fin origin 
and lateral line; dorsal portion of head, cheek and opercle cov- 
ered with fine scales. One of the distinctions used by Weber and 
de Beaufort (1916:28) to differentiate L. pattens and L. macro- 
chir. pectoral-fin shape, is related to sexual dimorphism; in males 
the second pectoral-fin ray becomes greatly elongated. Two dark 
spots on the caudal-fin base of the holotype of L. patlens men- 
tioned and figured by Vaillant are present only on one side of 
the body and are probably artifacts (dirf^). 

Distribution. — Malay Peninsula (Pahang). Sumatra (Palem- 
bang). Western Borneo (Kapuas). .lava (Pepeh R. at Surakarta). 

Lepidocephalus spectrum new species 

(Figure 81) 

Holotype. — MZB 3533. 74.7 mm. rocky ledge in middle of Sungai Melawi 
near conHuence with Kapuas mainstream, about 0.5 km upstream from Sintang 
(Kapuas 1976-50), 

Paratypes, — MZB 3534, 53 3 mm. Kapuas mainstream 58 km NE ofSinlang 
,ind I km downstream from Sebruang (Kapuas 197h-45). CAS 49353 (2), MZB 



3535 (1). and USNM 230267 (1), 4: 38,7-59.8 mm. Kapuas mainstream opposite 
Silat (Kapuas 1976-48), 

Diagnosis. — Superficially similar to its only congener, L. 
macrochir, but without eyes. 

Body entirely creamy or pinkish white in life. A small, shallow 
depression above suborbital spine in area normally occupied by 
eye; no sign of superficial or underlying eyeball or visual pig- 
ment. Lateral line complete, straight, with about 90 short pore- 
bearing tubules. Scales minute, embedded in thick skin and not 
readily countable or observable. 

Distribution. — Kapuas. 

Etymology.- The name spcctriim (Latin) refers to the ghast- 
ly or ghostlike character of this species. 

Nemacheilus Bleeker, 1863 

Noemacheilus Kuhl and \an Hasselt in van Hasselt. 1823:133 (unidentifiable). 
Seniacheilus Bleeker. 1863i:34-3() (type species Cohinsfascialus Valenciennes in 

CuMcrand Valenciennes. 1846, by subsequent designation of Jordan, 1917: 

llh) 
Modigliania Perugia, 1893:246 (type species Modigltana paptllosa Perugia, 

[i9'i=?Neniachcihis fasciann. by monotypy). 

In terms of number of species and frequency of citation, this 
is by far the most important of the fish genera first recognized 
by Kuhl and van Hasselt. Its original description has been var- 
iously attributed to Kuhl and van Hasselt (or van Hasselt) and 
to Bleeker, frequently without following the original spelling of 
either author. Bleeker himself has contributed at least two un- 
warranted spelling variants (Nematochcitiis and Nemachilus), 
and others have contributed Nemacheilos and Noemachitiis. 
Thus it is particularly desirable to determine the correct original 
authorship and valid spelling. 

The statement by Weber and de Beaufort (1916:38) that A'^'- 
niachittis [sic] van Hasselt. 1823:133 is "without description" 
is not precisely correct, but the genus could not possibly be 
identified from van Hasselt's account, which may be quoted in 
its entirety: "Noemacheilus Nob. nadert door platte maxillen 
het genus Poecitia Schn.. de Zundanezen noemen hem Jelaer, 
hij lecft bij Buiten/org en de species heeft in onze teekening den 
nam fascialiis Nob," [In other words, Nocmacheiliis Nobis, i.e., 
of Kuhl and van Hasselt (as opposed to Mihi, of van Hasselt 
alone), is distinguished by flat maxillae (or upperjaw parts) like 
those of Poccitia Schneider in Bloch and Schneider, its Sun- 
danese name is Jelaer, and it is found at Buitenzorg (near Jakarta 



ROBERTS-WESTERN BORNEO FISHES 



107 




Figure 82. Nemacheilus tactogcneus. Kapuas 1976-45. 48.1 mm male (MZB 3542, holotype). 



or Bogor); and in our figure of it the species bears the name 
fasciatiis.] But the figure was not pubHshed. Only one taxonomic 
character is mentioned, i.e.. the shape of the maxillae, and with- 
out a figure the description is too subjective to be useful. Kuhl 
and van Hasselt's figures and specimens of Javanese loaches 
were later studied by Valenciennes, and consequently the iden- 
tity of N. fasciatus and the others became known. In the case 
o( N. fasciatiis. Valenciennes published an excellent description 
and fine illustration of one of the specimens but chose not to 
recognize Noemacheihis. placing the species instead in Cobilis. 
It remained for Bleeker (1863) to recognize the genus. Sleeker 
(1863:37) designated Noemacheihis fasciatus van Hasselt, 1823 
as type species but it is unidentifiable. The earliest valid type 
species designation appears to be by Jordan (1917). 

Six species of Nemacheilus occur in western Borneo. Three 
of these were discovered during the Kapuas survey of 1976; one 
has been described already (Kottelat 1984). and the other two 
are described herein. 

Nemacheihis. like all Nemacheilinae, lacks a subocular spine, 
but in many species sexually mature males develop a large, 
posteriorly directed subocular papilla immediately below the 
antero ventral margin of the eye. The papilla was discovered by 
Perugia (1893) who, not recognizing it as sexually dimorphic, 
regarded it as diagnostic of his new genus and species Modigli- 
ania papillosa (type locality Lake Toba, Sumatra). Weber and 
de Beaufort (1916:40-41), reported the absence of the papilla 
in type specimens of A/, papillosa they examined and referred 
the species to A', fasciatus. I have examined a series of N. fas- 
ciatus from Buitenzorg. the type locality (CAS-SU 20498, 20: 
23.0-49.3 mm) and confirmed that males do have a subocular 
papilla. Other species of the genus in which males with a 
subocular papilla have been examined by me include N. beavani. 
N. kapuasensis, N. saravacensis. N. longipectoralis. and N. se- 
langoriciis. 

Nemacheilus kapuasensis Kottelat, 1984 

Nemacheilus kapuasensis Kottelat, 1984:244 (type locality Kapuas). 

Material Examined. — Western Borneo; Kapuas 1976-24, 4: 51.5-54.9 mm 
(BMNH 1982.3.29.139-141, MZB 3556); Kapuas 1976-25, 31: 38.7-56.3 mm 
(FMNH 94235, IRSNB 19745, KUMF 2855, MZB 3557, MNHN 1982-700); 
Kapuas 1976-26, 3: 32.3-55.9 mm (MZB 3558. RMNH 28880); Kapuas 1976- 
27, 23: 32.3-59.1 mm (MZB 3559, UMMZ 209872, USNM 230271); Kapuas 



1976-29, 42: 41.7-55.6 mm (MZB 3560. 4004-5. CAS 47378. 49360 [including 
holotype and 3 1 paratypes of A', kapuasensis]). 

Distribution. — Borneo (Kapuas. Rejang). 
Nemacheilus lactogeneus new species 

(Figure 82) 

HoLort'PE. — MZB 3542, 48 1 mm, Kapuas mainstream 58 km NE of Sintang 
and 1 km downstream from Sebruang, 16 August 1976 (Kapuas 1976-45). 
Paratype.— CAS 49355. 44.6 mm. collected with holotype. 

Diagnosis. — A morphologically generalized Nemacheilus 
similar to A^. fasciatus in body proportions and in most other 
respects but slightly more elongate (with 38 vertebrae instead 
of 33-36 as in other Sundaic Nemacheilus): color in life milky- 
white. without vertical marks or bars on fins or body in live or 
preserved specimens (vertical marks or bars characteristic of all 
other Sundaic nemacheilins); upper half to two-thirds of body 
nearly uniformly covered with very fine, faint melanophores. 
lower part of body without melanophores; barbels and fins with- 
out melanophores or with very fine faint melanophores along 
inner surface of outer rostral barbel and on elongate anterior- 
most dorsal-fin rays; caudal-fin base with a dark, subdermal, 
midlateral oval spot. 

Holotype and paratype (immature males?) with small, fleshy 
fingerlike subocular flaps and weakly developed tubercles on 
posterior margin of peduncular scales; head 4.3-4.4; snout 1 1.7- 
12.8; eye 18.9-19.8; interorbital width 15,4-17.2; barbels elon- 
gate; inner rostral barbel 11.5-12,9. outer rostral barbel 7.1- 
7.4, extending posteriorly beyond hind border of eye and slightly 
farther than maxillary barbel; maxillary barbel 10.9-12.4; body 
depth 6.4-6.8, width 8.0-8.4; caudal peduncle depth 11.2-11 .7, 
length 5.8-6.5. Dorsal fin with 9-1/2 branched rays; length of 
first branched dorsal-fin ray 1 1.2-1 1.7; pectoral fin length 4.3- 
4.7; pectoral-fin rays i 1 2-1 3. unbranched outermost ray with 
short filamentous extension nearly reaching pelvic-fin origin, 
branched rays with shorter extensions; pelvic fin length 6.9; 
pelvic-fin rays i7; caudal fin moderately forked, lobes about 
equal. Scales very small, difficult to count, about 160 in lateral 
series, 50 predorsal, 50 circumpeduncular. 

Distribution. — Known only from the Kapuas type speci- 
mens. 

Etymology.— The name lactogeneus (Latin) refers to the milk- 
white coloration of this species observed in life. 



108 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 83. Nemachciliis mamlucps. Kapuas 1976-27. 78.9 mm (MZB 3?43, holotype). 



Nemacheilus cf. longipectoralis Popta. 1905 

Socmachcilus longipixtorali^ Popta, I9IIS l!\2 (t\pc locality upper Mahakam). 

Material Examined. — Western Borneo; Kapuas I97b--I5. 1: 37.6 mm (MZB 
4007). 



Nemacheilus maculiceps new species 

(Figure S3) 

Holotype. — MZB 3543. 78 9 mm. immature male, rocky channel in main- 
stream ofSungai Pinoh. 37 km S of Nangapmoh. 24 ,luly 1976 (Kapuas 1976- 
27) 

Paratvpes.-BMNH 1982.3 29 135. CAS 49356. R.MNH 28878. L'SNM 
230269. 7: 48.5-99.7 mm. collected with holotype; MZB 3544. 1 16 mm, rocky 
channel in mainstream of Sungai Pinoh at Nanga Saian. 45 km S of Nangapinoh. 
26 July 1976 (Kapuas 1976-29); CAS 49357. MZB 3545. 4: 25. 1-29.9 mm. rocky 
ledge in middle ofSungai Melawi near confluence with Kapuas mainstream, about 
(15 km upstream from Smtang. 18 .August 1976 (Kapuas 1976-50). 

Diagnosis. — A large, heavy-bodied Nemacheilus with small 
eyes and very fine scales; largest specimen, a gravid female, 1 16 
mm; head elongate, somewhat dorsoventrally depressed (head 
laterally compressed in most Sundaic nemacheilines); larger 
specimens with cheeks projecting laterally well beyond eyes; 
entire dorsal, lateral, and sometimes ventral surface of head 
(including lips, rostral barbels, gill cover, and sometimes bran- 
chiostcgal membranes) with numerous moderately large dark 
maculae or vermiculae; dorsal-fin rays iv8-l/2 (dorsal-fin 
branched rays almost invariably 9-1/2 in other Sundaic Ne- 
macheilus): dorsal-fin origin relatively posterior, only slightly 
nearer to snout-tip than to caudal fin base (dorsal-fin origin 
usually much closer to snout-tip in other Nemacheilus): pectoral 
and pelvic fins widely separated (pectoral fin extending poste- 
riorly to or beyond pelvic-fin origin in all other Bomean Ne- 
macheilus): males without digitiform subocular flap (present in 
all other Sundaic Nemacheilus'?). Scales in lateral series about 
215, in transverse series 50, circumpeduncular 70. Dorsal sur- 
face of head relatively flat and evenly sloped anteroposteriorly; 
head length 3.5-3.9; eye hon/ontally oval, diameter 29-38 (strong 
negati\e allometry); interorbital width 14.5-17.2; barbels mod- 
erately elongate; rostral barbels with bases approximate; inner 
rostral barbel length 18-25. outer 13.9-16.8; maxillary barbel 
extending posteriorly to beyond hind margin of eye. its length 
13.0-16.8; nasal valve with a short barbel-like process dorsally; 
posterior nostril narrowing to a slit posteriorly (posterior nostril 
round or slightly oval, usually not narrowing to a slit posteriorly 
in other Nemacheilus examined); conformation of lips and 



horny jaw sheaths as in generalized Nemacheilus including N. 
fasciatus: upper lip relatively smooth, without papillae or marked 
transverse furrows; lower lip interrupted medially; homy jaw 
sheaths moderately developed, lower without processus denti- 
formis; body stout, subcylindrical. width at pectoral-fin origin 
5.6-7.5. depth at dorsal-fin ongin 5.6-6.7; height of first branched 
dorsal-fin ray 5.8-7. 1 ; pectoral and pelvic fins widely separated, 
peh ic-fin origin distinctly posterior to vertical through dorsal- 
fin origin; posterior margin of pectoral fin rounded or lobate, 
without fringes due to freely projecting ray tips seen in many 
Nemacheilus): pectoral-fin length 4.8-6.0. pelvic 6.0-7.5; pec- 
toral-fin rays i 10-12. pelvic i7. Pectoral and pelvic fins with 
fleshy axillary flaps arising above fin origin; caudal peduncle 
compressed, short, and very deep, depth 7.5-9.0. without dorsal 
or ventral keel; caudal fin moderately forked, lobes rounded, 
lower somewhat larger than upper; principal caudal-fin rays 
10-f9; vertebrae 20-22+1 1-13 = 33-34. 

Larger specimens with about nine evenly spaced, dark, wavy 
vertical bars or blotches on dorsal and lateral surface of body 
and two similar vertical bars on caudal fin; in some specimens 
bars vertically complete, in others broken into separate portions 
on dorsal and ventral half of body (especially posteriorly). Ab- 
domen pale or whitish (covered with fine scales which may be 
embedded in skin or secretion-covered and difficult to detect). 
Dorsal, anal, pectoral and caudal fins with fin rays generally 
dusky except near tips which are plain; interradial membranes 
nearly plain; anterior margin of dorsal fin dusky; no dark spot 
on base of dorsal fin near origin. Smaller specimens may have 
a broad, dark midlateral longitudinal stripe on body; this is 
absent in all specimens abo\e 70 mm. Largest males, apparently 
immature. 23.0-78.9 mm. with a single row of small breeding 
tubercles on dorsal surface of outermost three pectoral-fin rays. 
No other secondary sexual dimorphism observed. 

Distribution. — Known only from the Kapuas. 

Et'imology.— The name maculiceps (Latin) refers to the 
spotted head of this species. 

Nemacheilus saravacensis Boulenger. 1894 

.S'cinuclicilin saravaicnsn Boulenger. 1S94;25I (type locality Senah. Sarawak). 
Material E.xamined— Kapuas 1976-8. 36 11 mm (MZB 3561). 

The identification of the single specimen of this species ob- 
tained during the Kapuas survey of 1976 was verified by M. 
Kottelat. It IS male, with a subocular papilla, single rows of 



ROBERTS-WESTERN BORNEO FISHES 



109 




Figure 84. \aillamclla. Above, 1. cucpiplcra. Kapuas 1976-37, 83.0 mm (CAS 49362); below, I . inaaisi. Kapuas 1976-29, 97.7 mm (MZB 3570). 



small breeding tubercles on the dorsal surface of the pectoral- 
fin rays, and weak tuberculation near middle of caudal peduncle. 
Distribution. — Sarawak, Mempawah. 

Nemacheilus selangoricus Duncker, 1904 

Nemacheilussetangoncus Duncker, 1 904: 1 75 (lypc locality "Umgcgend von Kuala 
Lumpur"). 

Material Examined. — Malay Penmsula; Perak, Plus R., Lasal, 40.6 mm (C.AS- 
SU 31080); Johore, Kota Tinggi, 8; 27.5-38.3 mm (CAS-SU 39396); Johore 
Simpang Rengam, 1 1 : 42.8-55.4 mm (CAS-SU 39397); Singapore, Mandai Road 
4: 37.8-54.7 mm (CAS-SU 32598); Smgaporc, 20; 25.8-50.5 mm (CAS-SU 31081) 
Northeast Borneo: Balung R., 10: 18.2-39.7 mm (CAS-SU 33609). Western Bor- 
neo: Kapuas 1976-7, 20.2 mm (AMNH 48937, MZB 3547); Kapuas 1976-10, 4 
17.3-23.0 mm (BMNH 1982.3.29.136-138, MZB 3548); Kapuas 1976-12, 30,7 
mm (MZB 3549); Kapuas 1976-25, 4: 37.9-45.8 mm (CAS 49358, MZB 3550) 
Kapuas 1976-37, 1 1: 25.0-31.4 mm (FMNH 94234, MZB 355 I ); Kapuas 1976- 
39, 68: 15,5-50,4 mm (CAS 49359, HZM 6476, KUMF 2854, MCZ 58351 
MNHN 1982-699, MZB 3552, RMNH 28879, ROM 38602); Kapuas 1976-42 
15: 22.5-37.8 mm (MZB 3553, USNM 230270); Kapuas 1976-47, 8: 25.0-37.6 
mm (MZB 3554, UMMZ 209922); Kapuas 1976-51, 4; 21.0-38.8 mm (MZB 
3555, ZMA 116.737). 

Diagnosis (modified from Hora 1941). — Body with about 1 2- 
14 sharply demarcated light vertical bands set off by dark pig- 
ment. Dorsal fin with a very dark round spot near its origin and 
three irregular rows of lighter spots. A dark suborbital spot. 
Barbels very long, almost reaching gill openings. 

This is perhaps the commonest species of Nemacheilus in the 
Malay Peninsula. Herre ( 1 940:34) previously reported the spec- 
imens from the Balung River as A^. selangoricus but this iden- 
tification was questioned by Hora (1941:59). Having compared 
these specimens with those from the Kapuas and the Malay 
Peninsula I find them closely similar and consider that they are 
all A', selangoricus. 

Distribution. — Malay Peninsula. Northern Borneo (Ka- 
puas). 

Vaillantella Fowler, 1905 

Vaillantella Fowler, 1905:474 (type species Sonachcilus cuepiptenis Vaillant, 
1902, by original designation and monotypy). 

Diagnosis. — Distinguished from all other loaches by its ex- 
traordinarily elongate dorsal fin, with 59-73 rays, and by the 



deeply forked caudal fin with its greatly elongated upper lobe. 
Principal caudal-fin rays 10 + 9. Vertebrae 57-60. Secondary 
sexual dimorphism unknown. For a detailed characterization 
see Nalbant and Banarescu (1977, especially pp. 100-101). 

The genus comprises only two species; both occur in the Ka- 
puas basin, but in different habitats: V. euepiptera in lowland 
streams and I '. maassi in highland streams. 

Vaillantella euepiptera (Vaillant, 1902) 

(Figure 84 above) 

Nemacheilus etiepipienis Vaillant. 1902:137 (type locality ■■Ponlianak, Kapoeas 

(Sintang)""). 
Wnllaniella ciiepipienn Weber and dc Beaufort, 1916:37. 
I'aillaiilella euepiptera Nalbant and Banarescu. 1977:101. 

Material Examined. — Western Borneo; Kapuas 1976-16, 22: 30.2-81.0 mm 
(AMNH 48938. BMNH 1892.3.29.142-144. CAS 49361, MZB 3562); Kapuas 
1 976- 1 7, 4: 36, .3-73.0 mm (FMNH 94236. MZB 3563); Kapuas 1976-2 1 , 2: 44.7- 
70.5 mm (IRSNB 19746, MZB 3564); Kapuas 1976-37. 25: 48,8-120 mm (CAS 
49362, MNHN 1 982-70 1 , MZB 3565. RMNH 2888 1 ); Kapuas 1976-39. 35: 36.7- 
82.8 mm (MZB 3566, UMMZ 209904, USNM 230272); Kapuas 1976-42, 5: 
56.9-79.7 mm (MZB 3567. MZB 116.538); Kapuas 1976-46, 55.1 mm (MZB 
3568); Kapuas 1976-47, 5: 50.9-80.2 mm (CAS 49363. MZB 3569). 

Diagnosis. — Differs from its only congener in having anal fin 
with only 7 rays (vs. 12-15 in I', maassi), and in its coloration. 
Distribution. — Kapuas. 

Vaillantella maassi Weber and de Beaufort, 1916 

(Figure 84 below) 

Vjillanlella maassi Weber and de Beaufort. 1916:38. fig. 18 (type locality Goen- 

oeng Sahilan on Kampar km River. Sumatra). 
l'aillanlellallamfasciataTv.'eedK. 1956:59, pi, 6b(type locality River Tahan, near 

Kuala Tahan, Pahang, Malay Peninsula). 

Material Examined —Malay Peninsula: River Tahan near Kuala Tahan, Pa- 
hang, 2; 97,2-125 mm (CAS-SU 50180, paratypes of r flarofasaala). Western 
Borneo: Kapuas 1976-29, 2: 97.7-129 mm (CAS 49364, MZB 3570), Kapuas 
1976-45, 6: 44.6-77.3 mm (BMNH 1982.3.29.145, MZB 3571, UMMZ 209917, 
USNM 230273); Kapuas 1976-50, 3: 67,2-76,2 mm (CAS 49365, MZB 3572, 
ZMA 116.539), 

Diagnosis. — See under V. euepiptera. 
Until now this species has been known only from the holo- 
types of I '. Diaassi. the type series of I '. flavofasciata. and a 



110 



CALIFORNIA ACADEMY OF SCIENCES 



single specimen from the Kapuas River at Sintang identified by 
Naibant and Banarescu (1977:101). These authors did not ex- 
amine the types of I '. Ilavofasciata and maintained it as a valid 
species. I have compared one of the paratypes with my Kapuas 
specimens and consider them conspecific. 

Distribution. — Sumatra (Kampar kiri River). Malay Pen- 
insula (Pahang). Borneo (Kapuas). 



Ariidae 

The "sea catfishes" or Ariidae, all or almost all species of 
which produce relatively small numbers of large eggs that are 
carried through hatching in the male's mouth, include a number 
of species endemic to fresh water. In large rivers of western 
Borneo occur three distinctive freshwater species which appar- 
ently are not closely related to each other, and their relationships 
to marine ariids or to freshwater ariids of other areas (such as 
New Guinea and Australia) is obscure. These are Alius mela- 
nochir. A. sloniiii. and Hcnupinuiodus honwcnsis. 

Specimens of three additional ariid species have been re- 
corded from Pontianak: Ariiis argyropleuron Valenciennes ;/; 
Cuvier and Valenciennes, 1840, A. oetik (Bleeker, 1846), and 
Batrachoccphalus nnno (Hamilton-Buchanan, 1 822). These are 
primarily marine or estuarine; their occurrence in fresh water 
is unconfirmed and likely to be limited to areas of tidal influence 
and the lowermost courses of large rivers near the sea or estu- 
aries. 



Arius Valenciennes, 1840 

Anus Valenciennes in Cuvier and Valenciennes. 1840:.S.^ (type species Pimclodus 
aiiiii Hamilton-Buchanan, 1 822. by absolute tautonym> ; sec Wheeler and Bad- 
dokwaya 1981:76'*), 

Cc'pha/ocassis Bleeker, 1858t:98 (type species C'cpluilniawn mclanmhir Bleeker. 
1858, by subsequent designation of Bleeker. 186.^</:9I) 

Hcxanemalichlhys Bleeker. 1858c: 126 (type species Biinius Mimlaiciis Valen- 
ciennes III Cuvier and Valenciennes, 1839. by monotypy). 

Hciiuanus Bleeker. 1863tj:90 (type species Ccphuloccissis slunuii Bleeker, 1858, 
h> monolypy). 

The synonymy presented here includes only names relevant 
to systematics of freshwater ariids found in western Borneo. 

Anus has always been a catch-all for large numbers of catfishes 
with poorly known relationships. As originally conceived by 
Valenciennes it was nearly Linnaean in scope and included species 
now assigned to Auchenipteridae, Bagridae, and Diplomystidae 
as well as Ariidae. 

The name Tachysurus Lacepede, 1803 (type species Tachy- 
surus sinensis Lacepede, 1 803) continues to be regarded by some 
authors as a valid senior synonym oC Anus Valenciennes. This 
nomenclalural problem is discussed in detail by Wheeler and 
Baddokwaya (1981). Their findings and conclusions, with which 
I agree, may be briefly summarized: I) the original painting 
upon which Tachysurus sinensis was based apparently is no 
longer available; 2) the engraving of the painting and original 
description by Lacepdde (1803) are obviously based on some 
kind of catfish but cannot be identified with specimens of any 
known species or genus or even with the family Ariidae as 
presently understood (sensu Regan 191 1); 3) Tachysurus sinen- 
sis Lacepede. 1803, appears to be an unrecognizable siluriform 
fish from fresh water in China. Although Wheeler and Baddok- 



waya do not employ the term, Tachysurus sinensis (and 
Tachysurus) is a nomen dubium. 

As currently understood Arius contains a large number of 
species from Central and South America, Africa, Asia, and the 
Australian region; presumably several genera are involved. Some 
of these already have received names, but anatomical or other 
information for diagnosing these genera and deciding which 
species should be assigned to them simply does not exist. 

The type species Arius arius is not well known; it was long 
regarded as a junior synonym of Arius gagora (Hamilton-Bu- 
chanan, 1 822) but is recognized as a distinct species in Jayaram 
(1982) and other recent works on Indian Ariidae. 

Arius melanochir Bleeker, 1852 

(Figure 85 below) 

Anus inelaiwclurBkckcT. 1852</:590 (type locality Palembang. m Iluviis). 
Ccpiialncassis melanochir Bleeker. 1858i:103. 
Anus melanochir Giinlher. 1 864: 161. 

Material Examined. — Western Borneo: Kapuas 1976-19. 11: 125-250 mm 
(CAS 49426. MZB 37 1 3, USNM 2303 1 1 ); Kapuas 1 976-40. 222 mm ( MZB 3714); 
Kapuas 1976-48. 3: 60.6-91.6 mm (BMNH 1982.3.29.193. MZB 3715. RMNH 
289(18) 

Distribution. — Sumatra (Palembang, Djambi). Borneo (Ka- 
puas). 

Arius stormii Bleeker, 1858 

(Figure 85 above) 

Ccphalocassis Slorniii Bleeker. 1858i:246 (type locality Sumatra. Palembang. in 

flumine Mussi). 
Hcimanus Storim Bleeker. 1862-63:29, 
,I//i/,s v((7n?;;/ Giinlher. 1864:162, 

Material Examined -Western Borneo: Kapuas 1976-19, 3: 282-105 mm (CAS 
49427, MZB 3716, USNM 230312) 

A moderately large ariid, attaining 500 mm, characterized by 
very large, exceptionally stout, and heavily serrate dorsal and 
pectoral-fin spines; a pointed, strongly projecting snout; large, 
sharp, backwardly projecting jaw teeth; premaxillary tooth band 
partially exposed when jaws closed; palatal teeth in two small 
round vomerine patches (vomerine and palatine patches on each 
side sometimes nearly or quite continuous; see Weber and de 
Beaufort 1913, fig. 1 17); maxillary and mental barbels moder- 
ately elongate; lateral line on head and body forming dense 
network of anastomosing canals. 

It should be noted that Bleeker (1863) proposed the mono- 
typic genus Heiiiiarius based on this species. Only one other 
species. Anus {f/einianus) danielsi Regan, 1908, has been re- 
ferred to Hemiarius. and this species is now placed in Cochle- 
felis. an unrelated carcinophagous genus endemic to large rivers 
in southern New Guinea and northern Australia (Roberts 1978: 
45). The species Anus cf stirhngi Ogilby, 1898 reported from 
the Fly River (op. cit. pp. 37-38) bears some resemblance to A. 
slonnii. but they probably are not closely related. 

Distribution. -Thailand (Chao Phrya). Sumatra (Palem- 
bang, Djambi). Borneo (Kapuas, Bandjermasin). Apparently 
confined to fresh water (large rivers). 

Hemipimelodus Bleeker, 1858 

/Icnupiiiicloihn Bleeker. I858r:236 (type species Pimclodus honwcnsis Bleeker, 
1851. b\ subsequent designation of Bleeker. I863ij:92). 



ROBERTS-WESTERN BORNEO FISHES 



III 




Figure 85. Anus. Above. A. slomu. Kapuas 1976-19. 405 mm (CAS 49427); below. A. melanochir. Kapuas 1976-48, 83.0 mm (MZB 37151. 



Hemipimelodus borneensis (Bleeker, 1851) 

Piinelodm horneensn Bleeker, I85l(:430 (type locality "Sambas. In fluviis"). 
Hemipimelodus borneensis Bleeker, I858c':238. 

Material Examined. — None. 

Distribution.— Thailand (Chao Phrya, Bangpakong). Su- 
matra (Palembang). Borneo (Baram, Sambas, Barito). 

Bagridae 

The Old World family Bagridae, with about 27 genera and 
205 species (Nelson 1984), includes some of the most gener- 
alized and some of the most specialized catfishes in western 
Borneo, such as the remarkable Bagrichthys hypseloptems. The 
generalized species, with numerous relatives in tropical Africa, 
India, China, and elsewhere in southeast Asia, belong to the 
subfamily Bagrinae. The main genus in southeast Asia is Mystm. 
with four or five species in western Borneo, including one new 
species obtained during the Kapuas survey of 1976. The most 
specialized bagrids in western Borneo, with related forms re- 
stricted to southeast Asia and China, belong to the Bagrichthyi- 
nae. A few bagrichthyins are plain or uniformly dull colored, 
but most of the genera and species are characterized by a highly 
distinctive pattern consisting of a series of offset, horizontally 
elongate light blotches on the upper and lower halves of the 
body usually separated by a pale midlateral longitudinal streak 
or stripe of variable width. Such coloration is shared by most 
species of Bagrichthys. Bagroides. Leiocassis. and Peheobagrus. 



At least nine or 10 species of Bagrichthyinae occur in western 
Borneo, including one Bagrichthys and one Leiocassis discov- 
ered in 1976 and described herein as new. 

The small, very rare and poorly known species currently iden- 
tified as Peheobagrus ornatus is morphologically distinctive and 
has an extraordinary color pattern entirely different from that 
of any other Pcllcobagrus or Bagrichthyinae; its generic, subfa- 
milial, and even familial placement are tentative. 

Bagrichthys Bleeker, 1858 

Bagrichthys Bleeker, 1 858i': 1 30 (t\pc species Bagrus hypseloplerus Bleeker. 1 852, 

by monotypy). 
Pwiidobagrichlhys Bleeker, 1862-63:9, 49 (type species Bagroides macropterus 

Bleeker, 1853, by original designation). 

Bagrichthys is distinguished from all other genera of Bagridae 
by its elongate and laterally compressed caudal peduncle, with 
13-20 peduncular vertebrae, and from all others except Bag- 
roides in having upwardly (rather than downwardly) directed 
serrae on posterior margin of dorsal-fin spine. In addition, all 
species except B. macracanthits differ from all other bagrids in 
having one or both mental barbels with uniquely convoluted 
and papillose margins (not at all like modifications of mental 
barbels in mochokid catfishes to which they have been com- 
pared by some authors). Anal-fin rays 14-16; pectoral rays 8- 
11; pelvic rays 6. Gill rakers 2-4-^3-11=5-15. Vertebrae 17- 
20-^24-30=42-49. 

Bagrichthys occur mainly in large muddy rivers such as the 



112 



CALIFORNIA ACADEMY OF SCIENCES 



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FrciiiRE 86. Iiai;rniilh\'s inucracanllius. oral dcnlUion Mahakam, i^J.? mm 
(RMNH 7834), holot\pc B raillann) 

mainstream of the Kapuas and its larger tributaries, and feed 
almost exclusively on detritus of higher plants. The entire length 
or long portions of the intestine is filled with vegetable detritus 
in nearly every one of the four B. hypscloplcrus. one B. macra- 
canthiis, 25 B. niacivpleriis. eight B. niicranodus, and four 5. 
sp. undet. (Mekong basin) in which I examined the gut contents. 
The detritus is generally loose, sometimes with silt particles but 
never clay. The stomach is generally empty but when full its 
contents arc similar to those of the intestines. The only other 
food item observed was a single large winged insect in a 95.8 
mm B. micraiiodiis from Kapuas 1976-45. The intestines are 
convoluted but only moderately elongate for a detritivore. A 
233 mm B. hypsclopterus has intestines 330 mm long and a 105 
mm B. niacropicnis intestines 190 mm long. 

Species of Bagnduhys have been the source of much taxo- 
noniic confusion. Problems have arisen from 1) assumption of 
extensive late allometric changes in 5. hypsclopicrus: 2) Bleek- 
er's figure of oral dentition of 5. niacroptcrus in which the num- 
ber of teeth is greatly exaggerated; 3) poorly understood ditTer- 
ences in color pattern reported or observed in Bagrichthys sp. 
or spp. 

The first problem has been resolved by the present finding 
that the unique morphology of 5, hypsclopicrus is fully devel- 
oped in individuals under 200 mm, that specimens of all sizes 
can be identified by 19-20 rather than 13-16 peduncular ver- 
tebrae (vertebrae entirely posterior to base of last anal-fin ray 
including hypural centrum as 1 ), and that B inacracanihiis has 
been misidcntified as young B. hypsclopterus. The second prob- 
lem has been resolved bv examination of the dentition in the 



holotype and other specimens of B. macroplerus. The third 
problem, not fully resolved, is narrowed down to color variation 
within B. macracanthiis or to differences in coloration among 
species closely related to B. macracanthiis. Geographical dis- 
tribution of species o( Bagrichthys is shown in Figure 88. 

Key to Bagrichthys of Borneo 

la Mouth opening relatively large and broad; oral dentition 
well developed, jaws and palate with numerous exposed 
teeth forming well defined tooth bands (Fig. 86); dorsal- 
fin spine moderately to extremely elongate, with IS or 
more serrae in adults 2 

lb Mouth opening relatively small and narrow; oral denti- 
tion extremely reduced, jaws and palate with a few scat- 
tered teeth (sometimes apparently absent) deeply buried 
in soft tissue; dorsal-fin spine relatively short, with 15 or 
fewer serrae 3 

2a Body form relatively generalized, nape and dorsal-fin base 
not greatly elevated; jaw teeth not notably heterodont, 
those of lower jaw only partially exposed from gum; gill 
rakers 10-14; dorsal-fin spine moderately long, with 18- 
29 serrae in adults; pectoral-fin rays 9; inner and outer 
mental barbels with straight (non-convoluted) margins; 
total vertebrae 42-43; caudal peduncle moderately elon- 
gate, peduncular vertebrae 14-15; upper and lower cau- 
dal-fin lobes pointed but not markedly filamentous 

B. macracanthus 

2b Body form highly modified, nape and dorsal-fin base ex- 
traordinarily elevated; jaw teeth markedly heterodont, 
teeth of lower jaw about 1.5 times as long and much more 
slender than those of upper jaw, and totally exposed from 
gum; gill rakers 14-15; dorsal-fin spine extremely long, 
extending to or beyond base of caudal fin when depressed, 
with 60 or more serrae in large juveniles and adults; pec- 
toral-fin rays 10-11; inner and outer mental barbels with 
extremely convoluted anterior margins; total vertebrae 
47-49; caudal peduncle extremely elongate, peduncular 
vertebrae 19-20; upper and lower caudal-fin lobes with 
long filamentous extensions B. hypsclopterus 

3a Inner and outer mental barbels strongly crenulated; at- 
tains at least 236 mm; males sexually mature at 2 1 5 mm; 
color in life pale brownish or tan with whitish or cream- 
colored light areas B niacroptcrus 

3b Inner mental barbel crenulated but outer mental barbel 
simple; largest known specimen 125 mm; males sexually 
mature at 95.8 mm; color in life dark brown or brownish 
black with whitish or cream-colored light areas 

_.._ B. niicranodus 

Bagrichthys hypselopterus (Bleeker, 1852) 

(Figure 87) 

BDgnis hypscloplcnis Bleeker. 1852</:588 (type localits Palemhang. in fluviis). 
Bagnchllns liypscloplcnis Bleeker. 1 S.SSf: 131 

Material Examined. — Sumatra'': Palembang', 253 mm. Bleeker (RMNH 6877, 
possible holotype); 172 mm. Bleeker (BMNH 1863.12.4.61). Western Borneo: 
Kapuas 1976-11. 7: 200-241 mm (CAS 49366. MZB 3573. USNM 230274); 
Kapuas 1976-44. 2: 184-228 mm (CAS 49367. MZB 3574). 

D\.\GNosis. — Bagrichthys hypsclopterus is characterized by the 
most highly modified dorsal fin and body form of any Bag- 



ROBERTS- WESTERN BORNEO FISHES 



113 




Figure 87. Bagnchihys hypseloplerus. Kapuas 1976-19, 234 mm male (CAS 49366). 



richthys and perhaps of any member of the family Bagridae. 
Anterior portion of body very compressed and deep, predorsal 
profile extremely steep. Dorsal fin extraordinarily elongate; dor- 
sal-fin spine extending to or slightly beyond level of caudal-fin 
base when depressed, its posterior margin with up to 60 serrac. 
Caudal peduncle excessively elongate, about 1.5-2 times longer 
than in any other Bagrichthys. Gill rakers 14-15, Peduncular 
vertebrae 19-20, total vertebrae 47-49. 

The 184 mm Kapuas specimen has the following: head length 
6,0; eye 6,9 (much smaller than in specimens of any other species 
of Bagrichthys of comparable size); snout 19,2; mouth opening 
23; gill rakers 4+11 = 15; dorsal-fin spine, broken near tip but 
still extending posteriorly when adpressed to beyond caudal-fin 
origin, its length 1 .5; dorsal-fin spine serrae 53 (not allowing for 
gaps between serrae, apparently due to serrae having broken off, 
which would bring the count to about 60); pectoral-fin spines 
broken off near base; humeral process 13.3; body depth 3.3, 
width 7,3; caudal peduncle length 2,6, depth 17,7; nasal barbel 
extending posteriorly well beyond eye, length 14.8; maxillary 
barbel extending posteriorly to somewhat in front of pectoral- 
fin spine, length 7.8; both mental barbels with excessively con- 
voluted anterior margins; outer mental barbel length 16.7, inner 
25.2; upper and lower lobe of caudal fin with long filamentous 
extensions. The other specimens from the Kapuas are very sim- 
ilar in all respects to this specimen except for their larger size. 
Adult or subadult specimens of smaller species of Bagrichthys 
with less modified dorsal fin and body shape have been mistaken 
for young B. hypseloptenis (Weber and de Beaufort 1913:346, 
fig, 146; Jayaram 1968:381, fig. 13; Mohsin and Ambak 1983: 
125, fig. 92). The smallest known B. hypseloptenis. 172 mm 
(BMNH 1863.12.4.61), is as extreme in its morphology as the 
largest known specimens. 



Through the kindness of Han Nijssen I have been able to 
examine material of the Bagrichthys which Weber and de Beau- 
fort (1916:346, fig, 146) mistakenly identified as young B. hyp- 
seloptenis. These specimens, ZMA 101.457, a sexually mature 
male of 108 mm and two gravid females, 182-193 mm, from 
Balang Hari at Djambi, Sumatra. 1 tentatively identify as B. 
inacracaiuhiis. They have the fins black or ver>' dark, and the 
body entirely or almost entirely dark brown or brownish black 
except for a pale or white axial streak; jaws wide, teeth in both 
jaws large and numerous (not heterodont); mental barbels sim- 
ple (not convoluted or papillose); gill rakers 4 + 9-10=13(1), 



90° 



100° 



110° 



120° 



130° 




-20° 



-10° 



-0° 



O hypseloplerus 

• macracanthus 

* macropterus 
m micranodus 



^^=^=3?S,, 



ct:S3 tOs^ 



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-10° 



Figure I 



Bu.i^nchihys. Geographical distribution. 



114 



CALIFORNIA ACADEM\' OF SCIENCES 




Figure 84. Biif;nchtli\y Above, B niaciii[ncnis. Kapuas 1476-45, 124 mm (MZB .1575); below, B miciuiuiclus. Kapuas 1476-45, 125 mm (MZB 3578, hololype). 



14(2); dorsal-fin spine broken in all three specimens, 108 mm 
male (dorsal spine broken near tip) with at least 23 serrae; pec- 
toral-fin rays 9; peduncular vertebrae 14(2), 15(1); total verte- 
brae 18-19 + 24-25=43(3). 

Distribution. — Sumatra (Musi). Borneo (Kapuas, Barito?). 
Records from Malay Peninsula (Mohsin and Ambak 1 983) and 
Java (.layaram 1968) are erroneous. 

Bagrichthys macropterus (Bleeker, 1853) 

(l-igure S41 

Biii;ri>idcs iihuniplcrus Bleeker, I85.'*/:5I5 (t\pe loialil\ Muara Rompch. Su- 

malra). 
P\t-ndohai^>i, lilh\'\ luairoptents Bleeker, 1862-63:50. 
HiiKiiiula iikhiiipicnii Gunlher, 1864,41. 
IhiiinclilhY^ nhuroplcrus ]ayaxam. 1468:380. 

Material ExAMrNED— Western Borneo: Kapuas 1476-19, 4: 174-236 mm (C.'XS 
44368, MZB 3577); Kapuas 1476-45. 27: 58,4-183 mm (BMNH 1982,3,24,146- 
147, FMNH 94237, IRSNB 631, KUMF 2856, MNHN 1482-702, MZB 3576, 
RMNH 28882, UMMZ 204918, IISNM 230275, ZMA 116.540). Locality un- 
known (= Sumatra, Muara Kompeh'^), 193 mm (RMNH 6876, probable holot\pe). 

Note on Holotipe, — The 193 mm specimen (RMNH 6876) 
probably is the holotype from Muara Kompeh. It is still in a 
jar apparently supplied by Bleeker, and labelled Fscudobag- 
nchthys macropterus. Only one specimen of the species is in- 
dicated in the A group of the auction catalog of Bleeker material 



(none in B, C, D, or E). Therefore this should be the only Bleeker 
specimen of fi, macropterus in the RMNH (as it apparently is) 
and the holotype. The total length of about 256 mm as measured 
by me agrees with Blocker's of 256 mm. The specimen has its 
mouth cut at rictus so it could be more readily opened, and 
tissue overlying the jaws has been partially removed. The upper 
jaw has only two minute teeth visible (one tooth on each side) 
and 3+ I teeth on lower jaw. The description and figure of the 
dentition of 5, macropterus given by Bleeker (1862, pi. 67, fig. 
2) thus seems to differ from that in the holotype and in the 
original description. It may be that the 1862 account is based 
on BMNH 1863.12.4.108. a 117 mm Bleeker specimen from 
Sumatra. This is apparently the only other Bleeker specimen of 
B. macropterus. 

Diagnosis (based on Kapuas specimens). — A Bagrichthys with 
a \ ery small mouth opening and greatly reduced oral dentition; 
inner and outer mental barbels strongly crenulated; dorsal-fin 
spine relatively short, 5.5-7.1. with up to 14 serrae (usually 12 
or fewer); body and fins with strongly defined light marks on a 
darker background. 

The present Kapuas material includes a 215 mm sexually 
mature male and a 236 mm gravid female. The rest of the 
specimens seem to be immature. Apart from the elongate genital 
papilla of males, there seems to be no sexual dimorphism or 
dichromatism. 



ROBERTS-WESTERN BORNEO FISHES 



115 



The following observations are based on 25 Kapuas speci- 
mens 58.5-236 mm: head length 4.0-6.0; eye 32.4-49.7; snout 
10.6-18.1; width of mouth opening 36.6-56.0; width of mouth 
including lips 1 5.2-28.3; nasal barbel, invariably thin and round, 
extending posteriorly to eye by a distance no more than twice 
eye diameter, length 10.8-17.0; maxillary barbel usually failing 
to reach base of pectoral-fin spine, length 4.6-8.5; outer mental 
barbel with weakly or moderately strongly convoluted anterior 
margin, length 10.2-18.9; inner mental barbel with strongly 
convoluted anterior margin, length 20.3-39.3; dorsal-fin spine 
length 5.5-7.1, serrae 7-14 (increasing in number with growth); 
pectoral-fin spine length 5.2-6.6, serrae 13-26 (increasing in 
number with growth); pectoral-fin rays 8(1), 9(22), 10(1); body 
depth 3.7-4.7, width 5.6-6.5; caudal peduncle length 3.4-3.9, 
depth 11.4-14.4; gill rakers 2 + 5=7(2); vertebrae 45(2), 46(6); 
peduncular vertebrae 14-15. 

In addition to the material from Sumatra and Borneo listed 
above, I have examined four specimens from the Mekong basin 
in Thailand previously identified as B. macroptenis (UMMZ 
186765, 186851, 88.8-169 mm). These are morphologically 
close to B. macroptenis but have dorsolateral surfaces of head 
and body uniformly brown (i.e., without the highly visible pale 
or cream-colored blotches and midlateral stripe characteristic 
of Sundaic B. iiiacroplerus). For the present I identify them as 
Bagnchthys sp. undet. (near B. macroptenis). 

Bagrichthys micranodus new species 

(Figure 89) 

HoLorvTE. — MZB 3578, 125 mm mature male. K.apuas mamstream 58 km NE 
of Sintang and 1 km downstream from Sebruang (Kapuas 1976-45). 

Paratypes.-BMNH 19823.29.148, CAS 49369, MZB 3579, USNM 230276, 
8: 59.6-95.8 mm, collected with holotype; MZB 3580, 108 mm mature male, 
Sungai Djentawang near Kctungau, 37-38 km NNE ofSmtangl Kapuas 1976-49). 

Diagnosis. — A Bagrichthys with greatly reduced oral denti- 
tion as in B. macroptenis from which it differs in much smaller 
adult size, generally longer barbels with ribbon-like margins 
(excepting inner mental barbel), slightly longer fin spines, slight- 
ly more elongate caudal peduncle and darker coloration. 

The following observations are based on the 125 mm holotype 
and seven paratypes 59.6-108 mm; head length 4.0-4.7; eye 
27-34.5; snout 10.9-16.2; width of mouth opening 15.3-19.5; 
width ofmouth including lips 1 5.3-19.5; nasal barbel in 3 largest 
specimens 95.8-125 mm (all males) extending posteriorly to or 
beyond gill opening, length 4.4-5.9, but much shorter in other 
specimens, length only 10.0-1 1.0; nasal barbels with expanded 
flat margins in three largest specimens; maxillary barbel ex- 
tending posteriorly to or beyond base of pectoral-fin spine in 
all specimens, its length 3.5-5.8; outer mental barbel with simple 
margins, length 6. 1-9.6; inner mental barbel with strongly con- 
voluted and papillose anterior margin, length 1 8.8-25.2; dorsal- 
fin spine length 5.3-6.5, serrae 6-15 (increasing in number with 
growth); body depth (at dorsal-fin origin) 4.2-4.7; body width 
(at widest point in humeral process) 5.5-6.0; pectoral-fin spine 
length 4.7-5.8; pectoral spine serrae 12-22 (increasing in num- 
ber with growth); pectoral-fin rays 8(3), 9(5); humeral process 
length 8.4-10.1; anal-fin rays 14(3), 15(3), 16(1); caudal pedun- 
cle length 3.4-3.7, depth 13.4-15.6; gill rakers 2 + 3-7=5(1), 
6(1), 8(1), 9(1); vertebrae 19-21+25-26=45(6), 47(1); pedun- 
cular vertebrae 15(2), 16(5). Color in life overall dark brown 
with pale abdomen, large regularly spaced cream-colored marks 



on nape and side of body, and a cream-colored midlateral streak 
extending length of body. Color pattern generally similar to that 
of 5. macroptenis but nape mark generally narrower and failing 
to meet dorsally. 

There are numerous slight differences between B. micranodus 
and B. macroptenis. In addition to those already noted, at com- 
parable sizes the dorsal- and pectoral-fin spines are slightly long- 
er and generally have 2-3 more serrae in B. macroptenis; eye 
and mouth tend to be slightly larger, body slightly wider. 

The three largest specimens, 95.8-125 mm, are fully mature 
males. These have an elongate genital papilla, length 1 2.5-1 7.4, 
and ripe testes with dozens of villus-like projections up to 5 mm 
long. In all other specimens gonads weakly developed and sex 
not determined. 

Etymology.— The name micranodus (Greek) refers to the 
small size and virtually toothless jaws of this species. 

Bagroides Bleeker, 1851 

Biigroides Bleeker, 1851/i:204 (t\pe species Bagroiiies melaplenis Bleeker, 1851, 
hy monotypy). 

D\\G>ios\s.— Bagroides differs from all other bagrids except 
Bagrichthys in having serrae on dorsal-fin spine retrorse rather 
than antrorse or absent; it differs from Bagrichthys in having 
fewer vertebrae; humeral process very large and broad-based; a 
relatively small adipose fin partly free from body posteriorly, 
and a greatly enlarged oval-shaped vomerine tooth patch with 
molariform rather than conical teeth. Dorsal and pectoral spines 
very stout, and heavily serrate; occipital process and predorsal 
plate very strong and in contact; anal-fin rays 1 9-20; vertebrae 
18 + 21-22=39^0, peduncular vertebrae 9-10 {Bagrichthys with 
43^8 total and 1 3-20 peduncular vertebrae). 

Jayaram (1968) placed Leiocassis hirsiitus Herre, 1934 (type 
locality Wuchow, Kwangsi province) in this otherwise mono- 
typic genus. I have compared the 230 mm paratype (CAS-SU 
13885) (same specimen examined by Jayaram) directly with a 
235 mm Kapuas specimen of fi. melapterus (CAS 49370) and 
conclude they are not congeneric. Leiocassis hirsiitus differs from 
Bagroides in having head much broader anteriorly, with a wide, 
broadly rounded snout, relatively wide mouth and interorbital 
space; predorsal profile gently concave rather than gently con- 
vex; posterior margin of united branchiostegal membranes with 
a relatively strong median indentation; vomerine teeth in a 
strongly arched band; dorsal- and pectoral-fin spines relatively 
slender; serrae of dorsal-fin spine relatively weak (as if worn) 
and antrorse rather than strong and retrorse; humeral process 
narrow based; serrae of pectoral-fin spine much smaller and 
probably more than twice as numerous; adipose fin longer, with 
much more extensive thickened portion anteriorly; anal-fin rays 
23 (first 4 rays minute); vertebrae 17 + 24=41. 1 tentatively refer 
Herre's species to Leiocassis. 

Bagroides melapterus Bleeker, 1851 

Bagroides melapterus Bleeker. 1851/r204 (type locality Bandjerniassing. in liu- 

viis). 
Bagroides niclanopierus Bleeker. 1852/':413 {unwarranted spelling change). 

Material Examined. — Western Borneo: Kapuas 1976-19, 4: 85.0-250 mm 
(CAS 49370, MZB 3581. USNM 230277); Kapuas 1 976-48. 97.7 mm (MZB 3582). 

Distribution. — Sumatra (Palembang, Kwantan R,, Indra- 
giri, Djambi). Borneo (Sambas. Kapuas, Barito. Kahajan), We- 



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CALIFORNIA ACADEMY OF SCIENCES 



Table 5. Meristic Characters of Some Sundaland Le/oc.4Ssis. For definition of vertebral counts see p- 





Gill rakers 


Anal-fin rays 


Pei 


:toral-fin 


rays 


Vertebrae 


L- annatits 
Kapuas 


6-8? 


14-15 




6 




14-164-17-18=32(2). 33(2) 


L of. ZiMtm Popta. 1904 
Johore (CAS-SU 39339) 


12 


12-13 




6-7 




17 -(-18=35(3) 


L leiM-amlms W and de B. 1912 
Johore (CAS-SU 31006) 


10 


15-17 




7-8 




16 + 20=36(2) 


L micwpogon 
Kapuas 


12 


17 




8 




18-1-21=39(1) 


L myersi 
Kapuas (types) 


16-18 


17-18 




7 




16-17-1-26-27=43(3), 44(2) 



ber and de Beaufort (1913:349) give Rajang R.. but the basis 
for this record is unknown. 

Leiocassis Bleeker. 1858 

Leiocassis Bleeker. 1 858c :59, 1 39 (type species Bagrus pucciloplerus Valenciennes 
idCuvierand Valenciennes. 1839. by subsequent designation of Bleeker. 1862- 
63:9). 

Z.j(Hins/v Gunther. 1864:65 (unwarranted spelling enicndation). 

'^P'.cuJomysnis Jayaram, 1968:359 (type species Bagriis slcnoiiim Valenciennes 
//iCuvierand Valenciennes. 18 39, by original designation; described as subgenus 

of /.<7<l("t/VV/v), 

Although Jayaram (1968) reviewed Leiocassis. his work is 
based mainly on a survey of the literature. The only compre- 
hensive revision of the genus is that by Regan (1913). based on 
relatively few specimens and comprising relatively brief de- 
scriptions without figures. Several species have been descnbed 
subsequent to Regan, but it is clear from examination of mu- 
seum collections that additional species remain undescribed. 

It seems likely that the numerous species presently included 
in Leiocassis are of diverse phyletic relationships (i.e., poly- 



phyletic). Jayaram ( 1 968) divided Leiocassis into two subgenera, 
Leiocassis and Pseiidoniysliis. Within Pseiidomyslus he recog- 
nized two groups, the "leiacanlhus complex" with leiacanthus, 
siamensis, and bicolor. and the "stenomiis complex" with sten- 
onitis, vaillanti. inonialus. mahakamensis. fiiscus, moeschii. 
hreviceps. and rohustiis. The "leiacanthus group." with only three 
species, may be natural, but the "steiioniiis complex" appears 
to be very diverse and probably polyphyletic. Jayaram (1968: 
359) remarked of this group that the species resemble those of 
the genus Mysliis. I have myself noted the superficial resem- 
blance of L. stenomiis, L. mahakamensis, and L. cf fiiscus to 
Mysliis (or Hemibagnis) and am inclined to believe it may in- 
dicate a true relationship. In L. ci, fuscus and L. mahakamensis 
the branchiostegal membranes of the left and right sides are 
strongly overlapping (as in Mystiis) whereas in all members of 
the subgenus Leiocassis and leiacanthus complex of Pseudo- 
inystus. so far as I have been able to determine, they are non- 
overlapping. 

Meristic characters of some Sundaland Leiocassis are given 
in Table 5. 




Figure 90. Leiocassis armatus Kapuas 1976-39, 24 8 male (CAS 49427). 



ROBERTS-WESTERN BORNEO FISHES 



117 




Figure 91. Leivcussis micropogon. Rapuas 1976-27, 85.2 mm (MZB 3586). 



Leiocassis armatus (Vaillant. 1902) new combination 

(Figure 90) 

Ikvsis armanis Vaillant, 1902:64 (type locality Tepoc, bonds du Mahakam). 

Material Examined. — Western Borneo: Kapuas 1976-21, 7: 19.2-2.1.5 mm 
(BMNH 1982.3.29.149-150. CAS 49371, MZB 3583, RMNH 28883); Kapuas 
1976-39, 1 1: 17.9-24.8 mm (CAS 49372, MZB 3584, USNM 230278). Eastern 
Borneo: Mahakam basin, Tepoe, 21.7 mm (RMNH 7844, holotypcof.-l. armaliis). 

Direct comparison of several Kapuas specimens with the ho- 
iotype and only previously known specimen of "Akysis" ar- 
matus reveals that they are identical in all respects. The species 
does not have the characteristic features of the family Akysidae. 

Leiocassis armatus is perhaps the smallest member of the 
bagrid genus Leiocassis (largest known specimen, a sexually ma- 
ture male with elongate genital papilla. 24.8 mm). 

Distribution. — Kapuas. Mahakam, 



Leiocassis micropogon (Bleeker, 1852) 

(Figure 4 1) 

Bagrus micropogon Bleeker. 1852c/:94 (type locality Blilong [ = Billilon], in flumine 

Tjirutjup). 
Lc'incassis micropogon Bleeker, 1858t:142. 

''Liocass!s baramensis Regan. 1906:67 (type locality Baram River) 
""Liocassis hosii Regan, 1906:67 (type locality Sibu, Sarawak), 
'^Liocassis merahciisis Regan, 1913:550 (type locality Merahch. North Borneo). 
''Liocas-sis donac Regan. 1913:551 (type locality Borneo). 
'' Leiocassis citaseni de Bt:^n(on. 1 933:34 (type locality Ulujelai, Malay Peninsula), 

See Hora and Gupta (1941:25). 
"^Leiocassis regain Jayaram, 1965:9 (type locality Sadong, N. Borneo). 

Material Examined. — Malay Peninsula: outlet of Lake Chin-chin Jasm, Ma- 
lacca. 148 mm (C.AS-SU 31005). Western Borneo: Kapuas 1976-20, 2: 129-183 
mm (C.\S 49373, MZB 3585); Kapuas 1976-27, 85.2 mm (MZB 3586); Kapuas 
1976-28. 8: 32.9-105 mm (BMNH 1982.3.29. 1 51-152. CAS 49374, MZB 3587, 
RMNH 28884, USNM 230279). 

Leiocassis micropogon is closely related to L. poecilopterus 
(generic type species) but differs in having a more elongate head 
(especially snout) and body (especially caudal peduncle). It is 
rather variable in such respects as relative development of oc- 
cipital crest and predorsal plate, development of a posterior 
median projection from the vomerine or palatal toothplate, depth 
of caudal peduncle, and coloration (from subdued brownish with 



relatively indistinct mottling to relatively well defined dark marks 
on an orangish or yellowish background). 

The 1 83 mm Kapuas specimen, a gravid female (CAS 49373) 
has head length 3.1; snout 9.4; eye more than 3 times in snout, 
about 3 times in interorbital space, its length 32.7; soft inter- 
orbital width 16.4; barbels uniformly thin; nasal barbel very 
small, length 61 (less than eye diameter); maxillary barbel ex- 
tending posteriorly a little beyond eye, length 8.2; outer mental 
barbel 1 2.0 and inner 42; palatal tooth band with a broad-based 
short posterior median extension; branchiostegal rays 7-8?; gill 
rakers 3 + 9=12; predorsal plate elongate, separated from occip- 
ital process by a distance about equal to eye diameter; dorsal 
spine with 20 serrae. length 6,0; pectoral spine with 20 serrae, 
length 6.7; pectoral-fin rays 8; humeral process extending pos- 
teriorly as far as middle of adpressed pectoral spine, length 10,2; 
anal-fin rays about 14 (4 simple and 10 branched); adipose fin 
moderately long, with a well defined posterior insertion but 
thickened anteriorly with origin poorly defined, length about 
5.0; caudal peduncle length 6.0, depth 13.8, 

The nominal species of Regan ( 1 906, 1913), preceded by ques- 
tion marks in the synonymy of L. micropogon presented here, 
are all known only from one or a few type specimens from single 
type localities, I examined these specimens and tentatively con- 
clude that they represent individual or local variations of L. 
micropogon. L. doriae was distinguished because the 177 mm 
holotype and only known specimen has a vomerine or palatal 
tooth band with an exceptionally large median posterior pro- 
jection; in other respects it appears to be identical to other large 
specimens of L. micropogon which tend to have a variably de- 
veloped posterior projection. The syntypes of L. hosii have a 
more elongate or slender caudal peduncle than other specimens 
examined but in other respects seem indistinguishable from L. 
micropogon. 

Distribution, — Malay Peninsula. Sumatra. Borneo (Sara- 
wak, Sambas, Kapuas). Banka. Billiton, 

Leiocassis myersi new species 

(Figure 42) 

Holotype. — MZB 3588. 71.2 mm. small forest stream flowing into Kapuas 
mainstream NE oTGunung Sclunggul. 53 km NW of Sintang (Kapuas 1976-47). 



118 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 92. Lciocassis mymi. Kapuas, 1476-47, Above. 71.2 mm (MZB 3.S88, hololype), middle and below, 46.8 mm (CAS 49375, paratypcl. 



Paratypes.-CAS 49.375 (2), MNHN 1982-703 (1). MZB 3589 (2). USNM 
230280(31.9: 38,9-87.8 mm, same colleclion as hololype, BMNH 1982,3,29,153- 
154 (2), MZB 3590 (1), RMNH 28885 (1), 4: 52,8-55,9 mm, Sungai Seriang, a 
tributary to Sungai Palm, Rapuas hasm, 37 km W of Putussibau (Kapuas 1976- 
42), MZB 3591, 44,9 mm, Sungai Gentu, near where it Hows into Kapuas main- 
stream, 55 km NE of Sintang (Kapuas 1976-46). 



Diagnosis. — A Lciocassis with a relatively broad, moderately 
depressed head; dorsal surface of cranium rugose, exposed or 
covered only by a thin layer of skin which may be easily abraded 
oflF; pores of cephalic laterosensory canals relatively large and 
highly visible due to depigmented rims; adipose fin elongate and 



ROBERTS-WESTERN BORNEO FISHES 



119 




• *■■■ t^'V*^ jfJk ■• "'''<.'■ 



isf**' 




5 mm 



Figure 93. Lciocassts millanli Kapuas, 107 mm (RMNH 7840, hololypc; Icfl, after Vaillanl 1902:61). 



relatively low, extending posteriorly onto anterior portion of 
ridge formed by upper procurrent caudal-fin rays (in all other 
Leiocassis examined adipose fin terminates posteriorly directly 
on dorsal surface of trunk or caudal peduncle); gill rakers 16- 
18; anal-fin rays 17-18; caudal-fin rays 24 + 8/9+16-17; ver- 
tebrae 16-17 + 26-27=43-44; color in life bluish-black; side of 
body with a thin pale or white midlateral streak, two horizontally 
elongate pale or white oval spots above midlateral streak and 
three similar oval spots below it; small pale marks at origin and 
termination of dorsal fin, but dorsum without pale transverse 
bands; rayed fins overall pale but with one or two dark transverse 
bands; adipose fin black with pale edges or marginal spots. 

The following observations are based on the 71,2 mm ho- 
lotype and four paratypes 55,1-87,8 mm: head length 3,3-3,7. 
gill cover elongate, extending posteriorly to above middle of 
humeral spine; snout broadly rounded, length 9,9-10,6; eye 



without free orbital rim, horizontal diameter 29,0-31,6; mouth 
width 9,2-10,7; interorbital space broad, slightly concave, width 
13,5-14,8; occipital crest and predorsal plate rugose and mod- 
erately elongate, of equal length and similar shape, and nearly 
or quite meeting each other; predorsal plate length 16,0-18,7; 
body depth 4,9-5,5 and width 4.4-4,9; barbels thin, moderately 
elongate; nasal barbel extending posteriorly well beyond eye, 
7,5-9,5; maxillary barbel extending almost to or slightly beyond 
base of pectoral-fin spine. 4, 1-5,4; outer and inner mental bar- 
bels respectively 6,1-8,8 and 8,5-12,7; dorsal-fin spine mod- 
erately elongate, 5.3-6,1; adipose-fin base length 2,7-3,2 and 
height 16,4-20,9; pectoral-fin spine length 5,0-5,5; pectoral ser- 
rae 1 1-16 (increasing in number with growth); pectoral-fin rays 
7; humeral process rugose, moderately elongate. 9,5-9,8; su- 
praclavicular spine well developed, pointed distally; pelvic fin 
large, extending posteriorly to anal-fin origin, length 6,1-7,2; 




Figure 94. Leiocassis sp. undet. Kapuas. 75.2 mm (MNHN 1891-475). 



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CALIFORNIA ACADEMY OF SCIENCES 



pelvic-fin rays 6; caudal fin moderately forked, with pointed 
lobes, upper slightly longer than lower. 

Distribution. — Known only from Kapuas. 

Etymology.— This species is named in honor of Professor 
George S. Myers, my teacher in ichthyology and himself a stu- 
dent of Asian fishes. 

Leiocassis vaillanti Regan, 1913 

(Figure 9.1) 

Leiocassis moeschii VaiWanl. I'^02:61 (non Boulcngcr. 18')0), 
Leiocassis vaillanli Regan. 191.1:54')(type localils cnibouchiiredu Raoun. Rapuas 
basin). 

Material Examined —Western Borneo: Kapuas basin, embouehuredu Raoun. 
107 mm (RMNH 7840, hololype). 

Leiocassis vaillanti is known only from a description and 
figures by Vaillant (1902:61-63. fig. 8. 9), who identified it as 
L. moeschii Boulenger, and a very brief description by Regan 
(1913), who recognized it as a new species, ofa single specimen 
from the Kapuas basin. In addition to reproducing Vaillant's 
figures I give an illustration of the oral dentition. 

This species has dorsal surface of head consisting mainly of 
exposed, rugose or granulated bone. As noted by Regan, the 
"supraclavicular"" (=supracleilhrum?) bone lacks the posterior 
spinous projection characteristic of L. mocschii and is in fact 
strongly concave where the spine would be; this condition is 
accurately illustrated in Vaillant's figures. The anterior margin 
of the exposed bony head shield lies far posterior to the level 
of the eyes, whereas in L moeschu it is continued anteriorly as 
a pair of elongate projections extending in front of the eyes. L. 
vaillanti also has a broader, less pointed snout, a wider mouth, 
and a more elongate body than L. niocscliii: gill rakers 3+10=13; 
pectoral spine with 16 strong serrae; pectoral-fin rays 8; pelvic 
6; anal 16. 

Leiocassis sp. undet. 

(Figure 44) 

Leiocassis siciioiniis Vaillant. 184.^7.1 (non Valeneiennes in Cuvicr and Valen- 
ciennes. 1839; Knapei. Kapuas basin). 

Material Examined —Western Borneo: Kapuas basin. Knapei. 75,2 mm 
(MNHN 189 1-475, specimen identified as L siciioiiius by Vaillant). The following 
additional lot of Kapuas specimens, examined briefly and not directly compared 
with the preceding specimen, may also be the same species: RMNH 7837. 12: 
20.0-70 7 mm. Sintang. 

Mystus Scopoli. 1777 

,\/rs7Ky Scopoli, 1777:451 (type species undetermined, lor discussion see Jayaram 

1962). 
Miicroiic^ Dumenl, 1856:484 (type species Baiini\ lanidin Valenciennes in Cuvier 

and Valenciennes. 1839, preoccupied by Macroncs Newman. 1841.Coleoptera). 
HcmilHigrus Bleeker. 1862-63:9 (type species Bagrus nemwm Valenciennes 

in Cuvicr and Valenciennes. 1839. by original designation), 
hpntohagius Bleeker. 1862-63:9 (type species Hagnis giilio Hamilton- Buchanan, 

1823, by original designation and monotypy), 
Hypsetotiagnis Bleeker. 1862-63: 10. 57 (type species Bagnis nuicn'ncina Bleeker. 

\%At^ Bagrus nignccps Valenciennes /// C'u\ier and Valenciennes, 1839, by 

original designation), 
Hclcrohagrus Bleeker. 1864:355 (type species llclcrol'agnis Ivcoiirli Bleeker. 1864. 

b> original designation and monotypy). See discussion under MysUis nignccps 
\ona Jordan. 1919:341 (substitute name for Macmncs Dumenl, 1856; preoccu- 
pied by Anna Baly. 1863, Coleoptera), 
Spciala Holly, 1939:143 (siihsiiiutc name tor iorui Jordan, 19)4, and therefore 



with same type species; Holly's designation of Siliinis vilialus Bloch as type 

species IS invalid), 
Aoruinlivs'Wu. 1939:131 (substitute name for , Jtinu Jordan, 1919), 
Macmnicliltiys White and May-Thomas, 1940:505 (substitute name for Aona 

Jordan, 19 |9). 

Mystus Scopoli, 1777 is a traditional or catch-all genus (re- 
vived by Fowler 1 928: 1 05 to replace the preoccupied traditional 
genus Macrones Dumeril, 1856) to which approximately 30 
relatively generalized Asian bagrid species are assigned. 

The nomenclatural history of Myslits is extremely compli- 
cated and requires further study before Jayaram's conclusions 
concerning its type species can be accepted. 

Some species of Mystus (e.g., M. nemurus. type species of 
Hemihagms Bleeker) are superficially extremely similar to the 
genus Bagrus (type species Siliinis docmak F6rskal, 1775). This 
African genus comprises about eight species which are mor- 
phologically relatively similar. They differ from Mystus in hav- 
ing 8-1 1 dorsal-fin rays rather than the generalized number (for 
catfishes) of 7. 

Bleeker ( 1 862-63). in characteristically bold fashion, divided 
Macroncs or Mystus into four genera, but these are not recog- 
nized here pending further study of their morphology and de- 
termination of the type species oi Mystus Scopoli, 1777. 

Mystus micracanthus (Bleeker. 1846) 

Bdgiu-. inicidcanlluis Bleeker, 1846i(:15l (type locality Batavia, in lluviis). 
JI]-psclohagriis micracanllnis Bleeker. 1862-63:59, 
Macrones micracanthus Giinther. 1864:76, 
\lysnis mtcracantlnis Fowler. l934/i:94. 

Material Examined, — Malay Peninsula: Pcrak, Telok ,^nson. 132 mm (CAS- 
SU 30988); Malacca. 57,7 mm (CAS-SU 30985); Johore. Mawai. 66.6 mm (CAS- 
SII 30986); Johore. Sungci Kayu, 105 mm (CAS-SU 32715); Singapore, 2: 67,6- 
113 mm (CAS 53308, CAS-SU 30987), Sumatra: Padang, 106 mm (CAS-SU 
8014): Indragiri River. 112 mm (ZMA 119,059); Batang Han. 6: 72,5-98.1 mm 
(ZMA 1 19.060); Gunung Sahilan. 4: 60,5-65,5 mm (ZMA 1 14.393); Lampong. 
107 mm (RMNH I 5858); Moesi River at Moera Klingi. 8: 40,7-102 mm (UMMZ 
155689. 155713^), Borneo: Kapuas 1976-8, 47 7 mm (MZB 3592); Kapuas 1976- 
16. 40,2 mm (MZB 3593); Kapuas 1976-17. 2: 40,4-82,0 mm (BMNH 
1982,3,29.155. MZB 3594); Kapuas 1976-19. 3: 62,7-68,0 (CAS 49376, MZB 
3595); Kapuas 1976-32, 4: 67,8-83.4 mm (FMNH 94238, MZB 3596); Kapuas 
1976-33. 5: 60,6-90,8 mm (MZB 3597. USNM 230281): Kapuas 1976-36. 77.1 
mm (MZB 3598); Kapuas 1976-37. 5: 73,8-89,5 mm (CAS 49377. MZB 3599, 
RMNH 28886); Kapuas 1976-39. 94,1 mm (MZB 3600); Kapuas 1976-42. 2: 
73 .5-84,1 mm (KUMF 2857. MZB 3601); Kapuas 1976-43. 41,1 mm (MZB 
3602); Kapuas 1976-44, 60,6 mm (MZB 3603); Kapuas 1976-51. 3; 48,1-54.1 
mm (MZB 3604. UMMZ 209926). Java: Batavia, 106 mm, Bleeker collection 
(RMNH 15847. neotypc);Krawang. Ill mm (RMNH 2947); Brantas River, 78,1 
mm (RMNH 5206); no precise locality, 2 101-120 mm (RMNH 2948. 3009); 
Bleeker collection, no locality: 27: 62,7-122 mm (RMNH 5216), 

DwGNOSxs,. — Mystus micracanthus is a narrow-headed, mod- 
erately large-eyed species with a very long adipose fin and very 
long barbels. In these respects it agrees with M. nigriceps, but 
IS less elongate, with a shorter cranial fontanel and different 
coloration. Maxillary barbel extending posteriorly to or beyond 
end of caudal fin; outer mental barbel extending posteriorly to 
origin of pelvic fin. Color in life overall brownish. Area overlying 
entire tympanum moderately darkened, and caudal peduncle 
with a large, triangular midpeduncular spot, these two moder- 
ately darkened areas not outlined or otherwise intensified by 
pallid or white contiguous areas. 

Designation and Description of Neot'ipe. — Bleeker did 
not specifically mention any specimen(s) utilized in the original 
description of .A/, micracanthus. The RMNH possesses a single 



ROBERTS-WESTERN BORNEO FISHES 



121 



Table 6. Mystus micjuca/^thvs. Geographical vanalion in iVcqucncics of gill rakers and vertebrae. 



Gill rakers 



Vertebrae 



20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 



36 37 38 39 40 



Malay Peninsula 
Tclok Anson, 132 mm 
Malacca, 57.7 mm 
Mawai, 66.6 mm 
Sungci K.ayu, 105 mm 
Singapore. 67.6-113 mm 

Sumatra 
Padang, 106 mm 
Indragin, 1 12 mm 
Batang Han, 98.1 mm 
Batang Hari, 72.5-77.5 mm 
G. Sahilan, 60.5-65.5 mm 
Lampong, 107 mm 
Moesi, 40,7-102 mm 

Borneo 
Kapuas, 40,2-94.1 mm 

Java 

Batavia, 106 mm (neotype) 
Krawang, 1 1 1 mm 
Brantas, 78.1 mm 
"Java," 101-120 mm 

Bleeker Col, (Java?) 
no loc, 62.7-122 mm 
no loc, 74.3-109 mm 



1 1 



1 1 



1 1 



1 I 



1 3 



1 3 



5 12 



14 3 2 3 1 



3 3 4 

2 6 7 1 



Blocker specimen from Batavia, the type locality. RMNH 15857. 
106 mm. with broken anal and caudal fins but otherwise in good 
condition. The specimen has a piece of parchment tied to it but 
the writing on the label is indiscernible. It may well have been 
the actual specimen (or at least one of those) upon which Bleek- 
er"s original description was based. The only other Bleeker spec- 
imens of ,U. niicracanthiis in Leiden are RMNH 5216. 27 spec- 
imens without locality data, I therefore designate RMNH 15857 
as neotype. The specimen may be described as follows: head 
3.8, snout 9,8. eye 16.8. bony interorbital width 1 1,8; cranial 
fontanel extending posteriorly to anterior border of supraoccipi- 
tal bone; nasal barbel reaching posteriorly almost to end of head 
its length 5,7, maxillary barbel reaching to middle of caudal 
peduncle, its length 1,14; outer mental barbel 2,4. inner 3.4; 
dorsal-fin spine 7,1. pectoral 4,7; pectoral-fin spine with 16 
serrae on posterior margin; humeral spine 9,1; body depth 4,3. 
width 5,0; caudal peduncle depth 10,0; adipose-fin base about 
3,2; dorsal-fin rays 7-1/2, anal about 13. pectoral 9. pelvic 6; 
gill rakers 6 + 21=27; branchiostegal rays 9/9; vertebrae 
21 + 18=39. 

Intraspecific Variation. — Mystus inicracanthus exhibits 
differences in frequencies of counts of gill rakers and vertebrae 
(Table 6) interpreted by me as intraspecific variation. In Su- 
matra one finds most specimens with relatively few gill rakers 
and more vertebrae, but there is one sample (from Batang Hari) 
with more gill rakers and few vertebrae. Specimens from the 
Malay Peninsula examined by me all correspond to the first 
type, while those from Borneo (Kapuas) correspond best to the 
second. This pattern suggests that two species might be involved. 
Specimens from the Malay Peninsula. Sumatra, and Borneo 
with few gill rakers and more vertebrae tend to be larger than 



those with more gill rakers and few vertebrae, which also sug- 
gests two species. The picture is complicated, however, by spec- 
imens from .lava and from the Bleeker collections (unfortunately 
without locality data) in which the range of gill raker counts is 
very wide but vertebral counts do not differ in specimens with 
low and high gill raker counts. Another complication is that 
specimens from the Kapuas with low vertebral counts exhibit 
almost as great a range of gill raker counts as those from the 
entire range oi Myslits micracanthus. It would be interesting to 
examine additional material, especially from the Malay Pen- 
insula and Borneo (including Sarawak), to determine whether 
the patterns described here really exist. 

Note on Mystvs Bi,\tACUL4Tus. — Myslus bimacidatm Volz, 
1904, was placed as a junior subjective synonym of M, micra- 
canthus by Weber and de Beaufort (1913:339) and has not been 
recognized subsequently as a valid species. Although closely 
related to .U, micracanthus. it is immediately distinguished by 
its striking coloration, overall chocolate brown, with a large 
black humeral spot (round and well defined) and a large black 
vertical bar on posterior portion of caudal peduncle and base 
of caudal fin; anterior margin of humeral spot and both margins 
of caudal bar set off by vertical cream or white bars, A color 
photograph of live A/, bimaculatus from Djambi appears in 
Vogt (1979), Specimens examined by me (ZMA 119,064, 2: 
38,8-39, 1 mm, Bagan si api-api) have 23-24 gill rakers and only 
33-35 vertebrae. The species apparently is restricted to Sumatra. 

Mystus nemurus (Valenciennes, 1839) 

Bagnis /witnims Valenciennes ni Cuvier and Valenciennes, 1 839:423 (type locality 

Java). 
Bagrw, Hoevenii Bleeker, 18460:154 (type locality Batavia, in fluviis). 



122 



CALIFORNIA ACADEVn' OF SCIENCES 



ftje/Hs Sicholdii Bleeker, lS46u:l55 (tspc lotalil\ Bala\ la. in fluviis). 
'Wlacrtines hmgan Popla. 1904 1X2 (typo lixalily "le Bongan" = Kapuas basin). 

See below. 
Macnvws pUiiuccp^ Vaillant, 1 402: 1 8. 20. 23 (Siliau. Raoen, Kapuas hasin. Bloeoe. 

upper Mahakam). 
Mvslus neiinirus Fowler, l')34/i:'^5- 
Mvsms johorensis Herre, l<)40u;13 (t\pe loealiu Siingai Ka>ii. lb miles north of 

Kota Tinggi, Johore). See below. 
Mrslus pahangeiisis Herre, 1M40,(:14 {i\pe loeality Sungai Garam, near Karak, 

Pahangl- See below. 
''htacriincs (Hcniihugnn) tiUii)]ciiiiis Fang and C ham in Chaux and Fang, 194'); 

200 (t\pe locality Camhodgc) 

MATERIAL Examined. — Thailand. Chantahun, Kao Sabap. 144 mm (CAS-SU 
28825), Malay Peninsula: Pahang, Sungai Garam near Karak, 196 mm (CAS-SU 
33025, holotype M- paliani;c)isis): Perak. Telok Anson. 2: 77,0-85,6 mm (CAS- 
SU 30999); Perak. Chanra Dam, 2; 216-231 mm (CAS-SU 32717); Perak, Bukit 
Merah, 3; 98.4-161 mm (CAS-SU 30997, 32714); Kuala Lumpur, Selangor, 3: 
83. .3- 1 43 mm (CAS-SU 327 1 6); Malacca. Lake Chin Chin. 2; 66. 8-95, 6 mm (CAS- 
Sll 30996); Johore. Sungei Kayu. 208 mm (CAS-SU 33026. holotype M jolwr- 
cmis): .lohorc. Sungei Kayu. 200 mm (CAS-SU 3272 1 ); Johore, Kota Tinggi. 140 
mm (CAS-SU 32718): Johore, Ayer Hiiam. 1 13 mm (CAS-SU 31000); Singapore, 
147 mm (CAS-SU 31001), Western Borneo: Kapuas 1976-6, 4; 106-192 mm 
(BMNH 1982,3,29,156-158, MZB 3605); Kapuas 1976-15, 2: 20,6-24,6 mm 
ICAS 49378, MZB 3606); Kapuas 1976-16. 5: 71.3-168 mm (CAS 49379, MZB 
3607, ROM 37406); Kapuas 1976-17, 2: 80.4-86.5 mm (FMNH 94239. MZB 
3608); Kapuas 1976-21, 80 5 mm (MZB 36(')9); Kapuas 1976-24, 2; 61. 9-123 
mm (IRSNB 19747. MZB 3610); Kapuas 1976-25. 11: 49.8-188 mm (MNHN 
1982-704. MZB 361 1. RMNH 28887); Kapuas 1976-27. 5: 44.6-85 7 mm (MZB 
3612. LISNM 230282), Kapuas 1976-28, 2: 80,1-89,0 mm (MZB 3613, ZMA 
1 16,541); Kapuas 1976-29. 76.7 mm (MZB 3614); Kapuas 1976-30, 6: 40,9-56,9 
mm (CAS 49380, MZB 3615); Kapuas 1976-42. 99.8 mm (MZB 3616); Kapuas 
1976-46, 79.1 mm (MZB 3617); Kapuas hasm. Nanga Raoen. 145 mm (RMNH 
7828); Kapuas basin, Sihau, 93 3 mm (RMNH 7829) Eastern Borneo: Mahakam 
hasin, Bloeoe, 2: 63 1-84 5 mm (RMNH 7827) 

Diagnosis. — A/r5/»,v iwDuinis (type species of HcDiihagfiis 
Bleeker, 1862) has a broacd, Hal head; moderately large eyes; 
wide intcrorbital space; cranium moderately rugose dorsally; 
cranial fontanel extending well into posterior half of supraoc- 
cipital bone, almost to base of supraoccipital crest; supraoccipi- 
tal crest extending posteriorly almost to predorsal bone, barbels 
long, maxillary barbel extending posteriorly almost to 
anal-fin origin to somewhat beyond it; adipose fin large but 
short, originating far posterior to dorsal fin, with its dorsopos- 
terior margin distinctly rounded and its base incised posteriorly; 
upper caudal-fin lobe often with a short filamentous extension; 
color in life overall dark brown, with a well defined, dark, thin 
midaxial streak extending length of body (absent in all other 
Mystiis); branchiostegal rays 1 1-13; gill rakers 3-5 + 9-14=13- 
19; vertebrae 22-23 + 22-23=43-45. 

The thin, dark midaxial streak, probably always present in 
fresh specimens, juvenile as well as adult, apparently occurs 
only in this species among Mystus. This otherwise diagnostic 
character may fade entirely after long periods in preservative. 
I have examined the holotypes (which are also the only reported 
specimens) of .1/, paluni^ensis and M. johorcnsis: both have an 
ob\ ious midaxial streak and agree in all respects with M. ne- 
inurus. 

Remarks on Related Species. — A/riY(« nemwus seems 
closely related to M. pkuuceps and M. haninwnsis {Regan 1 906). 
Mystus baramensis is known only from northern Borneo, where 
it occurs sympatrically with M. ncmunis (see Inger and Chin 
1962:138-141, 202-204). As noted by Inger and Chin, M. bar- 
i(/»(7(.v;.v differs from M lu'iiiiirus in coloration, adipose-fin shape, 
and gill raker counts. M. hanimcnsis has no midaxial streak. Its 
adipose-fin base is more elongate. 



Gill rakers vary considerably in both species, but generally 
are more numerous in Af. barai>icnsis (see Table 7 and discus- 
sion in Inger and Chin 1962). Finally, having examined several 
lots of both species, I find M. baramensis slightly stockier and 
less elongate, with a shorter or stouter caudal peduncle, and with 
its cranial roof more covered with skin and less rugose. The 
largest reported specimen of M. baramensis is only 172 mm 
standard length (Inger and Chin 1962), whereas Af. nemurus 
has been reported to 350 mm (total length?) (Weber and de 
Beaufort 1913:342). Thus AI. baramensis may be a smaller 
species, although limited data available (Table 7) indicate this 
is not reflected by marked decrease in vertebral number. 

Alystus planweps (Valenciennes //; Cuvier and Valenciennes, 
1839), type locality Java, is similar to A!, nemurus but differs 
in having shorter barbels, slightly shorter snout, broader inter- 
orbital space, less rugose dorsal surface of cranium, supraoccipi- 
tal process relatively short (separated from predorsal plate by a 
distance equal to about twice eye diameter, vs. usually less than 
one eye diameter in AI. nemurus). possibly a pale rather than 
dark midaxial streak, and more vertebrae. Gill rakers 16-17. 
Vertebrae 46-49 (Table 7), 

Alystus planieeps probably does not occur in Borneo, although 
it has been reported from northern Borneo by Boulenger (1894: 
247) and Inger and Chin (1962:141), from western Borneo by 
Vaillant (1893:38; 1902:18, 20), and from eastern Borneo by 
Vaillant (1902:23). I have examined the specimens reported 
upon by Vaillant ( 1 902) (RMNH 7827-29) and reidentified them 
as AI. nemurus. I have also examined some of the specimens 
studied by Inger and Chin (1962) (FMNH 68064, 68092-93). 
These specimens, from the Rejang basin, differ from AI. plan- 
ieeps from Buitenzorg. Java (CAS-SU 20505) and are unlike 
other Bomean specimens of Alystus examined by me. 

Distribution. — .\/vi7«,s nemurus is common and widely dis- 
tributed in Thailand (Smith 1945:386). Malay Peninsula. Su- 
matra, Borneo, and Java. It apparently occurs in almost every 
river basin in Borneo (Weber and de Beaufort 1913:342; Inger 
and Chin 1962). 

Mystus nigriceps (Valenciennes. 1839) 

/(i(i,T(n luf^nccps Valenciennes in Cuvier and Valenciennes, 1 839:4 1 2 (type locality 

Jaxa) 
/)'ij.'^'/»v niacniiu'iiiiis Bleeker. l846ii:15U (type locality Batavia. in lluviis). 
Ikiiirwi singanngai' Bleeker. 1846u;l50 (type locality Batavia. in fluviis). 
fhiK'tn helcninis Bleeker. 1846ij:151 (type locality Batavia. in fluviis). 
llvpsclohagnts niacroneina Bleeker. 1862-63:58, 
.Macronei /i/^nccps Giinther. 1864:77. 
Hypsclolwgnis nignccps Fowler. 1905:470, 
.MvsliLs nignccps Fowler. 1935i;:94. 
.Mysliis carasius Smith, 1945:94 (in part). 
Macroncs cavasius von Martens. 1876:400 (Kapuas, Danau Sriang), 

.Material ExAMtNED. — Mala\ Peninsula: Malacca, 3: 44.9-76,9 mm (CAS-SU 
30995); Perak near Sauk, 1 25 mm (CAS-SU 32720); Perak. Chandra dam. 2: 190- 
198 mm (CAS-SU 30992); Johore, Ayer Hitam, 87,9 mm (CAS-SU 30993); Pa- 
hang, 2: I 1 1-132 mm (CAS-SU 30998). Singapore, 66.7 mm (CAS 20003), West- 
ern Borneo: Kapuas 1976-8. 12: 67,1-152 mm (BMNH 1 982,3,29, 1 59-1 60. CAS 
49381, FMNH 94240, MNHN 1982-705. MZB 3618); Kapuas 1976-19. 2: 112- 
133 mm (MZB 3619, RMNH 28888); Kapuas 1976-20. 3: 109-119 mm (MZB 
3h:(l UMMZ 209863); Kapuas 1976-33, 105 mm (MZB 3621); Kapuas 1976- 
34 1 \^ mm (MZB 3622); Kapuas 1976-43,93,5 mm (MZB 3623); Kapuas 1976- 
44. 4 1 12-127 mm (MZB 3624. USNM 230283), Java: Buiten/org, 161 mm 
(CA.S-SLl 20492). 

Diagnosis. — .A/, nigrieeps is readily distinguished from other 



ROBERTS- WESTERN BORNEO FISHES 



123 



Table 7 Frequencies of Gill Raker and Vertebral Counts in Mystus nemvkus and Related Species. 



Gill rakers 



Vertebrae 



13 



16 17 



18 



\<i 



43 44 45 46 47 48 44 



Mystus baraincnsis 
Northeast Borneo 
Kabili R. (CAS-SU 32711,-13,-19) 
Sibugal R. (CAS-SU 33738) 
Sandakan (Inger and Chin, 1962) 
Kalahakan (Inger and Chin, 1962) 

Mysliis nemunis 
Thailand 
Kao Sabap 
Singara 

Malay Peninsula 
Pahang (hololype pahangensis) 
Telok Anson 
Bukit Merah 
Chandra Dam 
Malacca 
Kuala Lumpur 
Johore (holotype johorensis) 
Singapore 

Western Borneo 
Kapuas 1976-16 
Kapuas 1976-30 

Mvstiis planuvps 
Java 

Builen/org 
\tys!i{s cf. planuvps 
Sarawak 
Rejang R. (FMNH 68092) 



1 1 

1 1 



1 2 



1 3 



4 12 2 



Mysliis in the Malay Peninsula and Indonesia by its long adipose 
fin contiguous with the dorsal fin (origin of adipose fin at a 
distance from dorsal fin in other species). Barbels very long, 
maxillary barbel sometimes extending to or beyond caudal fin. 
Branchiostegal rays 7-8. Gill rakers 21-25. Vertebrae 41-44 
(Table 8). 

Smith (1945:94) treated M. nigriceps as a junior synonym of 
M. cavasiiis (Hamilton-Buchanan, 1822). While the two species 
are morphologically extremely similar and undoubtedly closely 
related, I agree with Bleeker and with Weber and de Beaufort 
(1913:3.^8) that they are distinct. The most obvious difference 
involves coloration. Bomean and apparently Malaysian speci- 
mens are uniformly dark brownish in life, without distinct marks 
on body. Specimens from Nepal are greyish in life, with a more 
or less well defined midlateral longitudinal stripe, sometimes 
also a less well defined or more diffuse longitudinal stripe ventral 
to the midlateral stripe, and a dark humeral spot emphasized 
by a white or pale area along its ventral margin. The longitudinal 
stripes tend to fade but not the humeral spot even in very old 
preserved specimens. Such coloration, longitudinal stripes and 
a humeral spot outlined by pale or white areas, is characteristic 
of several Indian species but does not occur in any Malaysian 
or Indonesian species. Most of these Indian species have no 
obvious close relatives in Malaysia and Indonesia, but A/, ca- 
vasiiis constitutes an exception. 

Mystus nigriceps and M. cavasiiis generally can also be dis- 
tinguished by gill raker counts (Table 8). M. nigriceps from the 



Malay Peninsula and Kapuas have 22-25 rakers while most Af. 
cavasiiis examined from Pakistan, Nepal, India, Sri Lanka, and 
Thailand have only 15-21. This generally clear-cut difference 
in gill raker counts is complicated however by samples from 
Bombay and Burma. The Bombay specimens (CAS-SU 35854, 
8: 54.3-89.7 mm, Deolali, Bombay Presidency) have 20-24 
rakers. These specimens, although somewhat faded, exhibit the 
typical coloration of A/, cavasiiis and do not seem to differ except 
in gill raker counts from other samples of that species. The 
Burmese specimens, on the other hand, are rather distinctive 
and may possibly represent a species distinct from either A/. 
cavasiiis or A/, nigriceps. 

The bagrid Heterobagrus bocoiirti. characterized by an en- 
larged dorsal fin with an extremely elongate non-serrate spine, 
has been recognized as a valid genus and species since its original 
description by Bleeker ( 1 864), Bleeker made the following state- 
ment about the relationships of Heterobagrus: "Ce type nouveau 
tient le milieu entre les genres Hypselobagrus [type-species Mys- 
tus nigriceps]. Psciidobagriclithys et Bagrichtbys. Si Ton voyait 
que la tete, on n'hesiterait pas de le rapporter au Hypselobagrus. 
Si, au contraire. Ton ne tcnait compte du dos et des nageoires 
dorsales, on ne pourrait manquer d"y voir un Bagrichihys [type 
and only included species B. hypseloptenis]. Et si Ton n'en ob- 
servait que la moitie posterieure du corps, tous les Silurologistes 
y verraient un P.u'udobagriclithys." Resemblance between Het- 
erobagrus bocourii and Bagrulilhys (including Pseiidobag- 
richtbys) is in my opinion entirely superficial. A host of spe- 



124 



CALIFORNIA ACADEMY OF SCIENCES 



Table 8. Mrsrus fJiuRictps and Closely Related Species. Frequencies of gill rakers and vertebrae. 



Cjill rakers 



Vertebrae 



16 



18 



\'i 



?0 



34 



43 



44 



.Mvsliis hocoiirli 
Pasak R., CAS-SU :88:6 

Mwsiui Ciivas/us 
Deolah, CAS-SU 3485 
Bisranipur, CAS-SU 41068 
To/pur, CAS-SU 41065 
Calcutta. CAS-SU 34859 
Hugh R, CAS-SU 41069 
Naraini R., CAS 50275 
Rapti R.. CAS 50312 
Dudara R . CAS 50327 
Indus R.. CAS 24246 
Pulla. CAS-SU 34850 
"lakvala, CAS-SU 30556 
Nikaventiya, CAS-SU 30150 
Bangkok. CAS 53213 

MvMiis near cavasnis #1 
HIegu. CAS-SU 34851 
Minhla Tank, CAS 53339 
Rangoon. CAS-SU 33788 

Mvsliis near cavasiiis #2 
Rangoon, CAS-SU 

Pahang, CAS-SU 30998 
Telok Anson, CAS-SU 30994 
Pcrak, Sauk, CAS-SU 32720 
Perak, Chandra, CAS-SU 30992 
Malacca. CAS-SU 30995 
Ayer Hitam, CAS-SU 30993 
Kota Tinggi, CAS-SU 39341 
Kapuas, CAS 49381 
Buitenzorg, CAS-SU 20492 



1 I 

I 

1 

12 4 

I 



4 1 



cialized characters oCBagnchiliys including several modifications 
of the axial skeleton, well developed retrorse dorsal-fin spine 
serrae, convoluted and papillose barbels, and Lctocassis-Uke 
coloration are absent in Hcterohai^iiis. On the other hand, direct 
comparison of specimens of Hclcriihagrus and Mystus lugnceps 
reveal ihey are extremely similar in virtually every respect ex- 
cept size of the dorsal fin and its elongate spine. I consider 
llctcwhagrus bocourti a valid species congeneric with Mystus 
(or Hypselohagnis) nigrlceps. For merislic data on ,\f. hocoiirli 
see Table 8. The species occurs in the Chao Phrya and Mekong. 

Distribution. — .^/, nignceps is very widely distributed in the 
Malay Peninsula, Sumatra. Borneo, Java; it may also occur in 
Thailand, hut the few Thai specimens examined by me seem 
to be M. CLivasms. 

Popta (1906) described five new species and a new variety of 
Macroncs (=Afysiiis) from the Kapuas and Mahakam basins. 
Weber and de Beaufort (1913:34) examined the type specimens 
of these taxa and placed them all in the synonymy of M. ne- 
niiinis, but the type material should be more carefully re-ex- 
amined and compared with specimens of other species. I ten- 
tatively place only M. bongan as a synonym of M. nemiirm. 
Popta's other new species, M. howong. M. bo, and M. kajan. 
all from the Mahakam basin, may be synonyms of M. bara- 
))k7;i7i(Regan 1 906). descnbed from the Baram River, Sarawak. 
Mysius baranwiisis resembles M iicnninis but is less elongate. 



with a much shorter snout and a slightly shorter or deeper caudal 
peduncle, head much broader and covered with smooth skin 
(rather than relatively exposed and rugose dorsally, at least in 
larger specimens). 

Mystus olyroides new species 

(Figures 95. 96) 

HoimvpE. — MZB 3625. 155 mm female with ripening o\ aries. Sintang Market, 
liil\ 1976 (Kapuas 1976-19), reportedly caughl in Sungai Kebian (see Kapuas 
197h-20). 

PxRATYPE. — CAS 49382, 202 mm mature male with well de\eloped testes and 
moderately elongate genual papilla, obiained wilh holol\pe. 

Diagnosis. — A very elongate Mystus with a short flat head; 
small eyes; very long barbels; very short and weak dorsal spine; 
very long adipose fin; caudal fin with upper lobe elongate and 
lanceolate, lower lobe rounded and much shorter; gill mem- 
branes broadly overlapping, branchiostegal rays 9-11; gill rakers 
14; pectoral rays 8; anal rays 13; vertebrae 24 + 24-25=48 or 
49; body and fins without markings; color in life uniformly dark 
brown dorsally and laterally, abdomen dusky. 

The holotype and single paratype (in parentheses) oi Mystus 
olyroides may be further characterized as follows: head length 
3.7 (3.9). dorsal surface covered with smooth skin; snout round- 
ed, length 10.6 (10.9); eye small but entirely free from orbital 
rim. horizontal diameter 35.2 (40.4); interorbital space broad 



ROBERTS-WESTERN BORNEO FISHES 



125 




Figure MS. A/ysliis olyroKks- Kapuas 1 976- 1 '^. 155 mm (MZB 3625. hololype). 



and flat, width 13.6 (13.7); mouth broad, width 8.9 (8.9); upper 
and lower jaw teeth and palatal teeth in slender bands of rela- 
tively generalized conformation for Myslus. except lateral arms 
of palatal tooth patch extend relatively far posteriorly (Fig. 96); 
gill rakers elongate, 3+1 + 10 (2+ I + 1 1) on first gill arch, all 
well developed; an unusually long portion of upper limb near 
origin and of lower limb near insertion without gill rakers; pos- 
terior margms of first and second gill arches without gill rakers 
and of third and fourth with gill rakers; barbels very elongate 
and thin; maxillary barbel extremely elongate, extending pos- 
teriorly to base of caudal fin (or to end of anal fin base), its 
length 1.0 (1.2); nasal barbel, thinner than other barbels, ex- 
tending posteriorly almost to occipital crest (well past eye), its 
length 6.2 (9.2); outer mental barbel, slightly longer than inner, 
extending posteriorly nearly to end of pectoral fin, its length 2.7 
(3.1); body depth 6.0 (6.5), width 5.9 (6.0); dorsal spine weak, 
almost flexible, and very short, length 1 2.0 ( 1 2.6); occipital pro- 
cess, overlain by thick smooth skin, apparently very short, its 
length about equal to eye diameter; predorsal plate greatly re- 
duced and deeply embedded; distance between occipital process 
and dorsal-fin origin about 8.5 (9.3); pectoral spine short, length 
8.4, with 12 well developed moderately large serrae on posterior 
edge (pectoral spines broken off near base on both sides of 
paratype); humeral spine short, length 16.5 (15.2); pelvic fin 
with 6 rays and large, length 6.25 (6.5) but with a relatively 
narrow base, width at base 36.9 (35.5); adipose fin extremely 
long, originating a distance about equal to eye diameter behind 
base of last dorsal-fin ray and extending posteriorly dorsal to 
caudal peduncle and onto anterior half of ridge created by upper 
procurrent caudal-fin rays, its length 2.2 (2.1) and maximum 
height 14.9 (17.7); caudal peduncle very elongate, length 4.7 
(4.9) and maximum height excluding adipose fin 12.3 (13.6); 




Figure 'J6. Myslus iilyroidcs, oral toolh bands. Rapuas 1976-19, 202 mm 
(CAS 49382. paratype). 



126 



CALIFORNIA ACADEMY OF SCIENCES 



caudal fin with 9 + 8/9+ 13-14 rays; upper lobe length 2.7 (2.4); 
posterior swim bladder chamber of 202 mm paratype dorso- 
ventrally flattened and oval, its narrowed posterior end well 
before level of pelvic origin; posterior swim bladder chamber 
with rigid walls; internally a ngid dorsoventrally and longitu- 
dinally complete median septum divides it into separate right 
and left chambers, floors of which are further complicated by a 
series of skinlike partial septae; head (including barbels) and 
body (except ventral surface) dark brown; adipose fin and fin 
rays of all fins brown; interradial membranes of all fins except 
pectoral varying from dusky to almost black; interradial mem- 
branes of pectoral fin clear. 

Remarks. — Afystiis olyraides docs not seem particularly close 
to any other species. In some respects it resembles "Lciocassis" 
mahakaniensis but differs considerably from the holotype and 
only known specimen of that species (type locality Mahakam) 
in coloration, more elongate adipose fin. and rounded (rather 
than lanceolate) lower caudal-fin lobe. It also superficially re- 
sembles and may be related to the genus Olyra of India, Burma, 
and western Thailand, which is usually assigned to a separate 
family, Olyridae. but apparently is a bagrid. 

Distribution. — Kapuas. 

Etymology.— The name olyroidcs refers to the similarity of 
this species to members of the Asian catfish genus Olyra 
McClelland. 1842. 

Mystus wolffii (Bleeker. 1851) 

Bairns W'olthi Bleeker. 1851/'. 205 (type locality liandjcrmassing, in fluvus). 
Hypviiihagnn li'oltti Bleeker, 1862-63:58 (Bandjermassing, Ponlianak, Palem- 

bang). 
Macronci wnltfii Giimlicr, I8h4 82, 

Material Examined —Peninsular Thailand: lirackish lagoon near Songkhla, 7: 
inS-l.^h mm (CAS 55554) Borneo: no lurther loeality, .1 77.S-I 15 mm (RMNH 
2445) Bleeker collection: locality data lost. « 47 8-128 mm (RMNH 6866). 

The only report of this species from western Borneo appar- 
ently is that of Bleeker (1858f: 161, 1860f/:19)from Pontianak. 
The material upon which this record is based cannot be located, 
and neither can the holotype. Bleeker ( 1858i.';16l. 1862-63) 
indicated havmg 16 specimens of .U walffii. The auction catalog 
of Bleeker's collections also indicates 16 specimens, 12 in lot A 
and one each in lots B-E. Lot A. which should have included 
the holotype, is represented by RMNH 6866 which now has 
only nine specimens. None of them has a total length of 131 
mm as reported for the holotype. 

The Bleeker specimens of M. woljfii (RMNH 6866) have gill 
rakers 6-9+18=247(1), 25(2). 26(2), 28(2), and 29(2) and ver- 
tebrae 18-19+18-217=37(2), 38(6), and 39'^(1). The RMNH 
Borneo specimens agree closely in all respects observed with the 
Bleeker matenal. They have gill rakers 7-9 + 21-23=28( I), 29( 1), 
and 32(1) and vertebrae 18 + 20=38. The CAS specimens from 
Thailand have been compared directly with the Bleeker and 
Borneo specimens and agree closely except in having more nu- 
merous gill rakers. The Thai specimens have gill rakers 9-1 1 +29- 
30=39(2), 40(3), and 42(2) and vertebrae 18-19+19-20=37(3) 
and 38(4). 

DisTRiBL'TiON. —Thailand: lower courses of ri\ ers flowing into 
Gulf of Thailand. Sumatra (Palembang). Borneo (Pontianak, 
Bandjermasin). I suspect that this species is restricted to the 
lower courses of rivers, as indicated by Smith (1945;384). and 



that it may be a brackish water species. The specimens from 
Thailand were taken with a large number of specimens tenta- 
tively identified as Mysliis gidio (CAS 55555). Smith indicates 
a published record from Malacca, on the Indian Ocean coast of 
the Malay Peninsula, but I have not found the source of this 
record. A record from the upper Mahakam by Popta ( 1 906:253) 
is evidently based on a misidentified specimen examined by me 
and tentatively re-identified as M. ncmwiis (RMNH 7835). 

Mystus wyckii (Bleeker, 1858). 

Biiiinis W'ycku Bleeker. 1858i:156 (type locality Java, in tlumine Tjiiarum. pro- 

\ inciae Preanger, propc vicum Parongkalong), 
llciuihai;rm llVci// Bleeker, 1862-63:57, 
Macroncs wycku Giinlhcr, 1864:83. 
Mvsim H'lc*.'/ Smith, 1429:12, 

Material Examined —Western Borneo: Kapuas 1976-29. 4: 35,2-206 mm 
(C^S 49383. MZB 3626): Kapuas 1 976-45. 4: 3 1 ,7- 1 5 1 mm (MZB 3627, USNM 
230284); Kapuas 1976-48. 2: 18,6-20,2 mm (CAS 49384, MZB 3628); Kapuas 
1976-50, 2: 27,5-29 7 mm (CAS 49385, MZB 3629), 

Diagnosis. — A large species of Mystus (Hemibagnts) with 
distinctive coloration and relatively numerous vertebrae. Dorsal 
and lateral surfaces of head and body and all fins largely black 
in color but with well defined white markings on tips of pectoral 
and dorsal fin; leading edge of adipose and anal fins, upper and 
lower margins of caudal peduncle, and caudal fin (including 
filamentous extensions of unbranched primary caudal-fin rays); 
gill cover black but gill membranes pale, almost white; nasal 
barbel dark and mental barbels pale, but elongate maxillary 
barbel (extending posteriorly beyond level of pelvic-fin origin) 
with anterior margin white and posterior margin black; bran- 
chiostcgal rays 9 + 8; gill rakers on first gill arch 3-4 + 9-10=12- 
14, anterior portion of lower limb rakerless near origin; verte- 
brae 51. 

Pelteobagrus Bleeker, 1865 

Pi-llcobaarus Bleeker. 1865k:9 (type species Siliinis Liilviiniis Basilewski, 
l855 = F(»k'/o(/(« fiilvidracn Richardson, 1845''), 

Pelteobagrus ornatus (Duncker, 1904) 

(Figure 97) 

.l/;i(s sp, indet, Vaillant, 1902:71 (Le Kapoeas, ''Sintang), 

l\ciidiihagnis onialiis Duncker. 1904:173 (type locality Muar-Fluss bei Tubing 

tinggi) 
rehciibLtfinii onuilus Jayaram, 1968:297. 

Material Examined — Malay Peninsula: Muar River at Tubing tinggi. 2: 24,0- 
25 5 mm (HZM S654, svntypes of P onuilus]. Western Borneo: Kapuas basin, 
Sintang, 29,1 mm (RMNH 7826, previously identified as liius sp, indet,). 

Placement of this very small southeast Asian catfish with large 
e>es in cither Psctidohagiiis or I'clicobagnis. otherwise exclu- 
sively east Asian (China, Korea, Japan, Siberia) genera of rel- 
ati\el> large species with small eyes, is highly tentative. 

The species was known only from Duncker's two syntypes 
from Muar until Kottelat (1982:434) reported two specimens 
from southeastern Borneo. A single specimen from the Kapuas 
basin, unfortunately in rather poor condition, also appears to 
be this species. This specimen has not been compared directly 
to the syntypes but is apparently very similar to them in most 
respects: snout noticeably overhanging; horizontal diameter of 
eye fully 3 times in head length (eye relatively smaller in syn- 



ROBERTS-WESTERN BORNEO FISHES 



127 



types); tooth band of upper jaw nearly straight transversely, 
palatal tooth band about half as wide and moderately curved; 
nasal barbel absent? (minute nasal barbel present in syntypes); 
maxillary barbel extending to pectoral-fin origin (shorter than 
head in syntypes); mental barbels elongate; gill rakers 4+10; 
dorsal-fin branched rays 6; pectoral-fin spine with 8 strong ser- 
rae; pectoral branched rays 7; total anal-fin rays 19 or 20 (22- 
23 in syntypes according to Jayaram 1968:297). 

The extraordinary color pattern of the Kapuas specimen, 
identical or very similar to that of Duncker's syntypes (Fig. 97) 
in all respects, includes a horizontal stripe about half as wide 
as eye arising on side of head near snout-tip, extending laterally 
at mid-eye level to cover both nostrils, middle portion of gill 
cover, and part of cleithrum; a second stripe, about half as wide, 
arises near snout-tip but more dorsally and extends between 
eyes and onto occipital region; single roundish black spots at 
dorsal-fin origin, near middle of adipose-fin base, on abdomen 
behind each pelvic-fin base, and on anal-fin base near origins 
of anal-fin rays 4-7; caudal peduncle with a narrow midlateral 
horizontal stripe extending to middle of caudal-fin base; upper 
and lower halves of caudal-fin base with vertically oriented el- 
liptical or oval dark marks. 

Distribution. — Malay Peninsula (Muar). Borneo (Kapuas; 
Barito basin?). 

Clariidae 

This distinctive air-breathing family of Africa and Asia is 
readily distinguished by its elongate, entirely spineless dorsal 
fin (sometimes extending entire length of body), elongate anal 
fin. and distinctive overall appearance. Head flattened; four pairs 
of well developed barbels; eyes small; body cylindrical, mod- 
erately to extremely elongate; arborescent air-breathing appa- 
ratus arising from gill arches 2 and 4; cleithrum with knoblike 
bony process projecting into gill opening; caudal fin rounded. 

Clarias Scopoli, 1777 

Clarias Gronovius. 1763:100 (non-binomial, not available for zoological nomen- 
clature; Opinion 261. Official index of rejected and invalid works on zoological 
nomenclature. 1938). 

Chlanas Scopoli. 1777:455 (misspelling of C/dniu Gronovius, 1763; type species 
Silurus angiiillaris Linnaeus. 1758, by present designation). 

Clarias Cuvier. 1816:206 (type species Si/ttru\ ungutllaris Linnaeus. 1758. by 
present designation )- 

NoMENCLATURAL NoTES. — Although portions of Scopoli are 
binomial, the treatment oi Chlanas [=Clanas\ is non-binomial 
and does not include any reference to a binomial taxon. From 
the standpoint of the genus Clarias it is unfortunate that Gro- 
novius ( 1 763) is unavailable for zoological nomenclature, since 
the binomial Silurus anguillans is among the species listed under 
the genus and would therefore automatically be the type species 
by monotypy. The first authority to assign binomial taxa to 
Clarias is Cuvier (1816), who mentioned two species, Silurus 
anguillans Linnaeus, 1758 and Silurus halrachus Bloch, 1794 
[=Silurus halrachus Linnaeus, 1758]. Thus the generic type 
species must be one of these two species. In the apparent absence 
of subsequent designation of either Silurus anguillans or Silurus 
halrachus as type species of Clarias. I hereby designate Silurus 
anguillaris Linnaeus, 1 758 as type species o( Chlanas [=Clarias'\ 
Scopoli, 1777 and of C/anas Cuvier, 1816. 




Figure 97. Pelleohagrus ornalus. Malay Peninsula. Muar River (HZM 8654, 
syntype; after Duncker 1904, pi. 2, fig. 13). 



There is doubt as to the number of Clarias species in western 
Borneo and their identity. The survey of 1976 obtained only 
two species, but at least four and perhaps five species are present 
in the Kapuas. Weber and de Beaufort (1913) reported five 
species but their work is largely compiled and I have experienced 
difficulty identifying specimens utilizing their key, especially in 
distinguishing between C. leiacanlhus and C leysmanni. 

Clarias cf batrachus (Linnaeus, 1758) 

'.Silurus halrachus Linnaeus. 1758:305 (t>pe locality Asia). 

''Macropteronolus inagiir Hamilton-Buchanan. 1822:146 (type locality Ganges). 

Clanas halrachus Bleeker. 1858t:343 (Sambas), 

Clarias magur Vaillant, 1893:37 (listed from western Borneo). 

Material Examined.— Western Borneo: Kapuasbasin, Smitau, 212 mni(RMNH 
7799); Kapuas basin. Sintang, 2: 159-197 mm (RMNH 7800). 

The species tentatively identified as Clarias halrachus is one 
of the most important food fishes in southeast Asia (especially 
in Thailand, where it is extensively cultured). 

According to Weber and de Beaufort (1913:1 90) C. halrachus 
differs from other species of Clarias found in Borneo and In- 
donesia in having generally fewer dorsal- and anal-fin rays (60- 
76 vs. 68-106 dorsal and 47-58 vs. 52-95 anal). 

Clarias leiacanthus Bleeker, 1851 

Clarias leiacanlhus Bleeker. 1851/:430 (type locality Sambas, in lluviis). 
Clarias liacanlhus Giinther. 1864:20 (unwarranted spelling change). 

Material Examined. — Western Borneo: Kapuas 1976-6.4: 80.2-135 mm (CAS 
49422. MZB 3704. USNM 230308); Kapuas 1976-17. 131 mm (MZB 3705); 
Kapuas 1976-30. 96.8 mm (MZB 3706); Kapuas 1976-31, 14.7 mm (MZB 3707). 

Clarias meladerma Bleeker, 1 846 

Clarias nieladenna Bleeker. 1846i(:178 (type locality Batavia. in paludibus). 
Clarias melasoma Bleeker. 1852/):427. 
Clarias nwlantidcrma Bleeker. 1858i.:341. 
Clarias mclanosuma Bleeker. 1858-59ii:l. 

Material Examined. — Western Borneo: Kapuas 1976-51. 5: 88.6-242 mm 
(CAS 49423. MZB 3708, USNM 230309). 

NoMENCLATURAL NoTE. — Although virtually all authors sub- 
sequent to Bleeker (1858) have referred to this species as C 
melanoderma, the original spelling of C meladerma'\%Qmp\oyed 
here. The spelling C meladerma appears twice in Bleeker's orig- 
inal account (Bleeker 1846a: 178, 188) thus indicating it was 
intentional and not a typographical error. 

Clarias meladerma is distinguished by a series of strong serrae 
projecting from leading edge of pectoral-fin spine. In all other 



128 



CALIFORNIA ACADEMY OF SCIENCES 



Chinas the leading edge is smooth, rough, or with a few rela- 
tively weak serrae. 

Clarias nieuhofi Valenciennes, 1840 

C'laruis nicuhdti Valenciennes in Cuvier and Valenciennes. 1 840;3S6 (type locality 

not indicated). 
Clarias peniaplcnis Bleeker, l85l/;;206 (type locality Bandiermassing). 
''Clai-ias lagtir von Martens, 1876:302. 39>i (not ot Hamilton-Buchanan. 1822; 

Danau Snang. Kapuas basin). See Weber and de Beaufort (I913;l<*4). 
Clanas Nieuhofi Vaillant. 1893:63 (Kapuas basin. Knapei et Sebroeang). 

Material Examined. — Malay Peninsula: Tinggi. .lohore, 14(3 mm (CAS-SU 
32694). Western Borneo: Kapuas basin. Raouen R., Mandai. 2: 171-172 mm 
(RMNH 7685): Kapuas basin, upper Sibau R., 3: 204-230 mm (RMNH 7684); 
Kapuas basin. Sintang. |91 mm (RMNH 7805). 

Diagnosis. — r/am;.'; ;)/«//((.)/( differs from all other southeast 
.\sian Clarias in having more numerous dorsal- and anal-fin 
rays (87-106 vs. 60-76 dorsal and 69-95 vs. 47-53 anal) and 
dorsal and anal fins united to caudal fin at least at its base (Weber 
and de Beaufort 1^13). 

Clarias teysmanni (Bleeker, 1857) 

Clarias Tciismanni Bleeker. 1857/):344 (type locality ■Tjikoppo. pro\inciae Bui- 

ten/org. 900 metr. supra mare." .lava). (Typographical error lor 'Ifyuiiuiini ) 
Clarias Teysmanni Bleeker, 1862-63:104. 
Clarias Tevsmanni Vaillant. 1893:62 (Kapuas). 

Both MNHN and RMNH have a number of lots identified 
as C tcysmannt from the Kapuas basin. I briefly examined the 
RMNH material but was unable to come to any firm conclusion 
as to the correctness of its identification or the distinctness of 
C. icysmanni from C leiacantlnis. Two characters utilized by 
Bleeker and Weber and de Beaufort to distinguish the species 
seem highly variable and difficult to use: shape of occipital pro- 
cess and distance between occipital process and dorsal-fin base. 
The latter can differ by 20 percent or more in the same specimen 
depending on whether the head is flexed dorsally or ventrally. 

It may be noted in passing that substitution of "ij" for "y" 
is a common typographical error throughout Bleeker's publi- 
cations. Thus Cypnnidae is often printed 'Typrinidae." 

Encheloclarias Herre and Myers, 1937 

F.nchchclarias Herre and Myers. 1937:66 (type species llelcrnhraiuhin lapeinnp- 
icriis Bleeker. 1852. by onginal designation and monotypy) 

Encheloclarias. the only Asian clariid genus with an adipose 
fin, differs from African clariids with an adipose fin (Hctero- 
braiiclnis. Dinolopicrus. some species of Clanas) in head and 
body shape, relationship of adipose and anal fins to caudal fin, 
morphology of neural spines, vertebral formula, and size. Over- 
all form, when viewed from above, not notably tapered from 
head to tail; head very small, relatively short and narrow, only 
slightly wider than anterior part of body (in African forms with 
adipose fin overall form strongly tapered; head very large, much 
longer and broader). Adipose and anal fins broadly united to 
caudal fin (both fins always or almost always separate from 
caudal fin in African forms). Vertebrae about 16 + 41=57, those 
immediately behind Weberian apparatus relatively elongate 
{Heterohranchits with vertebrae about 23-25 + 34-36=57-61, 
those immediately behind Webenan apparatus strongly com- 



pressed) (vertebrae of Weberian apparatus difficult to count in 
radiographs). Largest known specimen 124 mm total length 
{Hetcrohranchus and DincHopicrus commonly reach 1 m and 
sometimes nearly 2 m). 

Although Herre and Myers (1937) stated that the adipose fin 
of Encheloclarias is supported by "extremely weak extensions 
of the neural spines," this is in error. I have examined the same 
specimen they did, and a radiograph clearly reveals that the 
neural spines are confined to the body proper (i.e., do not enter 
the adipose fin at all). Examination of the adipose fin with trans- 
mitted light reveals fine ray-like structures (ceratotrichia?) about 
4-5 times more numerous than neural spines. For further dis- 
cussion and illustrations of the elongate neural spines in the 
adipose fin of African clariids see Teugels (1983). Detailed com- 
parison of Encheloclarias and Heterobranchus probably will re- 
veal additional distinctions. The size, shape, and orientation of 
some cranial bones apparently differ. The anterior tubular nos- 
tril of Encheloclarias has not been studied. Only a single species 
is recognized. 



Encheloclarias tapeinopterus (Bleeker, 



;152) 



HclerohraiK-liii', lapcinitplcrin Bleeker. I852n:732 (type locality Banka); Bleeker, 

I S58<:337 (Sambas); Weber and de Beaufort, 1913:194. 
luhiicloclarias lapcinoplcriis Herre and Myers. 1937:66 (Mawai, Johore). 

Material Examined —Malay Peninsula: Mavvai. Johore. 63.3 mm (CAS-SU 
32004) 

Distribution. — Malay Peninsula (.lohore). Western Borneo 
(Sambas). Report of this species from Sarawak by Weber and 
de Beaufort (1913:195) (copied by Herre and Myers 1937:66) 
apparently is based on Bleeker's record of it from Sambas, which 
IS in western Borneo, not Sarawak. 

Schilbeidae 

Schilbeids are smooth-skinned catfishes with two to four pairs 
of usually elongate barbels (vestigial or absent in Silonia); large 
laterally or ventrolaterally directed eyes; large gill openings with 
branchiostcgal membranes separate from isthmus; gill rakers 
moderately to very numerous; swim bladder with bilaterally 
expanded anterior chamber frequently visible through large thin- 
walled tympanic membrane; posterior swim bladder chamber 
occasionally extending posteriorly above anal-fin base; dorsal 
fin small, with or without a spine and up to 7 branched rays, 
or absent; adipose fin small and h\aline or absent; pectoral fin 
with a thin stiff spine usually serrate on posterior margin; anal 
fin long, not confluent with caudal; anal-fin rays 24-90; caudal 
fin deeply forked, lobes usually pointed, usually (always?) with 
8 9 principal rays. Usually whitish or pearly, often with longi- 
tudinal dark stripes, a few species mottled brownish. 

The family comprises about 16 genera and 40 species equally 
divided between Africa and Asia. Most of Asian taxa restricted 
to Indian subcontinent, A single genus, Pseudeuiropius. in In- 
donesia. (Laiiles. sometimes placed in Schilbeidae, is arbitrarily 
referred to Pangasiidae in the absence of compelling evidence 
concerning its familial placement.) 

Pseudeutropius Bleeker, 1862 

PHnulctilrnpiiis Bleeker. 1 862398 (type species Eiiiropiiis hraclivpdpleriis Bleeker, 
18-^8. b\ original designation and monotypy). 



ROBERTS-WESTERN BORNEO FISHES 



129 




Figure 98. Pseudeutmpius. Above, P. hrafhvpopterus. Kapuas 1976-32, 73.8 mm (MZB 3683K below, R inoolenburghae. Kapuas 1976-32. 67.2 mm (MZB 3686). 



Pseudeutropius brachypopterus (Bleeker, 1858) 

(Figure 98 upper) 

Eutiopius brachypoplenis Bleeker. 1858i;169 (type locality Sumatra, Palembang. 

in fluviis). 
Pscudeulropius bnichypoplerus Bleeker. 1863^:107. 

Material Examined. — Western Borneo; Kapuas (1976-20. 8: 73.2-87.8 mm 
(CAS 49410, MZB 3682), Kapuas 1976-32, 46: 47.8-92.3 mm (BMNH 
1982.3.29.178-182, CAS 53290, FMNH 94244, MNHN 1982-711, MZB 3683, 
RMNH 28904, ROM 38617. UMMZ 209882, USNM 230301); Kapuas 1976- 
33, 8; 75,0-102 mm (CAS 49411, MZB 3684); Kapuas 1976-44, 3: 56,4-84.4 
mm (MZB 3685, ZMA 1 16,542), 

Diagnosis. — Head relatively long, 3.4-3.9; barbels relatively 
short, extending posteriorly about to level of dorsal-fin origin; 
gill rakers extremely long and numerous (22 + 71=93 in a 102 



mm specimen); anal-fin rays 31-35; vertebrae 12- 
26=37(5) or 38(8). 



13 + 24- 



Pseudeutropius moolenburghae Weber and de Beaufort, 1913 

(Figure 98 lower) 

Pseudeutropius iiioolcnhurghae Weber and de Beaufort, 1913:249, lig. 100 (type 
locality Batang Hari, Sumatra). 

Material Examined— Western Borneo: Kapuas 196-32, 95; 49.7-69.8 mm 
(AMNH 48939. BMNH 1982.3.29.183-187. CAS 49412. IRSNB 19748, MCZ 
58353, MNHN 1982-712, MZB 3686, RMNH 28905. ROM 38622, UMMZ 
209883, USNM 230302, ZMA 1 16,543); Kapuas 1976-33, 3; 50.8-80,3 mm (CAS 
49413, MZB 3687); Kapuas 1976-44, 2; 55,3-67.0 mm (MZB 3688, USNM 
230303); Kapuas 1976-46. 68.5 mm (MZB 3689). 

Diagnosis. — Head relatively short, 4.4-4.6; barbels relatively 
long, extending posteriorly to a vertical behind anterior fourth 



130 



CALIFORNIA ACADEMY OF SCIENCES 




(;\ 






/! 


\\^ 






\^v 






y / 


\ -■ 


. 


,^ 


■^ / 


\ 


^/"' 




y 


\^ 


^J_ 




/ 














10 mm 



Figure '^'J. Oral dentition ofSundaland Pangasnis. Species wuh complex palatal loothbands. a, P. dezwaaiu Weber and dc Beaufort, 1912, Sumatra, Taluk (ZMA, 
holotype); b, P nasiitm. Kapuas 1976-19, 215 mm (CAS 494071; c, /'. polywanodon, Kapuas 1976-19, 227 mm (CAS 49408), d, P. ponderosus Herre and Myers, 
1937, Malay Peninsula, Chanderoh dam. Perak, 600 mm (CAS-SU 14162, holotype) 



or third of anal-fin base; gill rakers very long, 14-1-29-31=43- 
45; anal-fin rays 42-49; vertebrae 10-11+ 30-32=4 l(5)or 42(1 2). 

Pangasiidae 

Pangasiids are elongate, moderately compressed, smooth- 
skinned midwater catfishes of large size; mouth subterminal, 
moderately to very wide; maxillary barbels and a smgle pair of 
mental barbels usually present, nasal barbels and second pair 
of mental barbels invariably absent; dorsal- and pectoral-fin 
spines well developed, with serrate internal margins; dorsal fin 
with 7 soft rays, situated anteriorly; adipose fin well developed 
but with a restricted base; anal fin elongate, with 26^2 rays; 
caudal fin deeply forked, principal caudal rays 8 + 9; posterior 
chamber of swim bladder large, flattened and shield-shaped, 
with rigid walls supported by numerous internal fibrous struts; 
vertebrae 41-52 (Pangasius). 

Many catfishes have paired pectoral or axial glands lying in 
the body wall near the pectoral-fin base and sometimes partially 
overlain by the humeral (or postcleithral) process. This gland is 
one of the specializations of siluroids that has no known homo- 
log in other fish groups. In most catfishes each gland opens to 



the exterior by a single axillary pore or poms pcctoralis lying 
just below the humeral process and above the origins of the 
innermost pectoral-fin rays. In Pangasius the gland is sometimes 
unusually large; it tends to be as long as the humeral process. 
In addition to a large poms pectoralis in the usual location, some 
Pangasius have a large number of additional pores more or less 
coextensive with the portion of body wall overlying the entire 
pectoral gland. Another peculiarity of the pori peclorali of Pan- 
gasius is that the openings are not simple, as in other catfishes, 
but loculose (i.e., divided into sections by internal partitions). 
In many catfish families, especially in South America, the se- 
cretion of the pectoral gland is milk-white or creamy and at 
least in some tastes like milk or cream. In Pangasius the secre- 
tion is clear and viscous, and tastes like mucus. In southeast 
Asia as in South America it is believed locally that the pectoral 
gland provides nourishment for the young. It is remarkable that 
so little is known about these glands. 

The family comprises five or six genera and about 30 species, 
the great majority of which belong to the genus Pangasius. It 
ranges from the Indian subcontinent (including the Indus basin 
but not Sri Lanka), through mainland southeast Asia (including 
the Mekong basin). Sumatra, and Borneo. Although Helicopha- 




FiGLiRE 100. Pangasius humcrahs Kapuas 1976-19, 389 mm (RMNH 28903, paratypc). 



ROBERTS- WESTERN BORNEO FISHES 



131 




*.,•■••■•■• '^■^■^ 




10 mm 



20 mm 




10 mm 



Figure 101. Oral dentition of Sundaland Pangasius. Species with simple palatal loothband. a, P lithostoina. Kapuas 1976-19, 214 mm (CAS 49406, paratype); 
b, P. meuwenhuisi. Mahakam, nvifre Bo, 400 mm (RMNH 7546, holotype); c, P humcratis. Kapuas 1976-19, 215 mm (CAS 49408, paratype); d, />, sp. undet., 
northeastern Borneo, Kinabatangan, 224 mm (FMNH 68041, formerly identified as P ntcuwcnhuisi). 



gus occurs in Thailand, Sumatra, and southeastern Borneo, only 
Laides and Pangasius are known in western Borneo. 

Laides Jordan, 1919 

Lais Bleeker, 1858c: 170 (type species Panf;asius hcxanciiiu Sleeker, 1852, by 

monotypy). Preoccupied by Lais Gistcl, 1848, Tunicata. 
Laides Jordan. 1919:293 (replacement name for Lais Bleeker. 1858). 

Laides differs from all other pangasiids in having two pairs 
of mental barbels instead of only a single pair. Maxillary barbel 
elongate, reaching posteriorly to middle of body. Adipose fin 
minute. Anal fin with 39-42 rays. Vertebrae 44. A single species. 

Laides hexanema (Bleeker, 1852) 

Pangasius hexancma Bleeker. l852(/:588 (type locality "Palembang. Balavia, in 

fluviis"). 
Lais hexanema Bleeker, 1 858t': 171. 

Pangasius longibarbis Fowler, l934/i:87 (type locality Mekong R, at Chieng Sen). 
Laides hexanema Smith, 1945:372. 

Material Examined. — Western Borneo: Kapuas 1976-9, 96.4 mm (MZB 3676); 
Kapuas 1976-14, 816 mm (MZB 3677), Thailand: Mekong R. at Chieng Sen. 2: 
51,6-95,0 mm (ANSP 59441. 59442, holotype and paratype of F, longibarbis). 

Distribution, — Thailand (Mekong). Malay Peninsula. Su- 
matra (Palembang, Lahat. Lematang Emm, Ringat, Sidjund- 
jung, Djambi). Borneo (Kapuas). 

Pangasius Valenciennes, 1840 

Pangasius Valenciennes in Cuvier and Valenciennes. 1840:45 (type species Pi- 

metodus pangasius Hamilton-Buchanan, 1822, by monotypy and absolute tau- 

lonymy). 
Pseudopangasius Bleeker, 1 862:399 (type species Pangasius polyuranodon Bleeker, 

1852, by original designation and monotypy), 
Pseudolais Vaillant, 1902:51 (type species Pseudolais lenancma VaiUant, 

\9Q2=Pangasius micronema Bleeker, 1847, by monotypy). See Weber and de 

Beaufort (1913:261). 
Neopangasius Popta, 1904:180 (type species Ncapungasius mciiwcnhuisii Popta, 

1904, by monotypy). 

Division oi Pangasius into three genera based on differences 
in palatal dentition (Pangasius with two vomerine and two pal- 
atine toothplates, Pseudopangasius with a single vomerine and 



two palatine toothplates, and Neopangasius w\lh a single median 
palatal toothplate) seems to result in artificial or non-phyletic 
assemblages of species and has been rejected by most authors. 
Nevertheless the toothplates are useful in distinguishing some 
species, and those of most of the Pangasius known from Sun- 
daland are illustrated in Figures 99 and 101. 



Pangasius humeralis new species 

(Figures 100. lOlc) 

HoLOTVPE. — MZB 3680. 173 mm. Kapuas basin, fish market at Sintang (Kapuas 
1976-19). 

Paratypes.-CAS 49408. 204 mm, RMNH 28903, 389 mm, same data as 
holotype. 



DiAGNOsis. — P. humeralis is distinguished from all other Pan- 
gasius in western Borneo by the very elongate humeral process 
which extends fully two-thirds to three-quarters of distance to 
end of pectoral-fin spine. Pectoral gland large, with numerous 
pectoral pores (14 or more) coextensive with area ventral to 
humeral process. Eye small. Palatal teeth in a single median 
patch. Gill rakers 7-^0-1 + 12-12=19-22. Anal-fin rays 30-31. 
Vertebrae 19-20 + 28=47-48. Pectoral fins jet black. Body with- 
out distinctive markings. 

P. humeralis may be further characterized as follows: snout 
broadly rounded, length 9.8-1 1.1; head length 4.2; exposed por- 
tion of eye 29-35; margin of lower jaw strongly curved, mouth 
width 10.2; body depth 4.3, width 4.7; predorsal length 2.6; 
dorsal and pectoral spines moderately strong; pectoral spine 
length 5.2; posterior pectoral serrae about 1 7, or about 29 count- 
ing rudimentary basal serrae; humeral process 6.7. 

Remarks. — The only other Pangasius known to me with a 
humeral process as long as that off. humeralis is P. meuwenhuisi 
Popta, 1904 from the Mahakam basin of eastern Borneo. This 
species also has palatal teeth consisting of only a median tooth- 
patch. 

Direct comparison of the 400 mm holotype off. nieuwen- 
huisi (RMNH 7546) with the 389 mm paratype of P. hu- 
meralis reveals that the snout shape is very different, that of P. 
meuwenhuisi being much more pointed; the angle subtended by 



132 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 102, I'cinvciMiis lilhoslmmi Rapuas l')7b-l'^, 1*^6 mm (MZB 3678. holotype). 



snout-tip and anterior border of eye is much greater in P. nicu- 
wcnhiiisii. In other respects these two species are rather similar. 

Distribution. — Known only from the Kapuas R. of western 
Borneo. In addition to the three specimens obtained at the mar- 
ket in Sintang, the species was observed in the Sungai Mandai 
(Kapuas 1^)76-38). 

Etymology.— The name luinierulis (Latin, shoulder) refers 
to the exceptionally large humeral (postcleithral) process of this 
species. 

Pangasius lithostoma new species 

(Figures lOla, 1112) 

HoLOTVPE.-MZB 3678. 1V6 mm, market at Sinlang (Kapuas 1476-1'*), 
Paratypes,-CAS 44406, RMNH 28W1, USNM 230249, 3: 181-214 mm, 
market at Sintang (Kapuas 1476-14) 

Diagnosis. — Distinguished from all other Pan,i;asius by a 
greatly expanded and strongly convex vomerine toothplate which 
projects strongly downwards from the roof of the mouth like a 
millstone. Anal-fin rays 40-41; vertebrae 2 H- 3 1 =52 (in three 
specimens). 

The holotype (and three paratypes) may be further described 
as follows: snout broadly rounded, length 10,7(10,0-1 1.1); head 
length 4.2 (4.2-4.3); exposed portion of eye 32.7 (26.6-30.6); 
mouth opening strongly curved, width 1 1.7 (1 1.2-11.9); body 
depth 34.6 (4.1-4.7). width 5.8 (5.7-5.9); predorsal length 2.6 
(2.5-2.6); caudal peduncle depth 12.1 (11.4-12.4); dorsal and 
pectoral spines moderately strong; dorsal spine length 5.8 (5.4- 
6.1); pectoral spine length 5.6 (5.4-6.1); pectoral serrae 16(17- 
19). Humeral process exposed, extending slightly less than half- 
way to end of pectoral spine, its length 1 2,0 ( 1 2, 1-1 2,8); pectoral 
pores conhned to area above bases of pectoral-hn rays; gill rakers 
6_8-(-14-19=22-26;finsdusky, body without distinctive marks. 

Distribution, — Known only from the Kapuas R, of western 
Borneo. 

Etymology.- The name lithosionia (Greek) refers to the 
stonelike palatal loothpatch of this species. 



Pangasius micronema Bleeker, 1847 

l'air^a\ni\ Diicnmciihi Bleeker, 1847:8 (type loeality Java). 

I'uni:ii\iii\ iios Bleeker, l851/;:205 (type locality Bandjermassmg, in lluviis). 

I'iiii\;usiiis iins Vaillant, 1843:71 (Knapei and Sebruang, Kapuas). 

Pscudolais iclrancma Vaillant, |402:.<i2 (type locality Tepoe. bords du Mahakam). 

See Weber and de Beaufort (1413:261). 
Pangasius nos Vaillant, 1402:24 (Kapuas), 
Pangasius mis Popta, 1406:28 (Kapuas) 
''Pauuasnis luvksi Hardenberg, 1448 (type localil> Kapuas). See below. 

Material Examined— Malay Peninsula: Chandra or Chandcroh dam, Perak 
River, 7: 1 17-263 mm (CA.S-SU 31031. 32725. 34430): Perak. 164 mm (CAS- 
SU 31032), 

Numerous large Pciiii;asitis, probably this species, were ob- 
served in the fish market at Sinlang during the Kapuas survey 
of 1976 but none was preserved. 

Diagnosis. — F. micronema is distinguished by very large eyes, 
greatly reduced maxillary and mental barbels, truncate snout, 
outline of lower jaw angular rather than rounded, and very short 
skin-covered humeral process. 

Four specimens from the Perak River (CAS-SU 32725) have 
the following: snout 10.7-12.3; head 4.5-4.9; exposed portion 
of eye 16.8-20.5; mouth width 10.6-12.5; body depth 4.5-4.9, 
width 5.5-5.9; predorsal length 3.0-3.2 (intermediate between 
P. polvwanodon. 3.3. and other Kapuas Pangasius. 2.5-2.6); 
depth of caudal peduncle 1 1.2-12.7; dorsal and pectoral spines 
slender (as in P. polyurauodon): dorsal spine 7.0 (intact only in 
one specimen); pectoral spine 5.3-5.8; pectoral serrae (including 
rudimentary anterior serrae) about 24-30; humeral process cov- 
ered with skin, extending posteriorly less than midway above 
pectoral spine length, its length 12.4-13.8. Pectoral pores about 
1-4, tightly clustered above origin of pectoral spine. 

The identity of P. hocksi pu/zled me until I saw specimens 
of P, micronema with very black pectoral fins. The 275 mm 
(total length) holotype and only known specimen of P. hoel<si 
was not illustrated and my efforts to locate it have not been 
successful. The original description may be quoted in its en- 
tirety; 



ROBERTS-WESTERN BORNEO FISHES 



133 



D.II.7; A.32; PT.I3; V6. 

Elongate, rather strongly compressed, dorsal profile nearly straight 
behind dorsal, sloping down in a straight line to snout before dorsal. 
Snout rounded, mouth subterminal. Height 4, head 4.5 in length. Eye 
4 in head, once in snout, almost 4 in interorbital space. Pupil large. 
Head totally covered by soft skin. Dorsal spine (damaged) totally smooth 
in its lower part. Length of dorsal equal to snout and half eye. Adipose 
fin small, above 22 anal ray. Anal about as long as 1.5 head. Height 
of anal somewhat less than eye and snout. Length of pectoral equal to 
head. Ventrals reaching anal. Height of caudal peduncle equal to width 
of mouth. Maxillary barbels short, not surpassing eyes, mandibulary 
ones shorter. 

Colouration brownish above, lighter below. Pectorals black. Caudal 
blackish. Other fins blackish with a black border. 

One specimen with a total length of 275 mm from the Kapuas River 
(W. Borneo). Legit Dr B. M. Hoeks. 

Of the five species of Pangasiiis known from the Kapuas this 
description fits only P. micwnema. The statements about barbel 
length and eye size could apply only to P. niicroiwiiia and rule 
out the possibility of P, hoeksi being identical with P. hiimeralis 
or P. lithostoma. Until the holotype is located or additional 
specimens are collected P. hoeksi should be considered a prob- 
able synonym of P. imcronema. 

Distribution. — Sumatra (Djambi). Borneo (Kapuas, Barito, 
Mahakam). Java (Surakarta, Wonogiri, Kediri, Gempol, Sura- 
baya). 

Pangasius nasutus Bleeker, 1862 

Pungasius nasiKiis Bleeker, 1862:72 (type locality Bandjermassin, in fluviis). 

Material Examined. — Western Borneo: Kapuas 1976-19, 7: 148-215 mm 
(BMNH 1 982.3.29. 177, CAS 49407. MZB 3679, RMNH 28902. UMMZ 209855. 
USNM 230.300). Sumatra: Palembang. 270 mm (UMMZ 155721). 

Distinguished from all other Bomean Pangasius by its sharply 
pointed snout, very small eyes, relatively stout dorsal- and pec- 
toral-fin spines, and fewer anal-fin rays (26-30 vs. 30 or more). 

Two Kapuas specimens. 148-178 mm (CAS 49407). have the 
following characteristics: head length 3.7-3.8; exposed portion 
of eye 42.3-49.4; mouth width 8.0; snout 9.1-9.4; maxillary 
barbel 5.3-6. 1 ; mental barbel 7.0-8.0; predorsal length 2.6; body 
depth 4.3-4.6. width 4.8-4.9; depth of caudal peduncle 12.8- 
13.8; dorsal spine 5.7-7.0; pectoral spine 6.0-6.6; humeral pro- 
cess exposed, extending slightly beyond middle of pectoral spine, 
its length 10.9-1 1.2; pectoral pores 6-8, confined to area above 
bases of pectoral-fin rays; gill rakers 6+13=19 and 8-1-13=21; 
fins dusky; body without distinctive marks. Vertebrae in Kapuas 
specimens 18-20-1-25=43(1), 44(5), or 45(1); in Palembang 
specimen 18-1-26=44. 

Distribution. — Sumatra (Kwantan, Batu Ridial, Indragiri. 
Batang Hari, Djambi, Palembang). Borneo (Kapuas, Barilo). 

Pangasius polyuranodon Bleeker, 1852 

Paiii;asiiis polyiiratiodon Bleeker. 1852/i:425 (type localits Bandjermassing. in 

flu\ iis). 
Piiiii^u^nts itiai'o Bleeker. 1852(^:589 (type localit\ Palembang. in fluviis). 

Material Examiimed. — Western Borneo: Kapuas 1976-19. 2: 227-294 mm (CAS 
49409. MZB 3681). 



This species is distinguished by its elongate body, blunt snout, 
large eye, distinctive palatal dentition, thin dorsal- and pectoral- 
fin spines, and long anal fin with relatively numerous rays (35- 
40). Dorsal-fin origin relatively far anterior, predorsal length 3.3 



(vs. 2.5-32 in other Kapuas Pangasius). Palate with a large, 
median vomerine toothpalch with enlarged submolariform teeth 
and small lateral toothpatches with relatively fine conical teeth 
(Fig. 99c), median palatal tootpatch about 4 times as long 
anteroposteriorly as premaxillary toothbands (anteroposterior 
length of premaxillary : vomerine toothpatches from l;l to 1:3 
in other Pangasius). Vertebrae 184-32=50. 

The 227 mm Kapuas specimen (CAS 49409) may be de- 
scribed as follows: snout relatively blunt, its length 12.8; head 
length 5.0 (3.7-4.3 in other Kapuas Pangasius); eye relatively 
large, exposed portion 22.5 (26.6-49.4 in other Kapuas Pan- 
gasius except P. micwnema); mouth opening only slightly curved, 
mouth width 12.8; barbels very thin; maxillary barbel length 
6.6; mental barbel length 16.9; body depth 5.0. width 6.3; caudal 
peduncle depth 1 1.2; dorsal- and pectoral-fin spines relatively 
slender; dorsal spine length 5.7. pectoral 6.1; pectoral serrae. 
32; humeral process normally covered by skin and relatively 
short, extending not quite to middle of pectoral spine, its length 
14.4; pectoral pores confined to a small area above bases of 
pectoral-fin rays; gill rakers 12+17=29; no distinctive marks 
on body or fins. 

Distribution. — Thailand. Sumatra (Banguasin, Palembang). 
Borneo (Baram, Limbang, Kapuas, Barito, Mahakam). Java. 

Sisoridae 

Sisoridae is an exclusively Asian family of bottom-dwelling 
catfishes, most with more or less thickened leathery skins with 
specialized unculiferous tubercles or polygonal plaques (Roberts 
1982a), typically inhabiting high-gradient lowland or mountain 
streams, with adults ranging in size from about 20 mm to 2 m. 
Many of the genera are disruptively or cryptically colored and 
have adhesive organs (labial or thoracic) with which they cling 
to the substrate. The family has its greatest diversity in the 
Indian subcontinent with about 17 genera and 63 species (see 
Jayaram 1981). The family is somewhat less abundant in China 
and mainland southeast Asia; in Borneo only two genera and 
fewer than eight species are known. 

Bagarius Bleeker, 1853 

Baganiis Bleeker, 1853/;:I21 (type species Piiuelodu.s bagarms Hamilton-Bu- 
chanan, 1822, by monotypy and absolute lautonymy), 

Bagarius differs from all other Sisoridae. and perhaps from 
all other catfishes, in having 1) each premaxilla evidently com- 
posed of two ossifications instead of a single ossification, and 
2) dentition of lower jaw markedly heterodont, with 2-3 outer 
rows of relatively numerous, close-set conical teeth, and 1-2 
inner rows of less numerous, widely separated, and much larger 
conical teeth. 

For a detailed account of this south and southeast Asian si- 
sorid genus and its three species see Roberts (1983). A single 
species occurs in Borneo. 

Bagarius yarrelli (Sykes, 1839) 

(Figure 103) 

Bagrus Yanclli Sykf:s. 1839:163, 

Sagarins yarrelli Sykes, 1841 :370 (type locality Mota Mola at Poona. India). 
Bagarius licaVoh. 1903<(:557 (type locality Moesi River al Palembang. Sumatra). 
Bagarius Nu'mvenlniisi Popta, 1904:190 (type locality Mahakam basin. Borneo). 



134 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 103, Butiuiiiis yanvlli Kapuas IQ7b-31. 129 mm (MZB 3695). 



Material Examined— Sumatra; Moesi Ri\cr at Palcmbang, 410 mm (radio- 
graphs only) (BNHM VO 37T. holotype of B lieu). Western Borneo: Kapuas 
1976-19,2:309-615 mm (CAS 54646. MZB 3694). Kapuas 1976-31.2: 129-14)i 
mm (CAS 44189. MZB 3695). Additional material examined from India. Bang- 
ladesh. Burma. Thailand, Vietnam. Sumatra, and .Ia\a is listed in Roberts ( 1983: 
441-142). 

Diagnosis. — Bagantis yanvlli. attaining 2 rn and therefore 
the largest member of the family Sisondae, is distinguished from 
other Bagarnis at all si/es by having pelvic-tin origin usually 
posterior to vertical line through base of last dorsal-fin ray, gill 
rakers 8-1 1 (vs, 6-10); pectoral-fin rays 1 1-14 (vs, 9-12), elon- 
gate neural spines (see Roberts 1983) 2-5, slender (vs, 4-9. 
distally expanded); abdominal vertebrae 21-24 (vs, 17-21); and 
total vertebrae 40-45 (vs, 38-42), 

DiSTRiBVTiON. — Bagariiis yanrlli is probably the most widely 
distributed freshwater catfish in .Asia, It is more or less contin- 
uously distributed in the Indus and Ganges drainages, most of 
southern India east of the western Ghats, Burma. Thailand, 
Kampuchea, Vietnam (including Red River), Sumatra, western 
and eastern Borneo, and Java, An apparent gap in its distri- 
bution occurs in peninsular Thailand and the Malay Peninsula. 
Unknown from north Borneo. 



Glyptothorax Blyth, 1 860 

Glypimhom.x Blyth. 1860:154 (type speeies (ilypio.slcrnoii sliiauii McClelland. 
1842, b> subsequent designation of Blccker, 1862:13), 

(Jlypioihoni.x are small to moderately large sisorids with tho- 
racic adhesi\ e organ consisting of unculiferous laminae arranged 



in a whorl and confined to abdomen immediately behind isth- 
mus and between pectoral fins (Roberts, 1982a), 

There is doubt as to the number of Glyptothorax species in 
Borneo. There may be as few as three or as many as six. Three 
species occur in western Borneo (including G. plaiypogonoides, 
hitherto known only from Sumatra), The status of (7. nieuwen- 
/;((/i7 (Vaillant, 1902), G. laak (Fopla. 1904), and G, tiongiVop- 
ta, 1904), placed as synonyms of Glyptosternum (=Glyptolho- 
ra.x) Diajiis by Weber and de Beaufort (1913), needs to be 
reconsidered. .Iky.sis hara»wn.sis Fowler. 1 904. placed in Glyp- 
tostcniuin (=Glyptoihora.\) by Weber and de Beaufort (1913). 
is a true .Akysis. 

Glyptothorax major (Boulenger. 1894) 

(Figure 1114 upper! 

IkvMs iiiaior Boulenger. I 894:246 (type locality Senah. Tagora. and Baram nvers. 

Sarawak). 
"i,lvpli<\icni('iiiplatyiiogi>iuiidi'-. Vaillanl. 1893 73 (Kapuas. Sebroeang). See Weber 
andde Beaufort (1913:267), 

Material E.xamined— Malay Peninsula: Perak. Tapah Fisheries Station, 41: 
28.8-55.4 mm (CAS-SH 39344); Perak. Batang Padang. 3: 17.5-56.7 mm (CAS- 
SlI 39343); Pahang. Benus River. 49.5 mm (CAS-SU 32678); Selangor. Kuala 
Lumpur. 49,8 mm (CAS-SU 32677); Johore. Muar. 42,1 mm (CAS-SU 31003); 
.lohore. Simpang Rengam, 4; 40.6-42,8 mm (CAS-SLI 39342); Johore. Kola Ting- 
gi. 3; 41.7-54,6 (CAS-SU 32675). Western Borneo: Kapuas 1976-12. 38,6 mm 
(MZB 3999); Kapuas 1976-17. 55.7 mm (MZB 4000); Kapuas 1976-47. 3:45.9- 
47 6 mm (CAS4')416. MZB 4001. USNM 230305), 

The identits of three nominal species of Glyptoiliora.x from 
eastern Borneo placed asjunior synonyms of (7. majorby Weber 
and de Beaufort (1913:267-268) requires further study. These 



ROBERTS-WESTERN BORNEO FISHES 



135 




Figure 104. Glyptothomx. Above, G. major. Kapuas 1976-17, 55.7 mm (MZB 4000); middle, O' plaiypogon. Kapuas 1976-30, 90.7 mm (CAS 49418); below 
G. p/alypogonoides. Kapuas 1976-24, 59.2 mm (CAS 49419). 



are G. laak (Popta, 1904) and G. iiieuwenhidsi (Vaillant, 1902) 
from the Mahakam and G. //o/;,^ (Popta, 1904) from the Kajan. 

Glyptothorax platypogon (Valenciennes, 1840) 

(Figure 104 middle) 

Pimelodus plalypogon Valenciennes in Cuvier and Valenciennes. 1840:152 (type 
locality Java), 



Pimelodits cyanochloros Sleeker, 1847:1 1 (type locality Java). See Bleeker (1858: 

218). 
Glyptothorax platypogon Gunther, 1 864: 1 87. 
Glyptosterimm Kiikeiithali Steindachner, 1901:448 (type locality Baram River). 

Mateiual ExAiviiNED, — Western Borneo: Kapuas 1976-24,42.2 mm (MZB 3697); 
Kapuas 1976-27, 16: 33.1-60.1 mm (BMNH 1982.3.29.188-189, CAS 49417. 
MNHN 1982-713, MZB 3698, RMNH 28906. USNM 230306): Kapuas 1976- 
30, 3: 31.1-90.7 mm (CAS 49418, MZB 3699). 



136 



CALIFORNIA ACADEMY OF SCIENCES 



Table 9. Meristic Characters of Akvsidae. For dclinilion of vertebral counts see p. 



Gill rakers 



Pecloral 
br. ravs 



Anal ravs 



Pnncipal caudal rays 



Vertebrae 



li rihlionimuchlhys cf inelanogasler 

Kapuas 
Akysis baramensis 

Baram R., CAS 
Akvsis hcnJncksoni Alfred, I ''66 

Pahang, CAS 4821? 
Akysis cC. macroncina BIccker, 1860 

Vietnam. Chau Doc, CAS 53206 
Akysis pscudohagarnis 

Kapuas, types 
Akysis vanegatus 

Java. Brantas R . CAS 53055 
Breilensteima cf. insignis 

Djambi 

Kapuas 



1+7=8 


7 


10 






1 + 7 = 8 




10 




15-16+19=34(11,35(1) 


1+4-5 = 5-6 


6 


9-11 


7 + 6 


15+16-18=31(11,32(1), 33(1) 


1+6 = 7 


7 


10-12 


7 + 8(4) 


16+19-20 = 35(2). 36(1) 


1 + 7 = 8 


6 


10 


7 + 7(l),6 + 8(2) 


16+19 = 35(2) 


1+6 = 7 


6 


8-9 


7 + 7 or 7 + 6 


14+16-18 = 30(1), 32(1) 






10 


6?+ 5 


20 + 22=44 


2 + 3-4 = 5-6 


6 


10 


5 + 5 


21+22=43 



Glyptothorax platypogonoides {Bleeker, 1855) 

(Figure 104 lower) 

Pimclodits platypogonoides BIccker. 1855/):272 (type locality Lahat. in lluvns). 
Olypioslenion plalypogonides Bleeker. 1858c:219, 
Glyptothorax platypogonoides Bleeker. 1 862-63:63. 
Glvptosiermtin platypogonoides Giinthcr. 1864:186. 

Ragiiniis haganus Vaillant. 1 902:72 (nee Hamilton-Buchanan. 1822. embouchure 
du Raoen = Kapuasl 

Material ExAMiNED.-Sumatra?: Lahat'.', 4: 55.1-68.7 mm (RMNH 6912. 
presumed syntypes G phitvpogonoidcs): Lahat', 3: 65.5-76.0 mm (RMNH 15289). 
Western Borneo: Kapuas 1976-24.4: 53.9-64.8 mm (BMNH 1982.3.29. 190. CAS 
49414. MZB 3700. RMNH 28907): Kapuas basin, embouchure du Raoen. 61,1 
mm (RMNH 7845. originally identified as Haganus haganus). 

This species illustrates well the sort of problems encountere(J 
in attempting to identify Bleeker type specimens, even in rel- 
atively simple instances where few specimens and localities are 
involved, and despite recent clarification of the history of Bleek- 
er's collections. The original description of G. platypogonoides 
refers to four specimens 70 to 85 mm total length from Lahat, 
Sumatra. These are equivalent syntypes. In his last published 
reference to the species Bleeker stated; "Habit. Sumatra (Lahat). 
in fluviis. Longitudo 4 speciminum 70'" ad 85". Jc ne connais 
cette espdce que des rividres de la partie orientale de Sumatra." 
The entry for G. platypogonoides in the auction catalog of Bleek- 
er's collections also indicates only four specimens (Hubrecht 
1879:37). Thus there is no published indication that Bleeker 
had more than four specimens, or specimens from any locality 
other than Lahat. The four specimens in the auction catalog 
were all in lot A, and thus were obtained for the Rijksmuseum 
van Natuurlijke Historic. In Leiden I found two lots of G. 
platypogonoides collected by Bleeker, RMNH 6912. 4: 55,1- 
68.7 mm (total length 70.5-84.6 mm) and RMNH 15289, 3: 
65.5-66.0, and 76.0 mm (total length about 85 mm for first two 
specimens. 97 mm for last). Collection records indicate that all 
seven specimens in these two lots originally had been included 
in RMNH 69 1 2. The implication is that someone, at a relatively 
late date, chose four specimens from the lot of seven which 
corresponded in length with Bleeker's syntypes. The problem is 
further complicated by the presence of one more Bleeker spec- 



imen of G. platypogonoides. unfortunately now dry. in London: 
BMNH 1 863. 12.11.154. approximately 56-58 mm or total length 
76-79 mm (A. Wheeler, pers. comm., June 1984). The size of 
this specimen indicates it may be one of the syntypes. At this 
late date it seems unlikely that we shall ever know which four 
specimens are the true syntypes. or even whether all eight Bleek- 
er specimens are from Lahat. I have examined RMNH 6912 
and 15289 and compared them directly with my Kapuas ma- 
terial of G. platypogonoides. The only other specimen of the 
species known to me is BMNH 1 863. 1 2. 1 1 . 1 54 (not examined 
by me). 

Distribution. — Sumatra (Lahat). Western Borneo (Kapuas). 
So far as I have been able to determine, all records of G. 
platypogonoides from India. Burma, Thailand, and the Malay 
Peninsula are due to misidentification. 



Akysidae 

Akysids are small to minute, cryptically colored, secretive 
catfishes with tiny eyes, encountered singly or in small numbers 
and poorly represented in museum collections. All have four 
pairs of barbels, no palatal teeth, strong dorsal- and pectoral- 
fin spines, adipose fin with a long base, and tough leathery skin. 
The entire surface of the head (including lips and barbels and 
excluding only the eyes), body and fins is covered with uncu- 
liferous plaques. The basic morphological similarity of the un- 
culiferous plaques of Akysidae and Sisoridae (as illustrated by 
scanning electron microscopic observations of .-icrochordon- 
ichthys and Baganits) may indicate that these two families are 
closely related (Roberts 1 982a:70-72, fig. 20-22). Akysids differ 
from sisorids and all other catfishes in having some of the un- 
culiferous plaques on the body greatly enlarged and arranged in 
longitudinal rows of fixed number and position, a median mid- 
dorsal row and four lateral rows. The ventralmost lateral row 
may be reduced or absent in the smallest species oi Akysis. 

As here understood Akysidae comprises three genera, .4cw- 
ctiordonichthys. .4kysis. and Brettensteinia. with about 1 5 species. 
The genus Parakysis. formerly placed in Akysidae. is assigned 



ROBERTS-WESTERN BORNEO FISHES 



137 








Figure 105. Acwchordomchtliys chamaleon. Kapuas (RMNH, synlype, after Vaillant 1902). 



to a new family Parakvsidae; Ak\'sis annatus Vaillant, 1902 is 



referred to the bagrid genus Leiocassis (p. 
acters of Akysidae are given in Table 9. 



. 17). Meristic char- 



Acrochordonichthys Bleeker, 1858 

Acrochordonichrhys Bleeker. 1858c:221 (type species Acrochordonichthys plaiy- 

cephaliis Bleeker. 1858. by subsequent designation of Bleeker. 1863a:105). 
Sosia Vaillant. 1902:82 (type species Sosia chameleon Vaillant. by monotypy). 

Bleeker (1858c) recognized six species of Acrochordonichthys. 
all described by himself and additional species were described 
by Vaillant (1902) and Popta (1904). bringing the total number 
of nominal species to 1 1. I believe that the naming of so many 
species is the result of failure to recognize intraspecific variation 
in 1) coloration, 2) sexual dimorphism, 3) development of epi- 
dermal structures, and 4) development of the supraoccipital 
crest. All of these problems have been exacerbated by the fact 
that most samples of Acrochordonichthys consist of a single 
specimen. Having examined a fair number of specimens I ten- 
tatively conclude that 1) extreme individual variation in col- 
oration is the rule in Acrochordonichthys: 2) all or nearly all of 
the larger specimens are gravid females; smaller specimens, usu- 
ally difficult to sex but presumably including mature males, tend 
to differ in proportions and sometimes have been taken for 
different species; 3) the keratinous epidermal structures (uncu- 
liferous plaques) may occasionally or periodically become great- 
ly hypertrophied (e,g,, in A. rugosiis and A. pachyderma) or 
greatly reduced (presumably due to sloughing and renewal); and 
4) as in some other catfish groups in which this character has 
been used, the development of the supraoccipital crest appar- 



ently is subject to considerable individual and populational vari- 
ation. 

As things now stand it is impossible to determine either the 
number of species or what names should be applied to them. 
Weber and de Beaufort (1913) recognized four species, placed 
two nominal species as junior synonyms, and listed five nominal 
species as doubtful. There seem to be two species in the Kapuas 
basin. The names and synonymies presented below are highly 
tentative. 

Acrochordonichthys chamaleon (Vaillant, 1902) 
new combination. 

(Figure 105) 

Sosia chamaleon Vaillant, 1902:82 (type locality Kapuas basin). 

Material Examined. - Western Borneo: RMNH 7848, 2; 68.5-69.5 mm. RMNH 
7849. 1: slightly dned. RMNH 7851. 3: 58.0-98.4 mm, Kapuas basin (syntypes 
of S chanialeon). 

Weber and de Beaufort (1913:370) suggested that Sosia cha- 
maleon is conspecific with Acrochordonichthys melanogaster. I 
consider it congeneric but not conspecific. In the four largest 
syntypes (RMNH 7848. 68.5-69.5 mm and RMNH 7851,71.1- 
98.4 mm) the head is substantially broader than in any other 
Acrochordonichthys examined; in the 98.4 mm specimen head 
width is 3.0 times in standard length (head width about 4 or 
more in other Acrochordonichthys). Other features which seem 
to set these specimens apart from all other described species are 
the peculiar white coloration of anterior portion of snout, across 
occiput, and on gill covers; extremely short nasal barbels; and 
absence of tubercles on nearly the entire caudal fin (principal 
caudal-fin ravs and interradial membranes covered for their 



138 



CALIFORNIA ACADEM'l' OF SCIENCES 




Figure 106. .liriichoiJ('iiii.h:liyi d. inclanogastcr Kapuas 1476-10. 73.2 mm (MZB 36'^0). 



entire length with small tubercles in all olher. Acmcinirdonichlhys 
examined). 



Acrochordonichthys cf. melanof>aster (Bleckcr. 1854) 

(Figure 106) 

''Acnichordonichthvi mclanogaslcr Blocker. 1 854i/;8M (lype locality Sumalrae on- 
cntalis. provmc. Palembang. uhi confluum tlumina Lamalang cl Emm) 

''At_riichordomchlh\\ pcichytlcrnhi Vaillant. l')02:66 (I\pc kKalil\ riMCrc Bloc. 
Kapuas basin) 

'^.\ci'nchi^r(ionich!h\'\ oh\i'ifni\ Popla. l'M)4'187 (typo locality rivicre Boiigan), 

'' U-riichordonhhihyi Buuikntcn Popla, 1404.187 (type locality nvicrcs Bongan 
and Bo). 

" \iittihi>rdi>iiiLhih\'s vai-iu\ Piipta, 1404:1X4 (type localilN rn lerc Bo). 

MArERi.*L Examined. — Western Borneo: RMNH 7556. 101 mm. rivi^re Bon- 
gan. Kapuas hasin (holotype of .4 ohsciinis): RMNH 7557. 126 mm. and 7558. 
2: 86.8-41.5 mm, rivieres Bongan el Bo. Kapuas basin (synty pes of. I hiiclliki^U'r:): 
RMNH 7554. 2: 95.7-124 mm. nviere Bo. Kapuas basin (syntypes of .1 nirius). 
RMNH 7850. 54.0 mm. Kapuas (syntypes of .S' cliumiilivii): MZB 3690. 73.2 
mm. Kapuas 1976-10. Easlem Borneo: RMNH 7843, 105 mm, nvierc Bloeoe. 
Mahakam basin (holotype of .-I puih\'th'ti}iii] 



Akysis Bleeker. 1858 

Akvsii Blocker. 1858( -234 (type species I'lniclodus varicgunis Blocker. 1847. by 
monot>p\ ) 

The Aky-iis of Indonesia and Kalimantan in particular are 
poorly known. The only species of Akysis previously known 
from Borneo, .-1. haramcnsis Fowler, 1904, has not been rec- 
ognized as an akysid since its incorrect placement in the sisorid 
genus Glyplostcrnum by Weber and de Beaufort (1913:265). I 
have examined the type specimens and find they represent a 
valid species oC Akysis. The only other species of Akysis de- 
scribed or reported from Borneo, A. arnialiis Vaillant, 1902, is 
in fact a bagrid and is herein assigned to Leiocassis. The Kapuas 
survey of 1976 obtained a single undescribed species of Akysis. 

Akysis pseudoba)jarius new species 

(Figure 107) 



HoLOTVPE. — MZB 3641. 48 2 mm. mauiic lemale' 
Mclaui at Sintang. 24 ,Iuly 1476 (Kapuas 1476-31). 



bar in mouth of Sungai 



ROBERTS-WESTERN BORNEO FISHES 



134 




Figure 107. Akysis psi'uJobuganus. Rapuas 1976-31 



mm mature female (MZB 3691. hololype). 



Paratypes.-CAS 49414. MZB 3692, 2: 21.4-23 3 mm. collected with holo- 
tvpe; UMMZ 155702. 47.2 mm. gravid female. Sumatra. Mocsi R. at Moera 
Klingi. A. Thienemann. 10 May 1929. 

Diagnosis.— An Akysis with exceptionally narrow and elon- 
gate head and body and a color pattern of well defined brown 
marks and bands on a cream-colored background. Snout slightly 
pointed (not broadly but evenly rounded), strongly overhanging 
lower jaw; premaxillary tooth band entirely exposed when mouth 
is closed; inner margin of pectoral-fin spine with 8-9 serrae more 
or less evenly spaced over its entire length (even in very small 
specimens); vertebrae 35. 

The following observations were made on the 48.2 mm Ka- 
puas holotype and 47.2 mm Moesi River paratype (these two 
specimens are very similar in all respects): head 3.9-4.0; eye 
39-48; interorbital space somewhat narrow. 22.4-25.4; snout 
9.5-9.7; width of mouth opening 10.1-11.2; gill rakers 1+6- 
7=7 or 8; nasal barbel extending posteriorly almost to posterior 
border ofeye or slightly beyond eye. length 1 1.2-1 1.3; maxillary 
barbel extending postenorly just beyond base of pectoral-fin 
spine. 4.1-4.8; outer mental barbel extending posteriorly to mid- 
dle or distal two-thirds of pectoral-fin spine. 5.4-6.0; side of 
body with 6 longitudinal rows of enlarged elongate epidermal 
tubercles or plaques (fourth row from top straddling lateral line 
canal); mid-dorsal row of enlarged tubercles weakly developed 
or absent; body depth 6.7-6.9, width 5.0-5.4; caudal peduncle 
length 4.7-5.1, depth 16.3-16.6; pectoral fin with short "fila- 
mentous" extension, extending posteriorly when adpressed al- 
most to middle of pelvic fin; more posterior than anal-fin origin; 
anal-fin rays ii or iii7-l/2; caudal fin deeply forked, lobes equal 



in length but lower lobe somewhat larger than upper; principal 
caudal-fin rays 6 + 8; vertebrae 16+ 19 = 35. 

The two small Kapuas paratypes, only 21.4-23.3 mm, arc 
very similar in nearly all respects to the two large specimens 
described above. The eyes and barbels have nearly the same 
relative proportions and the color pattern, although somewhat 
less well marked, is also the same; even the number of pectoral- 
fin spine serrae is similar (8 or 9). The longitudinal rows of 
epidermal tubercles or plaques are very evident, although rows 
3 and 6 are very weakly developed. The most striking differences 
seem to be that the basal membrane connecting the maxillary 
barbel to the check is relatively much larger, and the pectoral- 
fin spines proportionately longer, length 3.9-4.1. 

Live A. pseudobagarius have a handsome pattern of rich brown 
marks and transverse bands and cream-colored background on 
dorsolateral surfaces of head and body (ventral surfaces pale, 
without marks). Head with a brown blotch centered on each 
nostril, a narrow vertical brown bar below eye, and a broad 
transverse brown band (complete dorsally) extending over gill 
cover on posteriormost part of head immediately anterior to 
pectoral girdle and fin; dorsal surface of body with a large but 
poorly defined and somewhat irregularly shaped brown blotch 
anterior to dorsal fin, and two small middorsal faint oval brown 
marks, one midway between dorsal and adipose fin, the second 
posterior to adipose fin; lateral surface of body with a series of 
three well defined and nearly equally spaced broad transverse 
brown bands, the first centered below dorsal fin. the second 
below adipose fin, and the third on posterior half of caudal 
peduncle. A fourth transverse brown band, similar to those on 



140 



CALIFORNIA ACADEMY OF SCIENCES 




Figure lOX lin'ilcii^h-ima cf /«M,i;/i« Kapuas l97h-45. Above, ^3.0 mm (CAS 494151; below, 191 mm female (CAS 49415). 



body, occurs on distal half of caudal fin. All four bands with 
narrow midlateral anterior prolongations. Dorsal and adipose 
fins with brown coloration continuous with brown bands on 
side of body; pectoral fin with brownish mark basally; anal fin 
colorless or with a narrow brown band extending across middle 
of posteriormost four fin rays; pelvic fins colorless; nasal barbel 
with faint brownish marks (not well defined), barbels otherwise 
entirely or almost entirely colorless. 

The shape, coloration, and overall appearance of .4. pseudo- 
hagahiis is strikingly similar to that of Bagarius (before I ex- 
amined it, the paratype from Moesi River actually had been 
mistaken for Baganm). At Rapuas 1976-31 A. pscudobaganus 
and Bagarius yanelli were collected from a single seine haul at 
night and compared side by side while alive. The uncanny re- 
semblance includes color quality as well as nearly identical dis- 
tribution and shape of markings on head, body and fins. 

Alternating brown and cream-colored or yellowish banding 
comparable to that just described occur in many catfishes typ- 
ically found over gravel bars or riffles. Among these are African 
amphiliids and mochokids. South American pimelodids, and 
other Asian sisorids (e.g., Nangra and several species of Glyp- 
lothorax. including G. platypogoiuvdes which has also been mis- 
taken for Bagarius). There is also another Akysis. A. leucorhyn- 
(7;;« Fowler (1934:97. fig. 44,45). This species, from theMenam 
Chao Phr\'a, probably is not closely related to.-l. pscudobaganus: 
It differs in havmg snout more truncate, intcrorbital space broad- 
er, posterior portion of head laterally expanded, body less elon- 
gate, and adipose fin smaller. 

Distribution. — Known only from Moesi and Kapuas rivers. 

Et\mology. — The name pscudobaganus refers to the striking 
superficial (but phylogenetically false) resemblance of this species 
to members of the sisorid catfish genus Bagarius. 



Breitensteinia Steindachner. 1881 

Hrciicif.lcima Stemdachner, 1881i;:2l.l (type species Brcilcmlcinia insigiiisSXem- 
dachner, 1S81. b\ nionot\p\) 



Breitensteinia cf insignis Steindachner. 1881 

(Figure HIS) 

Hrcilcn^lcima insifiius Steindachner. 188 ii/:;!!.^ (type locality Teweh = Moara Te- 

weh. Barito River, southeast BoiTieo). 
BrcilenaciniLi insigiiis Va\\\arn. 1902:76. hgurcs 16. l7(K.apuasl. 

Material Examined. — Sumatra Djamhi, 77.3 mm (ZMA 104.653), Western 
liorneo: Sinlang, 176 mm female (RMNH 7847). Sanggau, 125 mm male (RMNH 
16048); Kapuas 1976-45, 4: 93,0-191 mm (CAS 49415, MZB 3693, USNM 
230304), Southeast Borneo: Moara Teweh, Banto-Flliss, I 33 mm (NMW 55042, 
holotype). 

The material examined represents all specimens of the genus 
Breitensteinia known to me. Through the kindness of the late 
Rainer Hacker I was able to examine the holotype of B. insignis 
during a visit to Vienna in October 1982. The holotype (sex 
undetermined) is in somewhat poor condition; perhaps it had 
been dead for some time before preservation. It is now faded 
or bleached to a uniform pale white presumably due to exhi- 
bition or storage in daylight. Although I examined specimens 
of Breitensteinia from Sumatra and Borneo before and shortly 
after examining the holotype, it was not possible to compare 
them directly. While the specimens from Sumatra and western 
Borneo appear to be conspecific, I am not certain that they are 
the same species as the holotype. 

The 125 mm specimen from Sanggau is of particular interest 
because it is a sexualK mature male; each testis has about 10 
elongate, digitiform lateral processes or caeca. The 176 mm 



ROBERTS-WESTERN BORNEO FISHES 



141 




Figure 109. Parakysis verrucosa. Malay Peninsula, Pahang, 42.5 mm gravid Icmalc (USNM 266589). 



specimen from Sintang and 191 mm specimen from Kapuas 
1 976-45 are gravid females. The gonads of other large specimens 
were not examined, but I suspect that they are females. Thus, 
the 125 mm specimen is the only known mature male of Brei- 
tensteima. This specimen differs so much from the more elon- 
gate females that it could represent a different species. The char- 
acteristics of the three mature specimens of known sex are 
summarized in Table 9. Two specimens had small prawns in 
their stomach (no other food items observed). 

Parakysidae new family 

In 1937 A. W. Herre collected some very peculiar little cat- 
fishes in Sarawak and in the southern part of the Malay Pen- 
insula which he subsequently described as a new genus and 
sptcKS of Miysxdae, Parakysis verrucosa (¥'\g. 109). Presumably 
Herre referred Parakysis to Akysidae on the basis of its highly 
tuberculate skin, cryptic coloration, small size, superficial re- 
semblance to Akysis. and occurrence in southeast Asia. He ap- 
parently did not compare the specimens of his new genus with 
any akysid specimens. It is possible that Herre never saw a true 
member of the family Akysidae: there were no specimens in the 
fish collection of the Stanford Natural History Museum during 



his time, and so far as I have been able to determine he never 
obtained any. 

Direct comparison of Parakysis with the three genera of Akys- 
idae— .•lcTOf/;o/'(/o/»r/;//?v.5. Akysis. and Breitensteinia — reweah 
little in common and numerous differences. Parakysis is highly 
specialized in several respects and does not seem closely related 
to any other catfishes. and therefore a new family is proposed 
for it. 

The family Parakysidae is characterized as follows: 

1. Head and body entirely covered with evenly distributed, 
rounded tubercles of nearly uniform size. 

The tubercles of Parakysis exhibit no indication of organi- 
zation into longitudinal rows which is such a striking charac- 
teristic of Akysidae. The rounded tubercles of Parakysis have 
not been examined with the scanning electron microscope but 
superficially they do not resemble the polygonal unculiferous 
tubercles observed in akysids and sisorids. In Parakysis the 
tubercles vary from pale to highly melanistic, while in Akysidae 
and Sisoridae they are generally (always?) unpigmented. At least 
some of the tubercles of Parakysis apparently have a depression 
(or pore?) at the summit, thus suggesting a possible sensory or 
secretory function absent in the poreless tubercles of Akysidae 
and Sisoridae. 



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CALIFORNIA ACADEMY OF SCIENCES 



2. Lips with transverse plicae; lower lip with a pair of ante- 
riorly projecting lateral lobes. 

The number of transverse plicae increases with growth. The 
largest specimen examined, a 59.8 mm P. verrucosa, has the 
main part of the upper lip with 7 complete transverse plicae 
and lower lip with 4 complete transverse plicae plus 4-5 plicae 
on each lateral lobe. The plicae superficially resemble those on 
the expanded lips of the cyprinid genus Osteochilus. In Akysidae 
and Sisoridae, as in catfishes generally, the lips are generally 
tubcrculate or papillose; no other Asian catfishes with trans- 
versely plicate lips have come to my attention. The lateral lobes 
of the lower lip of Parakysis are unique. 

3. Nasal, maxillary, and two pairs of mandibular or mental 
barbels well developed. Base of outer mandibular barbel with 
0-1 and of inner mental barbel with 0-} short accessory barbels. 
Maxillary barbel with a broad base originating at rictus of jaws 
and continuous with both upper and lower lips. 

Parakysis has the full catfish complement of four pairs of 
barbels, with the specializations noted above which are unique. 
Comparable accessory mental barbels are not found in any other 
Asian catfishes; the highly branched mental barbels of African 
Mochokidae and South American Doradidae are very different 
from the simple accessory mental barbels of Parakysis. In Akys- 
idae and Sisoridae. as in catfishes generally, base of maxillary 
barbel originates from or above upper lip only. 

4. Gill rakers absent. 

There are no gill rakers on any of the gill arches in Parakysis. 
In Akysidae gill rakers are reduced in number but are large and 
well developed and present on the leading edge of all arches 
except the last. The only other Asian catfishes as known to me 
in which the gill rakers are entirely absent belong to the highly 
specialized and unrelated family Chacidae. 

5. Dorsal and pectoral spines short, stout, and non-serrate. 
The dorsal-fin spine of Parakysis is exceptionally stout and 

wide for such a small catfish, much more so than the dorsal 
spine of any akysid or sisorid. The pectoral spine of nearly all 
akysids and sisorids bears large scrrae on its inner margin. 

6. Dorsal fin with 4, pectoral and pelvic fins with 5-6 soft 
rays. 

In catfishes the primitive or generalized complement of dor- 
sal-fin soft rays is 7. In small catfishes with a well developed 
dorsal-fin spine the number of rays is commonly reduced to 6 
or 5, rarely to 4 (never fewer?). In akysids the dorsal-fin rays 
are usually 5. 

7. Anal fin with 8-1 .^ rays, anteriormost .V8 rays simple and 
widely spaced. 

The number of simple or unbranchcd anal-fin rays m Akys- 
idae and Sisoridae, as in most catfishes, is usually no more than 
3-5; in catfishes with up to five unbranched rays, the first 2^ 
rays tend to be very close together (sometimes all based on the 
first anal pterygiophore). Perhaps the only Asian catfishes with 
numerous widely spaced simple anal-fin rays comparable to 
those of Parakysis are the members of Olyridae and Amblyci- 
pitidae. 

8. Humeral (or postcleithral) process absent. Pectoral gland 
present, with a single pectoral pore. 

The humeral process of the pectoral girdle is well developed 
in all Akysidae. In Parakysis there is a slight convexity on the 
posterior margin of the cleithrum near where the humeral pro- 
cess normally arises, but nothing like the well developed hu- 



meral process seen in nearly all Asian catfishes including Akys- 
idae and Sisondae. 

9. Caudal fin with upper lobe moderately elongate and point- 
ed, lower lobe shorter and rounded. Caudal fin with 3-5 upper 
and 3-4 lower procurrent rays and 7/6 or 7/5 principal rays. 

In Akysidae the upper and lower lobes are generally similar 
in shape, with the lower equal to or slightly longer than upper. 

In catfishes the principal caudal-fin ray formula is almost 
always n/n or n/n+1, with n ranging from 9 (the primitive 
number of catfishes, found only in Diploniysies) to 5. Many 
species have 8/9=17. The lowest principal ray count known in 
catfishes is 5/5=10, found in several highly specialized and un- 
related genera in different families. Although no exceptions to 
this formula were reported in the extensive survey of catfish 
caudal skeletons by Lundberg and Baskin (1969), at least three 
genera from southeast Asia depart from the rule; Parakysis with 
7/6 or 7/5, Akysis hiitchinsoni with 7/6, and the silurid Silu- 
riciithys wixh 7/6, 7/5, or 7/4. Akysidae other than .-I. hiitchinsoni 
usually have n/n or n+ 1 principal caudal rays (Table 9). 

10. Lateral line apparently absent from side of body. 

In most catfishes the lateral line is well developed on the side 
of the body. Akysidae have a series of lateral line pores at least 
on the anterior half of the body. .Acrochordonichthys has the 
lateral line with 23 pores extending to a point below end of 
adipose fin; Breitenstcinia has the lateral line with 27 pores 
extending to well beyond the anal fin. In .Akysis variegatus, the 
smallest akysid, the lateral line apparently is confined to the 
anterior half of the body, but has 5 well developed pores. No 
pores or other indication of a lateral line have been observed 
on the body of Parakysis. 

11. Vertebrae 16-19+11-14=30-32. 

In Parakysis the number of abdominal vertebrae is 2-7 more 
than of caudal vertebrae, while the reverse is generally true in 
.Akysidae (see Table 9). 

Additional features of Parakysidac. mainly primitive or re- 
duction characters of less help in establishing its distinctness or 
relationships, include the following: eyes minute, dorsal in po- 
sition; teeth of upper and lower jaws generalized, disposed in 
narrow bands; palatal teeth absent; gill openings restricted, ex- 
tending from just below to just above lateral prominence of 
cleithrum; isthmus broad; branchiostegal rays 5 + 5; anterior 
chamber of swim bladder with lateral lobes; adipose fin absent 
or represented by a long, low-lying dorsomedian ridge of almost 
uniform height beginning just behind dorsal fin and continuous 
with low ridge formed by upper procurrent caudal-fin rays; ma- 
ture males with moderately elongate genital papilla. 

The family Parakysidae comprises the genus Parakysis with 
two species. P. vcrnicimi from the Mala> Peninsula. Sarawak, 
and Sumatra, and P. aiioniah:>ptcry.\ new species from the Ka- 
puas basin. 

Parakysis Herre, 1940 

l'uruk\'\n Herre. l'J40u IJ! (lype species FarukyM.\ wiiiuvsa Herre. l'^40, by 
nionnispy). 

Parakysis anomaloptery\ new species 

HoLOTVPE — MZB .1702. 27.1 mm. Kapuas basm. Sungai Seriang. a forested 
tributary ollhe Sungai Palin. .17 km W of Putussibau (Kapuas 1976-42) 

Paratvpes.-MNHN 1 '(82- 7 14, 2: 2.1 0-24,1 mm. same collection as hololype; 



ROBERTS- WESTERN BORNEO FISHES 



143 




Figure 110. Chuca hankuiwmiS- Kapuas 1976-47, 101 mm (USNM 230310). 



AMNH 48940, BMNH 1982.3.29.151-192, CAS 49420, FMNH 94245, MCZ 
58352, MZB 3701, 18: 19,8-34,1 mm, Kapuas basm, Sungai Mandai Kctchil near 
confluence with Kapuas mainstream, 18 km WSW of Pulussibau (Kapuas 1976- 
39), CAS 4942 1 , MZB 3703, UMMZ 209923, USNM 230307. 1 8: 1 8,5-30.4 mm. 
Kapuas basin, small forest stream flowmg mto Kapuas mamstream NE of Gunung 
Selunggul, 53 km NW of Sintang (Kapuas 1976-47). 

Diagnosis. — f. anoinalopteryx is superficially similar to its 
only congener, P. verrucosa (Fig. 109), but differs from that 
species in fin ray counts, shape of upper lip, number of branches 
on mental barbels, and number of pores in mandibular branch 
of cephalic laterosensory canals. Pectoral-fin rays usually i5 (vs. 
16 in P. verrucosa); pelvic-fin rays 5 (vs. 6); caudal-fin rays almost 
invariably 7/5 (vs. usually 7/6); total anal-fin rays 10-13, 5-8 
simple + 3-7 branched (vs. total 8-11, 3-7 simple + 5-6 
branched). Upper lip with uniform margin (vs. upper lip with 
a discrete median lobe between anterior nostrils projecting ven- 
trally between lateral lobes of lower lip). Outer mandibular bar- 
bel almost invariably without branches, inner mandibular bar- 
bel almost invariably with a single branch (vs. outer mandibular 
barbel usually with one branch and inner with two or three 
branches). Mandibular laterosensory canal without a pore lateral 
to outer mandibular barbel (vs. with a pore). 

Distribution. — Kapuas. Its only congener, P. verrucosa, is 
known from Malay Peninsula (Pahang, Johore). Sumatra (Deli), 
and Sarawak. 

Etymology. — The name a//o»;iv/t)/V(vr.v refers to the unusual 
or "anomalous" fin ray counts characteristic of this species. 

Chacidae 

The south and southeast Asian angler-catfish family Chacidae 
is distinguished by its extraordinarily large, broad, flat head, 
nearly square in outline when viewed from above; mouth ter- 
minal, very broad; tongue very broad, occupying nearly entire 
floor of mouth; maxilla entirely included in gape; palate eden- 
tulous; gill arches greatly enlarged and elongate; gill rakers ab- 
sent; gill opening moderately large, extending onto isthmus only 
a short distance anteroventral to pectoral fin; isthmus extremely 
broad; branchiostegal rays 6-8; eyes minute; three or four pairs 
of minute or very small barbels (nasal barbel absent in C. chaca): 
numerous cutaneous flaps or cirri on dorsolateral surface of 
head, sometimes also on lower lip and on body. Skin extensively 



granulated. Dorsal fin with a strong spine and four rays. Anal 
fin with 8-10 rays. Pectoral fin with a short, strongly serrated 
spine and 4-5 rays; pelvic fins large, with 6 rays; caudal fin 
rounded with 5/6 or 6/5 principal rays; upper and lower pro- 
current rays greatly enlarged and numerous, 18-25/8-13. Ver- 
tebrae 14-16+17-20=31-35. 

A single genus, Chaca. with two species, C. chaca (Hamilton- 
Buchanan, 1 822) in the Indian subcontinent and C. bankanensis 
in the Malay Peninsula and Indonesia. For further information 
on the family see Roberts (1982/i). 

Chaca Gray, 1831 

Chaca Gray. 1831 (type species Plaiv\iacns chaca Hamilton-Buchanan. 1822, by 
monotypy and absolute tautonymy) 

Chaca bankanensis, Bleeker, 1852 

(Figure I 10) 

Chaca bankanensis Bleeker. 1852c:455 (type locality Banka, in lluviis). 

Material Examined.— Western Borneo: Kapuas 1976-16. 4: 46.0-89.2 mm 
(CAS 49424. MCZ 57598. MZB 3709); Kapuas 1976-14, 164 mm (MZB 3710); 
Kapuas 1976-42, 2: 15,2-17.9 mm (CAS 49425. MZB 371 1); Kapuas 1976-46, 
3: 3* 4-101 mm (MZB 3712. USNM 230310). 

Distribution. — Malay Peninsula (Johore, Perak). Sumatra 
(Indragiri). Borneo (Kapuas, Barito). Banka. Billiton. 

Siluridae 

The Eurasian Siluridae is one of the most distinctive and well 
defined families of catfishes. Head depressed, body compressed, 
skin smooth; maxillary barbels usually elongate; one or (in 
Parasilurus and Sthirus) two pairs of mental barbels (sometimes 
vestigial or absent); nasal barbels invanably absent; gill openings 
very wide; branchiostegal membranes, more or less broadly 
overlapping, with 8-21 branchiostegal rays; dorsal fin spineless, 
with 1-8 (usually 1-4) soft rays (sometimes absent), and exhib- 
iting a fixed position in relation to vertebral column unlike that 
in other catfishes (see below); anal fin very long with 41-1 12 
rays; abdominal cavity restricted; caudal vertebrae about 3-4 
times as numerous as abdominal; vertebrae 9-20 + 32-56=42- 
75. 

In all silurids. including those without dorsal-fin rays, well 



144 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 111, Hclntlonlichlhys dincma Kapuas 1976-1 'J. 246 mm (CAS 49386). 



developed dorsal-fin pterygiophores articulate with neural pro- 
cess of vertebra 7, or, in, the three closely related genera Hito. 
Parasilunts and Silurus. with neural process of vertebra 8 or 9. 
In other catfish families dorsal-fin pterygiophores generally ar- 
ticulate with neural processes of several vertebrae beginning 
with vertebra 4 or even with vertebra 1 or 2. 

The family comprises 9-10 genera and about 50 species (for 
systematic review see Haig 1952). The genera and species are 
most numerous in the Indian subcontinent, Mekong basin, and 
Borneo; seven genera and 2 1 species (two herein described as 
new) are known from western Borneo. Of particular interest are 
the highly specialized but poorly known endemic southeast Asian 
genera Bclodoniichlhys. Ceraloglanis. and Hennsilitrus. 

Belodontichthys Bleeker, 1858 

Hclotltiiinchlhys Blocker. 1 85Si:266 (lype species B(/i'i/('/»/i/?///n luacrnchir Blocker. 
\>siii=\\\illuiii> ilincnia BIcekor, 18M. hy monol>p\). 

A highly specialized and very distinctive genus. Head and 
body strongly compressed; predorsal profile strongly concave 
from tip of strongly upturned snout to above eye, then almost 
perfectly straight to dorsal-fin origin; dorsal body musculature 
extending very far forward onto head, to above middle of eyes 
(as in cyprinid Macrochirichlhys): depth of body strongly de- 
creasing from anal-fin origin to caudal-fin base; jaws long, com- 
pressed, strongly upturned, lower jaw strongly projecting, rictus 
below anterior margin of eye; jaw teeth large, lancet-shaped, in 
three irregular rows in both jaws, teeth in outermost row some- 
what smaller; vomerine teeth minute, restricted to a small patch; 
eyes large, lateral (not visible from below head), subcutaneous; 
ma.xillarv' barbels extending posteriorly to beyond level of pec- 
toral-fin origin and almost to pelvic-fin origin; mental barbels 
minute, vestigial; gill openings very large; branchiostegal mem- 
branes free from isthmus with 13-15 branchiostegal rays; gill 
arches extremely long; gill rakers short but relatively numerous 
(4-f 29=33 in 296 mm Kapuas specimen); dorsal fin small, with 
4-5 rays; anal fin long, with 88-96 rays; pectoral fin falcate and 
extraordinarily large, extending posterioriy well beyond level of 
anal-fin origin, with a basally rigid, distally flexible (i.e., non- 
pungent), non-serrate spine and 1 8-22 branched rays; cicithrum 
and pectoral muscles enormously expanded; humeral process 



absent; porus pectoralis and axillary gland absent; pelvic fin 
small, with 9-10 rays; caudal fin small, separate from anal fin, 
upper lobe slightly longer than lower with 8/9 principal and 
about 4/3 procurrent caudal-fin rays; vertebrae 13 or 14 + 53=66- 
67 (in one Kapuas specimen); color in life (Kapuas) iridescent 
greenish-blue dorsally, silvery or milky-white laterally and ven- 
trally; attains 550 mm. 
The genus comprises a single species. 

Belodontichthys dinema (Bleeker, 1851) 

(Figure 1 1 1 1 

Wallatio dim-ina Blocker. IKSI/i I4S, 2U2 (Upc locality Banjermassin). 
Bclodonlichlhvs inacmchir Bleeker. IS58i:2b6 (unwarranled subslilution of species 

name for If. dincma). 
Bclodoniichlhys diiieina Webor and do Beaufort. 1 913:204. 

WalUigo dincma Myers, 1938:98. 

Material Examined. - Western Borneo: Kapuas 1976-19. 3: 193-296 mm (CAS 
49386, MZB 3630. UMMZ 209859), 

Distribution. — Laos, Vietnam, Thailand (Mekong, Chao 
Phrya). Malay Peninsula. Sumatra (Palembang, Djambi). Bor- 
neo (Kapuas, Barito). 

Ceraloglanis Myers, 1938 

Ceraloglanis Myers. 1938:98 (type species Ilcnusilwm sclcroncnia Blocker, 1862, 
by ongmal designation and monotypy) 

Ceraloglanis is distinguished from all other silurids by its 
hook-shaped maxillary barbels (barbels permanently bent in a 
right angle near midlength) and more numerous anal-fin rays 
( 1 03-1 1 2 vs. less than 1 00 in all other silunds). Vertebrae 12 + 51- 
53=63-65. In shape of head and of its inferior, transverse jaws 
Ceraloglanis strongly resembles Hemisiliirus but its postenor 
nostril lies anterior to eye rather than posterior to it as in 
Henusilurus. Vertebral counts are virtually identical in Ceral- 
oglanis and Heniisilurus. 

Ceratoglanis scleronema (Bleeker, 1862) 

Ucmisihiiiis wicroncma Blocker. 1862-63:93 (type locality Krawang, .lava). 
Ceraloglanis scleronema Myers, 1938:98 

Material Examined. -Wcslem Borneo Kapuas 1976-19, 14: 226-348 mm 
(CAS 49387. MZB 3631. RMNH 28889. HSNM 230285). 



ROBERTS-WESTERN BORNEO FISHES 



145 



Kapuas specimens have gill rakers 2-3+ 1 1-12=13(2), 14(2), 
or 15(1); anal-fin rays 103-112; vertebrae 12 + 51-53=63(2), 
64(1), 65(2). 

Distribution. — Sumatra (Djambi). Borneo (Baram. Ka- 
puas). Java (Krawang). 

Hemisilurus Bleeker, 1858 

Hemisiliinis Bleeker. 1858cr295 (type species H'allago hcwrorhynclios Bleeker, 

1853, by subsequent designation of Bleeker, 1862:395). 
Diaslaloinycter Vaillant, 1891:182 (type species Diastatoinyctcr chaperi VaiUant, 

\%<^\ = WaUago heterorhynchos 'RXeeV.er. 1853, by monotypy). 

Hemisilurus are large silurids, attaining at least 80 cm (pers. 
obs.), distinguished from all other genera by having posterior 
nostril lying behind posterior margin of eye (rather than in front 
of eye) and in having maxillary barbel (simple, short, and thin 
in juveniles) becoming greatly enlarged with its distal half flat- 
tened and bearing a series of thread-like extensions in large 
adults. This condition of the barbels was reported by Weber 
andde Beaufort (1913:213, fig. 85) in a female H. moolenburghi 
presumably over 500 mm (length not stated). Haig (1952:64) 
assumed that the condition occurs only in females but this needs 
confirmation. I observed a number of very large specimens of 
Hemisilurus in the fish market at Sintang (Kapuas 1976-19), 
several of which had enlarged, modified maxillary barbels. It 
seems likely that these were H. heterorhynchos rather than H. 
moolenburghi; unfortunately the specimens were not obtained, 
and it was not possible to sex them or confirm the specific 
identity. The secondary growth and differentiation of the max- 
illary barbel apparently does not occur until fish are well over 
400 mm (and presumably have attained sexual maturity). Anal- 
fin rays 90-98. Vertebrae 12 + 51-52=63-64. 

Hemisilurus heterorhynchos (Bleeker, 1853) 

Wallago heWrorhyiichos Bleeker. 1853/:514 (type locality Moara K-ompeh, Su- 

matrae onentalis. in fluviis). 
Hemisilurus helewrhynchus Bleeker. 1862:94. 

Diaslalomycler chaperi Vaillant. 1891:182 (type locality Knapei. Kapuas basin), 
Hemisilurus chaperi Weber and de Beaufort, 1913:211. 

Material Examined. — Western Borneo: Kapuas 1976-19, 18: 215-350 mm 
(BMNH 1982.3.29. 161-1 62. CAS 49388. MNHN 1982-706. MZB 3632. RMNH 
28890, LIMMZ 209860. USNM 230286): Kapuas 1976-48, 136 mm (MZB 3633), 

Kapuas specimens have gill rakers 4 + 9-10=13(1) or 14(1), 
anal-fin rays 92-98, and vertebrae 12 + 51-52=63(1), 64(1). 

Although Weber and de Beaufort ( 1 9 1 3:2 1 1 ) and Haig (1952: 
94) treat Hemisilurus chaperi as a valid species, it is noteworthy 
that Vaillant himself treated it as a junior synonym. Vaillant 
had not examined specimens of either Hemisilurus heterorhyn- 
chos or Hemisilurus (=Ceratoglanis) when the original descrip- 
tion of his Diaslalomycler chaperi was prepared. Later, having 
examined type specimens of the three nominal species, Vaillant 
(1902:50) concluded that they represented only two species, H. 
helerorhynchos and H. scleronema. It is perfectly clear from the 
discussion that he recognized the identity of £). chaperi with H. 
helerorhynchos and that the statement "Diaslalomycler Chaperi. 
Vaillant=//. scleronema. Bleeker" is a lapsus calami. 

The 19 Kapuas specimens here reported as //. hclerorhynchus 
appear to represent a single species; they agree quite well with 
Bleeker's accounts of H. heterorhynchos and Vaillant's of D. 
chaperi. I therefore conclude that Vaillant was correct in rec- 



ognizing the specific identity of//, heterorhynchos and D. chap- 
en. 

Distribution. — Sumatra (Palembang, Djambi). Borneo (Ka- 
puas). 

Hemisilurus moolenburghi Weber and de Beaufort, 1913 

Hemisilurus miiolenhurghi Weber :ir\d de Beaufort. 19 13:2 12 (type locality Batang 
Han, Sumatra). 

Material Examined —Western Borneo: Kapuas 1976-19, 308 mm (MZB 3634). 

A single specimen of Hemisilurus obtained at the fish market 
in Sintang diflTers from all of the others obtained there in having 
a much smaller eye and more elongate but still very slender 
maxillary barbel. It thus agrees with //. moolenburghi, a species 
previously known only from the type specimens obtained in the 
Batang Hari, Sumatra. 



Kryptopterus Bleeker, 1858 

Kryploplerus Bleeker, 1858c:283 (type species Kryptopterus micropus Bleeker. 

l&iS=Silunus crvptopierus Bleeker. 1 85 1 . by subsequent designation of Bleeker, 

1862-63:18). 
Kryptoplenchlhys Bleeker. 1 858c :288 (type species Silunis palembangensis Bleek- 
er. \S52=Silurus hicirrhis Valenciennes in Cuvier and Valenciennes. 1839, by 

subsequent designation of Bleeker, 1862-63:18). 
Micronema Bleeker, 1858t:298 (type species Micronema lypiis Bleeker, 

\m=Silurus micronemus Bleeker, 1846, by original designation). 
Phalacroiiotus Bleeker, 1858i:302 (type species .Silunis leplonema Bleeker, 1852, 

by subsequent designation of Bleeker. 1862-63:18), 
Cryptoplerus Giinther. 1864:38 (unwarranted spelling emendation), 
Cryplopterella Fowler. 1944:1 (type species Cryptoplerella heldli Fowler, 

\')^'i=Kryptopterichihys macrocephalus Bleeker. 1858?). 

Bleeker (1858) divided the fishes here placed in Kryptopterus 
into two tribes, Kryptopterini and Phalacronoti, each with two 
genera. These taxa, based mainly on reduction characters such 
as presence or absence of dorsal fin and mental barbels, have 
not been followed by subsequent authors. Kryptopterini (with 
Kryptopterus a.nA Kryptopterichlhys) supposedly includes one or 
two large species and small species which have 8-12 branchios- 
legal rays, while Phalacronoti (with Micronema and Phalacro- 
nolus) includes only large species with 12-17 branchiostegal 
rays. Whether the taxa thus recognized are phyletically valid 
has not been sufficiently investigated. 

The genus has not been revised since Bleeker (I858r, 1862- 
63). The species are listed by Haig (1952). Counts of vertebrae, 
not previously given, and of gill rakers, rarely provided, appear 
to be characteristic or diagnostic for various species. Data on 
these and other meristic characters are presented in Table 10, 



Kryptopterus apogon (Bleeker, 1851) 

(Figure 112) 

Silurus apogon Bleeker. 1851/:67 (type locality Bandjermassing. in fluviis). 
Silurus leplonema Bleeker. 1852(/:584 (type locality Palembang, in fluviis). 
Silurus micropogon Bleeker. 1855</:418 (unwarranted replacement name for .S;- 

lurus apogon Bleeker. 1851), 
Phalacronolus leplonema Bleeker, 1858i:304. 
Plialacronolus micropogon Bleeker, 1858c:306. 
Crvplopterus micropogon Giinther, 1864:43, 
Crvptopierus apogon Weber and de Beaufort, 1913:221. 
Kryptopterus apogon Smith, 1933:75. 



146 



CALIFORNIA ACADEMY OF SCIENCES 



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ROBERTS-WESTERN BORNEO FISHES 



147 




Figure 112. Kryptopterus apogoth Kapuas 1976-19, 318 mm (CAS 49389). 



M,'\TERiAL Examined. —Weslem Borneo: Kapuas 1976-19. 4: 173-318 mm (CAS 
49389, MZB 3635, RMNH 28891, USNM 230287); Smtang. 204 mm (ZMA 
114.382). 

Diagnosis. — Probably the largest species of Kryptopterus. at- 
taining a total length of 770 mm (Weber and de Beaufort 1913: 
221); head more elongate than in any other Kryptopterus. its 
length 4.6-5.3 (vs. 5.5-7.4 in other species); lower jaw strongly 
projecting beyond upper when mouth is closed, teeth very large, 
those of lower jaw extending to external anterior margin of 
dentary; barbels greatly reduced, length of maxillary and mental 
barbels about 0.5-1.0 eye diameter. 

Kryptopterus bicirrhis (Valenciennes, 1839) 

(Figure 113) 

Silunis bicirrhis Valenciennes m Cuvier and Valenciennes. 1839:367 (type locality 
Java). 

Silurus palemhangcnsis Bleekcr, 1852(/:584 (type locality Palembang, Sumatra). 

Krypwplenchthys palemhangcnsis Bleeker. 1 858c:290. 

Kryploplcrichlhys bicirrhis Bleeker. I858i:292. 

Cryplopterus amboinensis Giinther, 1864:40, 429 (type locality ".'\mboyna.'" er- 
roneous). 

Material Examined. — Malay Peninsula: Bentong. Pahang, 3: 89.4-104 mm 
(CAS-SU 31980). Western Borneo: Kapuas 1976-14, 89.7 mm (MZB 3652); Ka- 
puas 1976-19, 2: 96.1-99.3 mm (CAS 49399, MZB 3653); Kapuas 1976-33, 95.7 
mm (MZB 3654); Kapuas 1976-34, 3: 93.5-125 mm (MZB 3655, RMNH 28897. 
USNM 230294); Kapuas 1976-44, 4; 67.4-87,8 mm (BMNH 1982.3.29. 172, CAS 
49400. MZB 3656); Kapuas 1976-45. 80.3 mm (MZB 3657); Sintang, 3; 60.8- 
78.6 (AMNH 7815). Java: 96.3 mm (MNHN 9932. liolotype of .S'. bicirrhis). 
Locality unknown: Bleeker collection, 122 mm (BMNH 1863,12.4.101); Bleeker 
collection, 4: 108-143 mm (RMNH 6837, presumably including holotype of .S' 
palembangensis). "Amboyna," 90,3 mm (BMNH 1855.3.24.14, holotype oft; 
amboinensis). 

D\AaNosis. — Kryptopterus bicirrhis is a medium-sized species, 
attaining perhaps 200 mm, transparent in life, with a long max- 
illary barbel, mental barbels greatly reduced or absent, gill rakers 
16-21, branchiostegal rays 8-10, dorsal fin well developed, pec- 
toral-fin rays 1 1-13. pelvic-fin rays 6, anal-fin rays 59-68, and 
vertebrae 46-49. 

Meristic data on this species and its synonyms are presented 
in Table 10. 

Kryptopterus cryptopterus (Bleeker. 1851) 

.Silurus cryploplcnis Bleekcr. I851(.:270 (type locality Bandjermassing in fltiviis). 
Kryptopterus micropus Bleeker, 1858t:284 (unwarranted substitute name for Si- 
lurus cryploplcnis Bleeker. 1851). 
Cryptopterus micropus Giinther, 1864:42, 
Kryptopterus cryptopterus Fowler, 1905:468, 



Material Examined —Thailand: Chao Phrya at Bangkok, 1 15 mm (CAS-SU 
28829), Western Borneo: Kapuas 1976-14, 4; 98,7-124 mm (BMNH 1982,3,29,168, 
CAS 49393, MZB 3640); Kapuas 1976-19, 3: 1 1 7-1 38 mm (FMNH 94242, MNHN 
1982-708, MZB 3641): Kapuas 1976-22, 126 mm (MZB 3642); Kapuas 1976-33. 
9: I 14-146 mm (MZB 3643, RMNH 28893, UMMZ 209890, USNM 230289, 
K!_IMF2858). 

Distribution. — Laos, Thailand (Mekong. Chao Phrya). Ma- 
lay Peninsula. Sumatra (Palembang, Djambi, Banjuasin). Bor- 
neo (Baram. Sambas, Kapuas. Bandjermasin). 

Kryptopterus lais (Bleeker. 1851) 

Silurus lais Bleeker, I851/:428 (type locality .Sambas m (luviis). 
Krypiopterichthys lais Bleeker. 1 858i:29 1 . 

Cryptopterus lais Giinther. 1864:62; von Martens, 1876:399 (Danau Snang, Ka- 
puas basin). 

Material Examined. — Western Borneo; Kapuas 1976-33. 2: 59.3-65.5 mm 
(CAS 49394. MZB 3644); Kapuas 1976-55. 5: 55,8-75,7 mm (BMNH 

1982.3.29.169, MNHN 1982-709, MZB 3645, RMNH 28894, USNM 230290). 

Distribution, — Borneo (Sambas, Kapuas. Kahajan, Barito). 
Kryptopterus limpok (Bleeker. 1852) 

Silurus limpok Bleeker, l852£/;583 (type locality Palembang, in lluviis), 
Kryptopterus limpok Bleeker. 1858c;286, 
Cryptopterus limpok Giinther, 1864:39. 

Material Examined —Western Borneo: Pontianak. 126 mm (RMNH 7814); 
Sintang. 122 mm (RMNH 7815); Kapuas. Knapci ct Sebruang, 2: 217-225 mm 
(MNHN 1891/456-457); Kapuas 1976-19, 15; 130-241 mm (BMNH 

1982.3.29.170. CAS 49395. MZB 3646. RMNH 28895. UMMZ 209861, USNM 
2.30291); Kapuas 1976-38, 294 mm (MZB 3647), 

Diagnosis. — Knptopterus limpok is a moderately large species, 
attaining 300 mm or more, immediately distinguished from all 
other Kryptopterus by its elongate mental barbel extending pos- 
teriorly to or beyond tip of pectoral fin. 

Kapuas specimens of this species have gill rakers 1 5-20, bran- 
chiostegal rays 10-12, dorsal fin present, pectoral-fin rays 13- 
15, pelvic-fin rays usuallv 9, anal-fin rays 76-85, vertebrae 57- 
59. 

Distribution. — Thailand (Chao Phrya. Pasak). Malay Pen- 
insula (Pahang). Sumatra (BatangHari, Kwantan, Musi). Borneo 
(Baram, Kapuas, Kahajan, Mahakam). 

Kryptopterus macrocephalus (Bleeker. 1858) 

(Figure 1 14) 

Kryptopterichlhy. niacroccphalui Bleeker, 185Si:293 (type locality Sumatra?. Pa- 
dang'', in lluvus). 



148 



CALIFORNIA ACADEMY OF SCIENCES 





FiGiiRF 1 1 V Krvi'lopli-'rii^ huirrhi^ Above, Kapuas 1976-34, '>,3,.S mm (MZB ,36.S5); helow, Java, 96.9 mm (MNHN 9932, holotype, radiograph by Janme Abel), 



CrYpUiplcni\ iihhinct'pliulii^ Gunther, 1864:41 

Cryploplcrus hiciirhis Popta. 1906:26 (upper Mahakam). 

Krypliiplcnis DUurocfphuliLS Herre and Myers. 1937:67 (Malay Peninsula). 

'X'nptiiplciYlla hcldli Fowler, 1944:2 (type localny Borneo). 

M.MERiAL Examined. — Peninsular Thailand: Singora (=Songkhla), 62.2 mm 
(CAS-SU 14840). Malay Peninsula: Perak, Bukil Merah, 66.0 mm (CAS-SU 3107 1 ). 
Sumatra: no further locality, 3: 51.5-62.4 mm (C,AS-Sl! 31071). Sumatra?: Pa- 
dang':'. 96.5 mm (BMNH I 863. 12.4. 101, holotype'). Western Borneo: Kapuas 
1976-5. 4: 50.2-5X 2 mm (CAS 49396. MZB 3648); Kapuas 1976-8. 10: 33.8- 
50 4 mm (C.-^S 49397, MZB 3649, USNM 2.30292). 

Diagnosis. — Krypicpierus niacroccplialus is distinguished 
from almost all other Kryptopterm by its strikingly striped or 
(alternatively) mottled coloration; small size, largest known 
specimen about 100 mm; long maxillary barbel, gill rakers 1 1- 
13, dorsal tin present, pectoral-fin rays 9-10, pelvic-fin rays 6, 
anal-fin rays 48-55, vertebrae 42-45. 

Kryploplcnis macrocephaltis has two quite distinct color va- 
rieties, striped and mottled. The commonest variety, and the 
one exhibited by the holotype, is striped, with three well defined 
longitudinal dark stripes extending length of body. The middle 
stripe tends to be broadest and best defined; its dorsal margin 



is limited by a very dark, thin midaxial streak. In the mottled 
color variety the longitudinal stripes are present but are variably 
broken up, and the clear spaces separating them are obscured 
by mottling which may extend onto the anal fin. In both color 
varieties there is usually observed a concentration of melano- 
phores forming a small spot at the base of each anal-fin ray, and 
a concentration of melanophores forming continuous thin lon- 
gitudinal line or streak for the length of the anal fin about one- 
third of the distance from its distal margin. The dorsal surface 
of the head usually bears two dark broad longitudinal bars sep- 
arated by a light interspace; in some specimens in which the 
entire dorsal surface of the head is darkened these are obscured. 
In the field I was inclined to believe that the striped variety 
(Kapuas 1976-5) and mottled variety (Kapuas 1976-8) were 
different species, but detailed comparisons of their color pattern 
and morphology leads me to conclude they are conspecific. Me- 
ristic data on these color varieties are presented in Table 10. 

Crypiopierclla hcldli is based on an aquarium specimen from 
Borneo (ANSP 71571. 93 mm) which I have not examined. The 
color pattern, as figured by Fowler, includes several features 



ROBERTS-WESTERN BORNEO FISHES 



149 




Figure 1 14. Kivplopicrus macwccphahis. Above, Kapuas 1976-5, 54.4 mm (CAS 49396); below, Kapuas 1976-8, 50.4 mm (CAS 49397). 



here considered diagnostic of A', macrocephalus including lon- 
gitudinal stripes (albeit faint), a thin dark midaxial streak, dark 
spots along anal-fin base, a thin longitudinal line on anal fin 
one-third of distance from its distal margin, and very large round 
spots on dorsal surface of snout. The maxillary barbels of the 
specimen are abnormally short because their ends have been 
bitten offCW, Smith-Vaniz and E, Bohlke. pers. comm.). 

Distribution, — Peninsular Thailand (Songkhla). Malay Pen- 
insula (Perak, Johore), Sumatra (Padang. Djambi), Borneo (Ka- 
puas, Mahakam), 

Kryptopterus micronema (Bleeker. 1846) 

Sihinis micronemus Bleeker. 1846/':2S9 (type locality Batavia). 

Sihinis phalacrofioliis Bleeker, 1851(:429 (type locality Sambas, in fluviis). 

Mhwnema typiis Bleeker, 1 858i :30(3 (unwarranted replacement name lor Sihinis 

nucroiiciinis Bleeker, 1 846). 
I'halacronolus micniroplenis Bleeker, 1 858c:307 (unwarranted replacement name 

I'or Sihinis phalacrnnolus Bleeker, 1851). 
Micronema phalacronotiis Bleeker, 1859a: 139. 
Cryploplenis niicroncimi Giinther. 1864:43 



Kryploplcnis deigiiani Fowler. 1937:136 (type locality Mepoon. Central Siam). 
See Smith (1945:344). 

Material Examined. — Western Borneo: Kapuas 1976-33, 3: 125-225 mm (CAS 
49398. MZB 3650): Kapuas 1976-44. 4: 116-156 mm (BMNH 1982.3.29.171, 
MZB 365 I, RMNH 28896, USNM 230293): Kapuas, Knapei el Scbruang, 2: 291- 
316 mm (MNHN 1891/452^53). 

Distribution. — Thailand (Chao Phrya). Malay Peninsula 
(Johore. Singapore). Sumatra (Palembang). Borneo (Baram. 
Sambas. Kapuas, Kahajan). Java (Batavia). 



Kryptopterus minor new species 

(Figure 115) 

HoLOTYPE. — MZB 3638, 68.5 mm. mainstream ol" Sungai Pinoh at Nanga 
Saian, 45 km S of Nangapinoh (Kapuas 1976-29). 

Paratypes.-BMNH 1982.3.29.163-167, 4, CAS 49390, 4, FMNH 94241, 4, 
MNHN 1982-707, 4, MZB 3636, 13, RMNH 28892, 4, USNM 230288, 4, 41: 
49.8-68.0 mm, Sungai Landak at Ngabang (Kapuas 1 976-9): CAS 49391,2, MZB 
3637, I. UMMZ 209865, 4, 7: 46,3-55,2 mm, Sungai Mclawi and mouth ol' 
Sungai Pmoh at Nangapinoh (Kapuas 1976-22): CAS 49392, 7, MZB 3639, 1,8: 
24,2-34,4 mm, Kapuas mainstream at Selimbau (Kapuas 1976-55). 



150 



CALIFORNIA ACADEMY OF SCIENCES 




FitiURE I 15. kr\pii>[fti-nL\ minor kapuas 1^76-2'), 68.5 mm (MZB .^638, paralype). 



Diagnosis. — AVv/'/o/i/t'CHi' Dinior is a plain colored, small 
species (largest known specimen 68.5 mm), transparent in life, 
differing from all other Kryptoptcrus by having usually or always 
fewer gill rakers, branchiostegal rays, and pectoral- and pelvic- 
fin rays. Dorsal fin present; maxillary barbel elongate; mental 
barbels minute; gill rakers 10-13; pectoral-fin rays 8-9; pelvic 
5-6, anal 46-60; vertebrae 9-10 + 33-35=43-46, Frequencies 
of meristic characters are presented in Table 10. 

This species is perhaps most closely related to A', bicinius. 
with which 1 initially identified it. Although both inhabit the 
mainstream of the Kapuas River and its larger tributaries, and 
were collected mainly by night-lighting, they were not collected 
together on any occasion. They differ greatly in nearly all me- 
ristic characters (Table 10). 

So far as I have been able to determine this species has not 
been previously reported under any other name. The only spec- 
imens known to me are those obtained by the Kapuas survey 
of 1976. 

Etymology. — The name minor (Latin) refers to the small 
size of adults of this species. 

Kryptopterus schilbeides (Bleeker. 1858) 

llciiiisiliinis «7»//>0(/('v Bleckcr. lS58i;J''7 (iNpc locality Palcmbang. Sumalra, 

and Bandjermassin. Borneo). 
Crrplnpicrus schilheulcs Gunlher, 1 8b4;4 1 . 
Knpioplcrus schilhciili's Haig, 1952:110. 

M,4TERiAL Examined. — Western Borneo kapuas I'lVb-U, 2: 82.0-82 6 mm 
(BMNH I982..V2<).17.1, MZB ?.h5iiy. Kapuas 1976-.1.1. 8: 61.4-88.6 mm (CAS 
49401. MZB .^651. RMNH 288Q8. 1.IMMZ 209891. LISNM 230295). 

Distribution. — Sumatra (Palembang, Djambi), Borneo (Sa- 
rawak, Kapuas, Bandjermasin). 

Kryptopterus sp, undet. 

Cryp!:iplcrii\ itucraucma Vaillant, 1902:48 (Pontianak) 
CrvpU'plcnis apiifiiin Weber and dc Beaiirorl, I9LV22I (part) 

Material Examined —Western Borneo: Pontianak, 67.9 mm (RMNH 7817, 
idenlilied as C luicrnncma hy Vaillant 1902) 

1 have not been able to identify this distinctively colored 
specimen. It is certainly not K. Ducroncma as originally deter- 
mined by Vaillant (1902), nor is it A', apogon as suggested by 
Weber and de Beaufort; it differs from both of these species in 
having a dorsal fin (overlooked by Vaillant) as well as fewer gill 



rakers and anal-fin rays. Mouth broad, lower jaw prominent; 
teeth large, sharp; maxillary barbel extending posteriorly to mid- 
dle of eye (left side) or to its posterior border (right side); mental 
barbels absent'^; dorsal fin present (rays broken off near base); 
pectoral-fin branched rays 1 3-14; pelvic-fin rays 8?; anal-fin rays 
77? (73 reported by Vaillant); gill rakers 2+14=16; branchi- 
ostegal rays 14 or 1 5?; entire head, body, and anal fin strongly 
marbled. 



Ompok Lacepcde, 1803 

Oinptik LacepMe. 1803:49 (type species Ompok siluroulcs LacepCde. \iiOi=Silnrus 
hinuuiihilifs Bloch, I 797, by monotypy ). 

Callhhroiis Hamilton-Buchanan, l822:l49(typespecies.S7/H/H.s(C'i)//;i'/ir()i«)/)u^- 
iki Hamilton-Buchanan. 1822. by subsequent designation of Bleeker, 1862:17). 

S/liiroili's Bleeker. I858{:271 (type species Siliirus hypophlhalmus Bleeker. 1846. 
b\ subsequent designation of Bleeker, 1862:177). 

P\ciuiosi!uim Bleeker, 1858t:275 (type species Silunn himutululus Bloch, 1797, 
b\ present designation; n.b.: designation o( Psciiiiosiliinis as an absolute syn- 
onym of Calluhroui Hamilton-Buchanan. 1822 by Bleeker. 1862:17 is invalid 
because Siliiru^ (Callichrous) pabda is not among the species included in the 
original account o{ Pwuciosiliirui) 

Four species of Ompok occur in western Borneo. Meristic 
data on Bornean Ompok are given in Table 1 1. 

Ompok eugeneiatus (Vaillant, 1893) 

Calhchwus cugcimalus Vaillant. 1893u:6l (type locality Kapuas). 
Silurotic\ I'liffi'niiinis Weber and dc Beaufort. 1912:13. 
Ompok fi(i;i-iu-niliis Haig. 1952: 105 

Material Examined, — Western Borneo: Kapuas 1976-32, 9: 76.5-94.8 mm 
(BMNH 1982.3.29.174, CAS 49402, MNHN 1982-710. MZB 3661, UMMZ 
209K8I): Kapuas 1976-44. 3: 122-129 mm (MZB 3662, USNM 230296). 

Ompok hypophthalmus (Bleeker. 1846) 

Siliinis hypophlhulmiis Bleeker, I846i;:149 (type locality Batavia). 

Sihinis macwnema Bleeker. 1 85 l/r203 (type locality Bandjermassing, in lluviis). 

Sihirodcs hypophthalmus Bleeker, 1858i-:272. 

Siliirodes macroncma Bleeker, I858c:273. 

Callichwus hypophlhalmus Gunlher, 1864:48. 

Callichwus macroncma Gunther. 1864:49. 

Ompok hypophthalmus Haig. 1952 105. 

Material Examinei> —Western Borneo: Kapuas 1976-20. 3: 144-177 mm (CAS 
494113. MZB 3663): Kapuas 1976-32. 2: I I 3-125 mm (MZB 3664, RMNH 28899); 
Kapuas 1976-33. 190 mm (MZB 3665); Kapuas 1976-42, 140 mm (MZB 3666); 
Kapuas 1976-44. 2: 117-196 mm (MZB 3667, USNM 230297). 



ROBERTS- WESTERN BORNEO FISHES 



151 



Table 1 1. Meristic Characters of Bornean Ompok For definition of vertebral counts see p. 22 





Branchiostegal 
rays 


Gill rakers 


Pectoral- 
fin rays 


Pelvic-fin rays 


Anal- 
fin rays 


Vertebrae 


O. eugeneiatus 
Kapuas 


9-10 


17-19 


12-13 


7(21,8(1) 


61-62 


11-1-39^0=50(1), 51(2) 


O hypophihalmus 
Kapuas 


12-13 


14-15 


13-14 


8(1) 


80-84 


1 1 +44=55(2) 


O. leiacamhus 
Baram 




10 


13 




55 


94-35=44 


O. sabanus 
Kinabatangan 
Kapuas 


9-12 


16-20 
12-14 


11-12 


6(4), 7(1) 


57-64 
53-58 


ll-t-37-39=48(l), 49(4), 50(1) 
ll-h35-37=46(2), 47(5), 48(1) 


O. weberi 
Kapuas, Kumai 










41-47 





Ompok sabanus Inger and Chin, 1959 

(Figure 1 16) 

Ompok sabanus Inger and Chin, 1959:282 (type locality Segama River, north 
Borneo). 

Material Examined. — Western Borneo: Kapuas 1976-39, 23: 57.0-75.8 mm 
(BMNH 1982.3.29.175-176, CAS 49404. FMNH 94243, MZB 3668, RMNH 
28900). North Borneo: Kinabatangan basin, 6: 68.9-108 mm (CAS 49745, FMNH 
68024. paratypes). 

Distribution. — Kapuas. Kinabatangan (northeast Borneo). 
Ompok weberi (Hardenberg, 1936) 

Caliichrous weheri Hardenberg, 1936:232 (type locality Padang Tikarbay, western 

Borneo); Hardenberg, 1937:9 (Kumai River, western Borneo). 
Ompok wcben Haig, 1952:106. 

Material Examined. — None. 

Ompok weberi is known only from Hardenberg's descriptions 
of the 79 mm holotype from Padang Tikarbay and a 50 mm 
(total length) non-type specimen from the Kumai. Both speci- 
mens are presumed lost. 

As described by Hardenberg it differs from all other Ompok 
in having only 41-47 anal-fin rays (vs. 53 or more). Eye covered 
by skin. 4 in head; maxillary barbel extending posteriorly to 
beyond anal-fin origin; mental barbel very short, length less than 
that of head; branchiostegal rays 13; pectoral-fin rays 8-9; cau- 
dal-fin lobes rounded. 

Silurlchthys Bleeker, 1856 

Silunchlhys Bleeker. 1856:417.418 (lype species Silunis phamsoma Bleeker. 1851. 
by monotypy). 

This distinctive southeast Asian genus is poorly represented 
in museum collections, and it is doubtful how many species 
should be recognized or how they may be distinguished. There 
are three species in the Kapuas, including one new species. 
Meristic data on these three species are presented in Table 12. 

Silurichthys hasseiti Bleeker, 1858 

(Figure 117) 

Silunchlhys Hassehu Bleeker, 1858t:270 (type locality Java, Tjisekat, in fluviis). 

Material Examined. — Malay Peninsula: Kota Tinggi, Johore, 4: 44.7-99.0 mm 
(CAS-SU 31076, 32698, 39335); western Borneo: Kapuas 1976-6, 81.5 mm (MZB 
3669); Kapuas 1976-26. 83.3 mm (MZB 3670). 



Silurichthys phaiosoma (Bleeker, 1851) 

(Figure 1 18) 

Silunis phaiosoma Bleeker, I851;:428 (type locality Sambas, in fluviis). 
Silunchlhys phaiosoma Bleeker. 1858(-:268. 

Material Examined. — Malay Peninsula: Telok Anson. Pcrak (CAS-SU 31076); 
western Borneo: Kapuas 1976-8, 38.4 mm (MZB 3671); Kapuas 1976-42, 70.9 
mm (MZB 3672). 

Silurichthys sanguineus new species 

(Figure 1 19) 

Holotype. — MZB 3673. 40.9 mm. Sungai Tekam, a small forest stream, where 
It enters Kapuas mainstream about 5-6 km upstream from Sanggau. 

Diagnosis, — A Siliirichfhys lacking pelvic fins and with only 
7-1-4=11 principal caudal-fin rays (all other Silurichrhys ex- 
amined with 12-13 principal caudal-fin rays); head relatively 
shorter and body more elongate than in any other Silunchlhys; 
upper lip extending posteriorly to a point behind vertical along 
posterior margin of eye; dorsal-fin origin far in advance of a 
vertical through anal-fin origin; vertebrae 13 + 43=56; color in 
life entirely blood-red or vivid purplish-red. 

The holotype and only known specimen of 5. sanguineus may 
be further characterized as follows: dorsal-fin rays 4; pectoral- 
fin rays 18; anal-fin rays 60; head length 6.3; maxillary barbel 
extending posteriorly to well beyond anal-fin origin, mental bar- 
bel to well beyond pectoral-fin origin; greatest body depth (at 
abdomen) 7.4; body depth at middle of anal fin 9.9; depth of 
anal fin 19.5; upper caudal-fin lobe distinctly longer than lower 
lobe. Specimens of 5. hasseltii and S. phaiosoma of comparable 
size observed in life were plain colored or mottled brownish 
without any coloration of the vivid red color observed in 5. 
sanguineus. 

Etymology. — The name sanguineus (Latin) refers to the 
blood-red color of the holotype of this species observed in life. 

VVallago Bleeker, 1851 

U'allago Bleeker. 1851c:265 (type species Silurus nnillcn Bleeker, lS46=Silunis 
aim Bloch and Schneider, 1801, by subsequent designation of Bleeker, 1862a: 
394). 

li'dllagonia Myers. 1938:98 (type species II V;//a,?o /«'n; Bleeker, 1851, by original 
designation). 

IVallago comprises large silurids, distinguished from all other 
genera in southeast Asia by having a relatively large dorsal fin 



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Figure 1 16. Ompok sahaiuis Above, Kapuas l'^76-39. Ti.O mm (FMNH 'J4243), below, nonhcaslem Borneo, KLnabalangan, 76.3 mm (FMNH 68024, paratype). 



with 5 rays (dorsal fin smaller, usually with 4 or fewer rays, or 
absent altogether, in other southeast Asian silurids). Eye with 
free orbital rim, lying entirely above level of corner of mouth, 
not visible from beneath head (eye subcutaneous in all other 
southeast Asian silurids, lying at or below corner of mouth in 
Ouipok. visible from below head in Krypioptcrus and most other 
silurids); gape oblique, wide, and very long, reaching to or be- 
yond anterior margin of eye (shorter in Onipok and most other 
silurids); branchiostegal rays 13-21; pectoral-fin rays 12-15, 
pelvic 8-1 1. anal 60-96; caudal fin deeply forked, with slightly 
pointed lobes, free from anal fin; principal caudal-fin rays 8/9. 
Three species of Wallago are recognized (Roberts 1982(), of 
which only one is found in western Borneo. 

VVallago leerii Bleeker, 1851 

WalUiiin Lcvni Bleeker. IS."! 1/427 (type loealily Samhabet Paicmbang. m lliiviis) 
H'lillufiii m7i/(/('.v»,v Vaillanl. 1402 46 (l\pe kualiu Tepoe. bords du Mahakam). 

.See Roberts (1<J82(1. 
'^WalUjgonia Iwccdia Hora and Misra /// Hora and Gupla, 144 1 : i S (t>pe loeality 
Kuala Tahan. Pahang) See Roberts (14S2i ) 

Material ExAMiNEi:) —MalaN Peninsiila Singapore, .10s mm (( ,-\S-Sl) 148.1M) 
Sumatra: Djambi. 21: I02-.31S mm (ZMA 114.4U7). Borneo: Pleyhane Ri\er, 
21S mm (RMNH \imy. Kapuas 1976-16. 2: I 7>^-202 mm (CAS 4940.S, MZB 
.3674), Kapuas 1176-46, 175 mm (MZB .'(675), Kapuas River, .321 mm (ZMA 
I 14. 411(1): Mahakam River. I.s2 mm (RMNH 7X12. hololype of If iichiildsiis). 

D[AaNos\s. — Wallago leerii is distinguished from all other 
\\'allag< I hy its jet black pectoral-fin membranes. Branchiostegal 
rays 15-18; gill rakers 12-16; anal-fin rays 64-75. 

Local fishermen interviewed in 1976 indicated that Wallago 
leeni (always known to them as ikan tapah, and not confused 
with any other species) attains 80 kg and is the largest fish in 



the Kapuas. They also stated that it used to form large migratory 
schools in the Kapuas mainstream but that such schools had 
become less noticeable, which they attributed to intensive gill 
netting, 

Belonidae 

The predominantly marine and estuarine needlefishes or Be- 
lonidae include a single freshwater genus in Asia, 

Xenentodon Regan, 1911 

Xi'nt'nloJon Regan, 191 1:332 (type species liclonc cuncila Hamilton-Buchanan, 
1S22. b\ suhsetiuenl designation or.lordan. I920:S40). 

Body cylindrical; jaws with large canine teeth alternating with 
much more numerous small conical teeth; infrapharyngobran- 
chial bones absent or toothless except for third pair (infrapha- 
ryngobranchials two through four present and tooth-bearing in 
all or almost all other Belonidae except the diminutive South 
•American Behniioir. Collette 1966); gill rakers absent; scales 
very small to extremely small; lateral line placed very low on 
body, not forming keel on caudal peduncle; dorsal and anal fins 
about equal and nearly opposite each other, with 14-19 rays; 
caudal fin truncate or rounded; vertebrae 33-40 + 21-23=55- 
62. 

The genus is widely distributed in fresh water (and brackish 
water?) in India, Burma, Thailand, the Malay Peninsula, Su- 
matra, and western Borneo but apparently is absent from north- 
ern and eastern Borneo and Java. Only two species currently 
are recognized but my observations indicate there are at least 
three. 



ROBERTS-WESTERN BORNEO FISHES 

Table 1 2. Meristic Characters in Silurichthys For definition of vertebral counts see p. 



153 



Anal-fin rays 



Principal caudal-fin rays 



Vertebrae 



S. hasseln 
Kota Tinggi 
Gunong Pulai 
Singapore 
Kapuas 

S. phaiosoma 
Telok Anson 
Kapuas 

S'. sanguineus 
Kapuas 



50-56 

4S 

52 
55-56 



55 

49, 56 



60 



7 + 5(3), 6 + 6(1) 
7 + 6? 

7 + 6(1) 



7 + 6(1) 



7 + 4 



10-1 1+35-37=46(1), 47(2), 48(1) 

11 + 36=47 

11 + 35=46 

10'' + 36-39=46?(l), 49?(1) 

11 + 38=49 
11+38^0=49(1), 51(1) 

13 + 43=56 



Key to Xenentodon 

la Dorsal-fin rays 16-18, anal-fin rays 17-19; predorsal scales 
more than 200, transverse scales (between dorsal- and 
anal-fin origins) 30-40; vertebrae 57-62 (small-scaled 
species) 2 

lb Dorsal- and anal-fin rays 14-17; predorsal scales about 
140-160, transverse scales 20-30; vertebrae 55-58 (large- 
scaled species) Xenentodon sp. or spp. 

2a Jaws relatively large and heavy-set. length of upper jaw 
2.6-3.2 times in body length measured from anterior mar- 
gin of orbit to end of hypural plate; enlarged teeth in 
upper jaw 9-21 (Fig. 120); dorsal-fin origin usually an- 
terior to a vertical through anal-fin origin; a heavy-bodied 
species reaching 40 cm in total length .\'. cancila 

2b Jaws relatively slender and delicate, length of upper jaw 
2.3-2.6 times in body length; enlarged teeth in upper jaw 
21-32 (Fig. 120); dorsal-fin origin usually over base of 
second or third anal-fin ray; a smaller, more slender species 
apparently not exceeding 30 cm in total length 

X. canciloides 

The large-scaled species of Xenentodon. of which there may 
be more than one. occurs in southern India, Sri Lanka, and the 
Malay Peninsula, probably in brackish as well as fresh water. It 
apparently does not exceed about 30 cm total length. The small- 
scaled A', cancila appears to be the only species of the genus in 
the Ganges-Brahmaputra. It may also be widely distributed in 



Burma and Thailand but this requires further study. Although 
a large-scaled species and X. canciloides both occur in the Malay 
Peninsula, onlv the latter is known from western Borneo. 



Xenentodon canciloides (Bleeker, 1853) 

Belone canciloides Bleeker. 1 853i :454 (type locality Pontianak. in llumine Kapuas; 

Pangahocang, provinciae Lampong. Sumatrae austro-onentalis, in tluviis). 
Mu'.kicemheliis cancilmdes Bleeker. 1866:223. 

Belone cancila von Martens. 1876:400 (Kapuas); Vaillant, 1902:31 (Kapuas). 
Xenentodon canciloides Weber and dc Beaufort, 1922:133, 

Material Examined. — Malay Peninsula: Johore, Kota Tinggi, 2; 98.5-123 mm 
(CAS-SU 32881 . 33862). Western Borneo: Kapuas 1976-8, 3: 108-245 mm (CAS 
49433. MZB 3735): Kapuas 1976-9, 148 mm (MZB 3736); Kapuas 1976-22. 4; 
62.4-128 mm (FMNH 94246, MZB 3737); Kapuas 1976-32. 6: 147-201 mm 
(MZB 3738. RMNH 28911. USNM 230315); Kapuas 1976-33. 161 mm (MZB 
3739); Kapuas 1976-37. 174 mm (MZB 3740); Kapuas 1976-39.2: 170-194 mm 
(BMNH 1982.3.29,203. MZB 3741); Kapuas 1976-42. 153 mm (MZB 3742) 
Kapuas 1976-44. 151 mm (MZB 3743); Kapuas 1976-46. 160 mm (MZB 3744) 
Kapuas 1976-47. 2: 69,2-103 mm (CAS 49434. MZB 3745); Kapuas 1976-51. 2 
168-180 mm (MNHN 1^82-715, MZB 3746). 

Distribution. — Malay Peninsula (Johore). Sumatra. Western 
Borneo (Kapuas). 



Hemiramphidae 

The circumtropical Hemiramphidae or halfbeaks comprise 
about 12 genera and 80 species (Nelson 1984). Three genera in 




Figure 117. Silimchlhvs Imsselti. Kapuas 1976-26. 83,3 mm (MZB 3670), 



154 



CALIFORNIA ACADEMY OF SCIENCES 



"*^yA'^.- 









Figure 1 18. Siliinchlhys phaiosoma Sumatra (after Weber and do Beaufort \'^\'S. fig. 77). 



southeast Asia (including Celebes) are viviparous and occur 
largely or exclusively in fresh water, two in western Borneo. 

Dermogenys van Hasselt, 1823 

Dennogenys van Hasselt in Kuhl and van Hasselt. 1823:131 (type species Der- 
mogenys pusillw, van Hasselt in Ruhl and van Hasselt, 1823. hy monotypy). 

Dermogenys. characteristic of brackish as well as freshwater 
habitats, is the most widely distributed of the viviparous south- 
east Asian halfbeaks, ranging from Burma and Thailand along 
the coasts of the Malay Peninsula and Greater Sunda islands to 
Celebes. Although Dermogenys occuv% far inland in some places, 
and its distribution surrounds that of Hcmlrhamphodon. the 
two genera apparently never occur sympatrically. The single 
collection of Dernu^genys taken by the Kapuas survey of 1976 
was obtained in a tidal creek near the Kapuas mouth far from 
any collections of Hemirhamphodon. Dermogenys is unknown 
from the Kapuas interior. I suspect that wherever it occurs the 
exclusively freshwater Hemirhamphodon has excluded or dis- 
placed Dermogenys. 

Dermogenys cf pusillus van Hasselt //; Kuhl and van Hasselt. 

1823 

Derniogenw^ pii.\illiis van Hasselt /// Kuhl and \an Hasselt, 1823: 1 3 1 (type locality 
Java). 

M,*TERiAL Examined. — Western Borneo: Kapuas l')7b-3. 6: 23.7-35.9 mm (CAS 
4Q428. MZB 3717. ZMA 1 16.544). 

Mohr (1936) revised Dermogenys. giving an extensive syn- 
onymy of D. piisillu.s and recording specimens of the species 



from numerous localities virtually coextensive with the entire 
range of the genus. Until Dermogenys can be revised again these 
records should be regarded with caution. 

Hemirhamphodon Bleeker. 1866 

Heinirlmitiphi'don Bleeker. 1866a: 140 (type species Hemirhamphus phaiosoma 
Blcckcr, 1852, by original designation). 

Di.^GNOsis. — Myrmecophagous halfbeaks with each side of 
beak (extended lower jaw) with two or three rows of anterolat- 
erally directed conical teeth (beak toothless in all other hemi- 
ramphids). Beak terminating in an elongate, flexible, fleshy ap- 
pendage. Scales in lateral line series about 90-100. Dorsal fin 
with 14-24 rays, its origin far anterior to that of anal fin, Ovo- 
viviparous. Males with an elongate genital papilla and distinc- 
tively modified anal fin with posteriormost four rays separated 
from anterior rays and morphologically specialized. 

Male Hemirhamphodon may have 1-3 minute spines pro- 
jecting laterally from the posterior margin of the scales over 
most of their body surface. These are relatively smaller but 
otherwise probably comparable to the scale spines or ctenii on 
males of the South American freshwater belonid Pseiidotylo- 
siiriis angiisiiceps (see Wiley and Collette 1970:189, fig, lOf) 
and numerous cyprinodonts. Such "breeding contact organs" 
have not been reported previously in hemiramphids. 

Key to Hemirhamphodon of western Borneo 

la Dorsal-fin origin considerably posterior to vertical line 
through anal-fin origin; dorsal-fin rays 14-15 
H. pogonognathiis 




Figure 119. .Siliiriehlhys iangmiieiis Kapuas 1976-16. 40,9 mm (MB 3673, holotype). 



ROBERTS- WESTERN BORNEO FISHES 



155 




CS 



"^Ticzrrx^:! 



X. cancila 




X. canciloides 

Figure 120. Xenenlodon. Above, A cancila. Nepal, Chitawan Valley, 1 1 1 mm (CAS 50358); below, \ canciloides. Rapuas W76-22, 1 12 mm (FMNH 94246), 



lb 



Ic 



Dorsal-fin origin considerably anterior to vertical line 
through anal-fin origin: dorsal-fin rays 22-24 

H. phaiosoma 

Dorsal-fin origin almost directly above or slightly pos- 
terior to a vertical line drawn through anal-fin origin; 
dorsal-fin rays 17-18 H. sp. undet. 



Hemirhamphodon phaiosoma (Bleeker. 1852) 

Hemirhamplius phanKoma Bleeker, 1852i;:26 (lype locality Billiton). 
Hemirhamphodon phaiosoma Bleeker, 1866u:168. 

Material Examined. — Western Borneo: Kapuas 1976-6, l.V 26.5-50.4 mm 
(CAS 49429, MZB 37 1 8), Rapuas 1976-1 2. 8; 37, 1-49.2 mm (MZB 37 1 9, USNM 
2303 1 3); Kapuas 1 976-25, 6: 32.0-47.5 mm (MZB 3720, RMNH 28909); Kapuas 
1976-26, 5: 26,9-48,6 mm (BMNH 1982.3.29.194-197, MZB 3721); Kapuas 
1976-30, 2: 41.0-47.9 mm (MZB 3722. UMMZ 209878). 

CoLORATiON. — Life colors of//, phaiosoma were recorded in 
the field at Kapuas 1976-12: overall body color violaceous or 
mauve: larger specimens usually with two (sometimes three) 
parallel longitudinal red streaks: smallest specimens with a sin- 
gle red streak: ventrolateral half of body without oval marks or 
blotches: dorsal, anal, and caudal fins, and gular membrane 



90° 



100° 



110° 



120° 



130° 




-20° 



-10° 



• phaiosoma 
A pogonognathus 



^^.- 



-10° 



Figure 121. Hentiramphodon. Geographical distribution of two species. 



yellow with thin red margins; abdomen pale, but in some spec- 
imens with two widely spaced parallel longitudinal red streaks 
extending from just behind head to anal fin origin. The only 
other color observation for this species known to me, of spec- 
imens from Billiton Island (de Beaufort 1939:192) agrees with 
mine except that he noted sexual dichromatism: "Two males, 
76 and 80 mm. and two females 68 and 69 mm [presumably 
total length]. Color of preserved specimens yellowish. The males 
with two narrow crimson lines, running parallel to each other 
along sides of body and two, the females with a faint pinkish 
lateral band, bordered above and below with dusky." 

Habitat, — In the Kapuas H. phaiosoma was found in smaller, 
relatively high-gradient, deeply shaded forest streams with nu- 
merous rocks and stones on the bottom. 

Distribution (Fig. 121). — Borneo (Palandok, Sadong Ma- 
tang. Kapuas). Banka. Billiton. The species has been reported 
from Singapore but I agree with Alfred (1966) that this record 
is suspect. There are no other records from the Malay Peninsula. 

Hemirhamphodon pogonognathus (Bleeker, 1853) 

(Figure 122) 

licinirhaniphns poi^onogmithiis Bleeker, IS53i/;193 (lype locality Banka Is.. Ma- 

rawang, in fluviis). 
Hcinirhainphodon pogonogiiarhii^ Bleeker. 1866^^169. 

'Hemirhamphodon kukenlhali Steindachner, 1 90 1 :450 (lype locality Baram Riv- 
er, Sarawak). See Mohr (1936:60-61). 

Material E.xamined— Malay Peninsula: Perak, Tapah Fisheries Station, 47.5 
mm (CAS-SH 40484); Johore, Kulai, 1 1 : 24,4-50,9 mm (CAS-SU 394 1 7); Johore, 
Gunong Pulai. 64: 22,2-52.5 mm (CAS-SU 30757); Johore, Mawai district 50- 
60 mi N of Singapore, 6; 30.2-44.1 mm (CAS-SU 30755); Johore, Kota Tinggi, 
1 1: 23,7-49.5 mm (CAS-SU 33619); Singapore, Mandai Road, 97: 29,0-56,4 mm 
(CAS-SU 33947, 33618); Singapore, 34: 26.5-49.7 mm (CAS 19994, CAS-SU 
30756. Sarawak: 15-20 mi E of Kuching, 24: 27.6-63,3 mm (CAS-SU 33564), 
Western Borneo: Kapuas 1976-5, 9; 25,9-55,0 mm (CAS 49430, MZB 3723); 
Kapuas 1976-6. 37.5 mm (MZB 3724); Kapuas 1976-8. 6: 26,6-49.7 mm (BMNH 
1982.3.29. 198-202, MZB 3725); Kapuas 1976-10,47,1 mm (MZB 3726), Kapuas 
1976-1 I, 2: 25.5-25.6 mm (CAS 49513, MZB 3727); Kapuas 1976-16, 35.5 mm 
(MZB 3728); Kapuas 1976-17, 7: 28.0-50.6 mm (MZB 3729. RMNH 28910); 
Kapuas 1976-367, 3; 31.6-43.2 mm (MZB 3730, UMMZ 209896); Kapuas 1976- 
3Q. 9: 17.9-44.3 mm (MZB 3731, USNM 230314); Kapuas 1976-46, 3: 24.0- 
25.0 (CAS 49431, MZB 3732); Kapuas 1976-51, 3: 29.7-36.5 mm (MZB 3733, 
ZMA 116.545). 

Coloration, — Life colors of//, pogonognathus were record- 
ed in the field at Kapuas 1 976-8, Overall body color pale bluish 
or violaceous. No midlateral longitudinal red streak. Ventro- 



156 



CALIFORNIA ACADEMY OF SCIENCES 





i-f*. ■ ■ ■-■ - ■ 



Figure 122. Hemirhamphodnn pogonognalhus. Above. Kapuas 1976-5. 55.0 mm male (CAS 4Q430); below. Singapore. Mandai Road. 54 mm male with upper 
jaw iransli,\cd by mandibles of 7.5 mm ant (CAS-SU 33947). 



lateral half of body with a row of four evenly spaced, bluish or 
bluish-green, vertically elongate oval spots or blotches from just 
behind pectoral fin to just in front of anal-fin origin (spot in 
front of anal fin sometimes faint). Abdomen white; no abdom- 
inal red streaks. Fins colorless or dusky e.xcept for white or pale 
blue margin of anal tin. Gular membrane red. Color photographs 
of the species have been published by Brembach ( I978«), Wick- 
man (1981). and Hyrtl (1983). The colorations illustrated agree 
generally with my observations but there are some difTerences 
worth noting. Wickman's fish was collected near Kuching; its 
coloration is basically similar to that described above except 
the overall body coloration is brownish or reddish-brown and 
the vertically oval spots dusky or dark; a thin midlateral lon- 
gitudinal reddish-brown streak present, scarcely distinguishable 
from overall brownish or reddish-brown body color. Caudal fin 
with a thin marginal coloration of pale blue and reddish flecks, 
otherwise plain. Opercle with two short, horizontal iridescent 
blue marks; gular membrane pale bluish with variably red mar- 
gins. Brembach's specimens and Hyrtl's specimen are without 
locality data. Brembach's have overall body color iridescent 
greenish dorsolatcrally and violaceous or pearly ventrolaterally; 
a thin, faintly visible midlateral longitudinal red streak; four 
bluish oval blotches; dorsal, anal, and caudal fins with thin pale 
bluish border, that of dorsal fin also flecked with red; gular 
membrane rosy, ventral border with a thin black margin. Hyrtl's 



specimen, a male, is strikingly beautiful: overall body color deep 
blue, somewhat violaceous ventrolaterally; midlateral red streak 
relatively well defined; four oval blotches rather indistinct, al- 
most same deep blue hue as overall body color; dorsal fin with 
thin red marginal band; pelvic, anal, and caudal fins with pale 
bluish margins; fins otherwise colorless; gular membrane en- 
tirely bright scarlet. 

Reproduction. — It has long been known that Hemirhaiii- 
phodoii is viviparous (or ovoviviparous), producing relatively 
large young, but there has been little documentation of the phe- 
nomenon. Soong ( 1 968). in an extremely brief communication, 
reported that in //. pnf;oiiognathtis: 1 ) males grow larger than 
females; 2) young are born in smaller batches than in Der- 
Diogcnys piisilliis. at an average (total?) length of 1 1 mm; 3) 
duration of gestation is about 6-8 weeks; 4) individual wild 
populations breed synchronously but diflerent populations breed 
at dilTerent times; and 5) there is no true (reproductive) season- 
ality. Brembach (1978(;) reported or indicated that; 6) pregnant 
females carry a number of young in diflferent stages of devel- 
opment; 7) young are bom in rapid succession, sometimes daily; 
8) the number of young produced is usually only 1 or 2 and 
sometimes 4 daily; 9) isolated females will generate young daily 
for a considerable period of time. My own observations on 
superfetating H. pogonognathus do not bear on items 3-4 but 
confirm or support 1-2 and 5-9. Data on males and females in 



ROBERTS-WESTERN BORNEO FISHES 



157 



Sarawak, Kuching 
16 Feb. 1937 
n:24 



R 




i 



H 



n 



Singapore Isd.. Mandai Road 

27 Feb. 1937 

n:41 




nnn 



Singapore Isd.. Mandai Road 

14 May 1937 

n:56 




Malay P.. Johore. Gunong Pulai 

August 1934 

n:64 




mm SL 



non-pregnant female ^M pregnant female I | male 

Figure 123. )lcmirliamphodon pogonognathus Length-frequency dislnbution of males, non-pregnanl females, and pregnant females from four population samples. 



D 



four population samples of H. pogonognathus are presented in 
Figure 123. These population samples were collected in Feb- 
ruary, May, and August. Smaller samples examined were col- 
lected in January. March, July, and October. Due to the early 
appearance of an elongate genital papilla in males and a swollen 
genital region in females it is possible to determine the sex of 
nearly all wild-caught specimens in my samples, including the 
smallest which are only 22 mm. In the four largest samples the 
male: female ratios are 12:12, 22:19, 27:29, and 30:34 or vir- 
tually 1:1. Males grow substantially larger than females. In three 
of four large samples the largest males are 9-12 mm longer than 
the largest females; 29 of the 3 1 largest specimens in the four 
samples are males. The largest male is 63 mm and the largest 
female only 52 mm. Females become sexually mature by 34 
mm; in all samples examined, most females above 34 mm are 
pregnant (60 out of 70 in four large samples) and almost all 



females above 38 mm are pregnant (49 of 52). From this ob- 
servation it follows that reproduction occurs throughout the 
year; i.e., that it is basically not seasonal. 

The developing young or fetuses are generally of about the 
same number and stage of development in the left and right 
ovanes. The following observations are based on the right ovary. 
The number of young in a single ovary varies from one to 12, 
but is usually 4-9. In addition there may be one to 10 or more 
unfertilized eggs or eggs to 2 mm in diameter in an early stage 
of development. The developing young are aligned in single file, 
each tightly wrapped around its yolk sac. Size at hatching is 
about 7.3-8,3 mm (the largest fetal young observed; in the small- 
est pregnant females the largest young are about 1 mm smaller 
than in the largest females). Successive young generally differ 
greatly in size. Standard length in mm of successive fetuses from 
the right ovary of several pregnant females is as follows: 37.4 



158 



CALIFORNIA ACADEMY OF SCIENCES 



90° 100° 110° 120° 130° 




o boa la 

• deokhatoides 
m heterosoma 

* martensii 



^^^^, 






-10° 



Figure 124, Doryuhlhys. Geographical distribution. 



mm female, 12 fetuses (CAS-SU 30757): 7.3, 7.3, 6.2, 5.1, 4.8, 
4.8. 4.5, 3,6, 3.6. 2.8, 2.4 (plus 4 eggs 1.2-1,6 mm in early 
development''); 37,2 mm female, 5 fetuses (CAS-SU 33618): 
6.7, 6.1, 4.2. 3.5, 1.1+2 eggs 1.1 mm in diameter (mimaturc?); 
41.3 mm female, 6 fetuses (CAS-SU 33618): 7.1, 6.2, 5.8, 4,6, 
2.7, 1.5 mm + 7 eggs 1.2-1.6 mm in a double row; 44,5 mm 
female, 7 fetuses (CAS-SU 33618): 7.7, 6.8. 5.2. 4.8, 3.0, 2,5 
mm + 1 egg 1.7 mm, with late cell divisions (no other eggs); 
5 1 .5 mm female. 1 1 fetuses (CAS-SU 30757): egg 2. 1 mm (mis- 
shapen, with small white core; evidently in process of resorp- 
tion), fetuses 8,3, 7.2 mm, egg 2.0 mm (very misshapen, large 
white core), fetuses 5.2, 5.4, 4,5, 4.5. 4.1. 3.4, 3,6. 2.8, 2.8 
mm + double row of 9 eggs (6 with morula- or gastrula-like 
development). From these and other observations I conclude 
that H. pogonoguaihus exhibits an extreme form of superfeta- 
tion, with each successive brood in a single ovary consisting of 
no more than three young and usually of only a single young. 
The difference in age of the young probably is due to individual 
fertili/alion of eggs by sperm stored in the female. An alternative 
hypothesis, that differences in fetal development arc controlled 
by a maternal or fetal hormonal gradient in the ovary, is con- 
traindicated by the observation of successive fetuses of identical 
length in pregnant females from Gunong Pulai. 

In summary, data presented here on H. pogonognathus in- 
dicate that females become sexually active at about 34 mm, and 
that from then on they produce very small numbers of relatively 
large young (7-8 mm) at more or less regular intervals through- 
out the year. A pregnant female may carry up to 8 discrete broods 
at once. If the gestation period of 6-8 weeks gi\en by Soong is 
correct, females presumably give birth at intervals as short as 
2-8 days. 

Food Habits. — Gut contents were examined in over 30 spec- 
imens representing nearly all available collections of//, pogo- 
nogihithus. Virtually every specimen contained at least one dis- 
crete food item. Almost all food items were terrestrial insects, 
sometimes too digested to be identified even to order. Never- 
theless, it is clear that H. pogonognathus feeds almost exclusively 
on terrestrial insects and predominantly ants. In more than half 



of the specimens examined ants were the only identifiable gut 
contents. Several specimens have 5-6 or more ants in their guts 
and no other items. One specimen, a 54.0 mm male (Fig. 122) 
has its upper jaw transfixed by the mandibles of a 7.5 mm ant 
it had tried to swallow. Other insects encountered (in small 
numbers, usually singly) include Diptera, Coleoptera. and Or- 
thoptera (mole-cricket). Food habits of the other species of 
Ilcmuiiamphodon apparently are comparable to those of H. 
pogonognathus. It seems likely that the rows of antrorse conical 
teeth unique to the beak of Hcnurhaniphodon enable it to snag 
the legs of live ants which are its predominant food source. 

Habitat. — In the Kapuas basin //. pogonognathus was found 
in small low-gradient streams with abundant logs and leaf litter 
but no rocks or stones on the bottom. 

Distribution (Fig, 121). — Malay Peninsula (Perak, Johore, 
Singapore). Sumatra (Palembang, Taluk, Gunong Sahilan, Sim- 
bolong mountains). Borneo (Baram'', Kapuas). Banka. Billiton. 

Hemirhamphodon sp. undet. 

IHcmirhamphodun chrYsnpunctaiu\ Brenibach. lQ78/i (type locality southeast 
Borneo). 

Material E.xamined. — Western Borneo: Kapuas I^76-I.V 5: 40.1—46-8 mm 
(CAS 49432. MZB 3734), 

A single small lot oi' Hcnurhaniphodon obtained in 1976 is 
not identifiable with either H. phaiosoma or H. pogonognathus 
(see Key above). It may be H. chrysopunctatus or an undescribed 
species. Color in life not recorded. 

Syngnathidae 

Syngnathidae (pipefishes and seahorses) are predominantly 
marine and brackish water inhabitants but endemic freshwater 
species occur in tropical west .Africa, southeast Asia, and perhaps 
elsewhere. The southeast .Asian freshwater genus Doryichthys 
comprises four species, all present in western Borneo. Their 
geographical distribution is shown in Figure 124. 



Doryichthys Kaup. 1856 

Dcrvhiiihv^ Kaup. 1853233 (nomen nudum) 

lli'iviiiuhys Kaup. I85(i:56 (t>pc species Dnrvuhlhys hihiicaliis Kaup. 

l8i(>=Syngihilluis dcukhuloulvs Bleekcr, 1853; h\ subsequent designation of 

Jordan and Evermann. 1896:773), 
Ktiiiiva Smith. I 963:533 (type species .9r/ii;/;i;?/!Ki hnuiu Bleeker. 1 85 I. by onginal 

designation) 

DiAONosis. — Opercle with a single longitudinal ridge; superior 
trunk ridge and inferior tail ridges not confluent; caudal-fin rays 
typically 9. .Apparently confined to fresh water. 

Doryichthys obtained during the Kapuas survey of 1 976 have 
been identified by C. E. Dawson, and some of the specimens 
utilized in his revision ofthe genus (Dawson 1981). The present 
account is based on this revision, to which the reader may refer 
for fuller information. 

Key to Doryichthys (after Dawson 1981) 

la Trunk rings 1 5-20; total rings 45-53; dorsal-fin rays 27- 
39; principal ridges of each ring without a prominent spine 



lb Trunk rings 22-26; total rings 56-64; dorsal-fin rays 43- 



ROBERTS-WESTERN BORNEO FISHES 



159 



69; principal ridges of each ring with a prominent spine 
in aduhs and subadults 3 

2a Trunk rings 17-20; tail rings 28-34; pectoral-fin rays 19- 
23; usually with large dark spots above lateral trunk ridge 
D. deokhatoides 

2b Trunk rings 15-17; tail rings 31-37; pectoral-fin rays 16- 
22 (usually 19 or less); usually with small dark spots on 
lateral trunk ridges D. martensii 

3a Trunk rings 22-24; dorsal-fin rays 43-65 D. boaja 

3b Trunk rings 25-26; dorsal-fin rays 65-69 D. heterosoma 

Doryichthys boaja (Bleeker, 1851) 

Syngnalhus boaja Bleeker. 185 la: 16 (type localilv Banjermassmg. in tluviis)- 
Doryichlhvs spinosiis Kaup, 1856:57, 75 (type locality Java, Borneo). 
Syngnalhus Jullieni Sauvage: 1874:338 (type locality Cochin China). 
Syngnalhus zonalus Karoli. 1882:185 (type locality Borneo). 
Doryichrhys helewsoma Vaillant. 1902 (misidcntification of specimen from Sin- 
tang). See Dawson (1981:16). 

Material Examined. — Western Borneo: Kapuas 1976-16, 191 mm(MZB3002): 
Kapuas 1976-40.338 mm(MZB 3003). These specimens arc illustrated by Dawson 
(1981, fig, 6). 

Distribution (Fig. 124). — Vietnam. Thailand (Mekong. Chao 
Phrya, Paltani). Sumatra. Borneo (Kapuas). Java (Brantas). 

Doryichthys deokhatoides (Bleeker, 1853) 

Syngnalhus deokhatoides Bleeker. I853g:5, 11.18 (type locality Palenibang, Su- 
matra, and Pontianak, Borneo). 
Doryichlhys deokhatoides Bleeker. 1859^:188. 

Material Examined. — Western Borneo: Kapuas 1976-3. 157 mm(MZB 3747): 
Kapuas 1976-16. 3: 112-136 mm (C'^S 49435. MZB 3748. USNM 218069): 
Kapuas 1976-17. 2: 114-131 mm (MZB 3749. USNM 218070); Kapuas 1976- 
51. 106 mm (MZB 3750). 

Distribution (Fig. 124). — Peninsular Thailand. Malay Pen- 
insula. Sumatra. Western Borneo (Kapuas). 

Doryichthys heterosoma (Bleeker, 1851) 

Syngnalhus heterosoma Bleeker. 1851/441 (type locality Sambas River, western 

Borneo). 
Doryichthys heterosoma Gunther. 1870:180. 

Material Examined. — None. 

This species is known only from the Sambas River and the 
Natunas islands. 



Doryichthys martensii (Peters, 1869) 

Syngnalhus martensii Peters. 1869:459 (type locality Pulo Matjan, Kapuas). 
IMicrophis caudatus Vaillant. 1893:62 (not of Peters, 1869; Kapuas). 
Microphis ignoralus Vaillant, 1902:28, 40. fig. 1.2 (type locality Mandai River. 

Kapuas). 
Doryichthys (Microphis) ignoralus Popta. 1906:21 1. 
Doryichthys Martensi Duncker. 1915:53. 
Doryichthys hrachyrhynchops Fowler. 1934/): 145 (type locality Chantaboon. 

southeast Siam). 

Material Examined— Western Borneo: Kapuas 1976-7. II; 36.5-104 mm 
(CAS 49436. GCRL 15759, MZB 3751. USNM 218071); Kapuas 1976-10, 2: 
72.5-82.5 mm (MZB 3752, USNM 218072); Kapuas 1976-13, 54.0 mm (MZB 
3753); Kapuas 1976-2 1 , 7: 52.0-97.5 mm (CAS 49437, MZB 3754, USNM 2 1 8073); 
Kapuas 1976-22, 105 mm (MZB 3755): Kapuas 1976-24. 70.5 mm (MZB 3756); 
Kapuas 1976-25. 1 14 mm (MZB 3757); Kapuas 1976-28. 2: 44.0-99.0 mm (MZB 
3758. USNM 218074); Kapuas 1976-29, 88.5 mm (MZB 3759). 



Distribution (Fig. 124). — Southeast Thailand. Malay Pen- 
insula. Western Borneo (Kapuas). 

Chandidae 

Chandidae (also commonly referred to as Ambassidae) is one 
of the few families of percoid fishes which exhibit greater di- 
versity in fresh water than in marine habitats. Seven of the eight 
genera currently recognized and about 21 of the approximately 
4 1 species are confined to fresh water in Madagascar, south Asia, 
southeast Asia, and the Australian region (including New 
Guinea). A systematic review of the family is given by Fraser- 
Brunner (1955). The fresh waters of western Borneo are inhab- 
ited by three genera and five species; three of the species are 
known only from fresh water in Borneo or in Borneo and Su- 
matra; the fourth species is known also from the Malay Pen- 
insula, and the fifth from Thailand. One of the species is a highly 
specialized new genus and species of scale-eaters discovered 
during the Kapuas survey of 1976. Another species is an oral 
brooder, the first member of the family Chandidae in which 
such parental care has been observed. Although generally known 
as glass-perches, of the species found in western Borneo, only 
Gynuwchanda filamentosa is actually transparent. 

Key to freshwater Chandidae of Borneo 

la Cheeks, gill cover, and body covered with scales; dorsal 
margin of prc-orbital bone entire or with small postero- 
ventrally projecting serrae; dorsal- and anal-fin branched 
rays 9-10, never filamentous; vertebrae 10+14-15=24- 
25; adults considerably larger than 33 mm 2 

lb Head and body scaleless; dorsal margin of preorbital bone 
with 3-4 enormous dorsally projecting serrae, dorsal- and 
anal-fin branched rays 12-14, filamentous in males; ver- 
tebrae usually 8+ 17=25; largest specimen 33 mm (males 
and females sexually mature at much smaller size) 

Gyinnochaitda filamentosa 

2a Dorsal profile gently sloped or but moderately steep; body 
depth at least 2.3; second anal-fin spine only moderately 
enlarged; gill rakers 22 or less 3 

2b Dorsal profile very steep; body depth about 2.0; second 
anal-fin spine enormously enlarged; gill rakers 29 

Paramhassis wolffii 

3a Dorsal profile gently concave at nape; cheek with fewer 
than 9 scale rows; dorsal preopercular margin serrate; 
lower jaw not strongly projecting 4 

3b Dorsal profile strongly concave at nape; cheek with 9-10 
scale rows; dorsal preopercular margin smooth; lower jaw 
strongly projecting Paramhassis macwlepis 

4a Scales in lateral series 40^7; jaws about equal, or lower 
jaw slightly projecting; second anal-fin spine larger than 
third Paramhassis apogonoides 

4b Scales in lateral series about 60; lower jaw much shorter 
than upper; second anal-fin spine about equal to or slightly 
shorter than third Paradoxodacna piratica 

Gymnochanda Fraser-Brunner, 1955 

Gymnochanda Fraser-Brunner. \'^ii:2\f){\\pe%pe<:K^(i\iiinochanda filamentosa 

Fraser-Brunner. 1955. by ongmal designation and monotypy). 
Gymnochanda Boeseman. 1957:75. pi. 3 (type species Gymnochanda filamentosa 



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CALIFORNIA ACADEMY OF SCIENCES 




Table 1 3. Comparison of the Two Scale-eating Genera of Freshwater 
Chandidae. 



Chanda 



Paratioxodacna 



Figure 125. Paradcxodacna piraluu. oral dcnlition- Kapuas 1476-33. 64.3 
mm (CAS 49440. paratype). 



Bocseman. 14S7. by original designauon and monoupv). N. B. Bocscman in- 
dependently chose the same genus and species name for this hsh. 



Gymnochanda filamentosa Fraser-Brunner, 1955 

(ivinnochiinda hianicnhtsa Fraser-Brunner. 1455:210. fig 4 (type locality "south- 
ern Malaya"; aquarium specimens) 

(iviiinnchandatildiuciilosa Boeseman, 1 45 7: 7 5 (type locality "Singapore''"; aquar- 
ium specimens). 

Materwl Examined. — Western Borneo; Kapuas 1976-8. 6; 4.1-23.9 mm 
(BMNH 1482,3.29.205-209. MZB 3767); Kapuas 1476-22, 47: 23.1-32.1 mm 
(AMNH 48441, C\S 44441. FMNH 44247, IRSNB 19749, MNHN 1982-716, 
MZB 3768, RMNH 28912. UMMZ 209866, I'SNM 230317, ZMA 116.546); 
Kapuas 1976-47. 22 7 mm (MZB 3769) 

Diagnosis. — Scales entirely absent. Dorsal profile strongly 
concave above eye; dorsal margin of preorbital bone with 3-4 
enormous dorsally projecting scrrac; dorsal portion of bony in- 
terorbital rim with about 10 strong postcroventrally projecting 
serrae. Jaws equal. Gill rakers about 16. Body transparent or 
translucent yellowish or yellowish-brown in life, filamentous 
rays of sexually mature males milk-white distally. 

Distribution. — Malay Peninsula (Johore). Western Borneo 
(Kapuas). 



Strongly prognathous 

Preorliital ridge with a large anterior 
spine directed dorsopostcriorly and 
9-10 smaller spines directed pos- 
tcrolaterally; no ventrolaterally 
projecting lamina 

Preorbital edge smooth 

Preopercular ridge smooth excepting 

terminal serra 
Lower edge of preoperclc with a lew 

irregular serrae, hind margin 

smooth 
Dorsal-lin rays 15-17. anal 14-17 
Cheek and gill cover scaleless, body 

scalclcss anteriorly 
Posterior nostril vertically oval, 

about twice as large as anterior 

nostril 
Vertically elongate humeral spot 
Vertebrae 10+15 = 25 (rarely 

10+16 = 26) 



Strongly retrognathous 

Preorbital ridge non-serrate, with a 
smooth-edged ventrolaterally pro- 
jecting lamina 



Preorbital edge with 9 strong poslero- 

ventrally directed serrae 
Preopercular ridge with 9 strong serrae 

Lower edge and hind margin of pre- 
operclc strongly and regularly serrate 

Dorsal-fin rays 9-10, anal 9 

Cheek, gill cover, and body fully scaled 

Postenor nostril round, about same 
size as anterior nostnl 

No humeral spot 
Vertebrae 10+14 = 24 



Paradoxodacna new genus 

Tvpe Specie^. — Piii'itdi>\odiicnd pnaliLii new species. 

Diagnosis. — A scale-eating chandid with retrognathous jaws 
(lower jaw markedly shorter than upper) and highly modified 
teeth; upper and lower jaw with two rows of large conical teeth, 
outer row with crowns labially directed, inner row with crowns 
lingually directed; upper jaw with a separate innermost row of 
very small conical teeth with lingually directed teeth, and lower 
jaw with similar small teeth in anteriormost portion of inner 
tooth row (Fig. 125), 

Discussion. — The dentition of Pa nu/oxoclucna is remarkably 
similar to that of Chanda Hamilton-Buchanan, 1822, the single 
species of which, C. nania Hamilton-Buchanan, 1822. is the 
only other known scale-eater in the family Chandidae (pers. 
obs.). Because of its scale-eating habits and dentition I thought 
at first that Paradoxodacna should be placed in Chanda but I 
have rejected this hypothesis because of their numerous differ- 
ences (Table 13). In short, the non-dental characters of Para- 
doxodacna suggest that its closest relatives are freshwater chan- 
dids endemic to Sumatra and Borneo, whereas the characters 
of Chanda seem to indicate that its closest relatives are other 
Indian freshwater chandids. The similarity of dentition in the 
two scale-eating genera is therefore interpreted as convergent. 

Etymology . — Paradoxodacna (feminine) from Greek "par- 
adoxos" and "dakno" (bite), in reference to the extraordinary 
jaw teeth. 

Paradoxodacna piratica new species 

(1 igures I 25. 126) 
iinhinsis nucrclcpis Vaillant. 1843 1 10 (in part. Kapuas) 

HoLOTYPE. — MZB 3762, 41,8 mm, Kapuas mainstream and mouth of Sungai 
Sekayam at Sanggau (Kapuas 1976-14). 

Paratypes.— CAS 44438, 31.8 mm, same collection as holotype; CAS 44439, 
MZB 3763, 2: 16.5-18,9 mm. Kapuas basin, sand and gravel bar in mouth of 



ROBERTS-WESTERN BORNEO FISHES 



161 




Figure 126. Faradoxodacna piralica. Kapuas 1976-14, 41.8 mm (MZB 3762, holotypc). 



Sungai Melawi at Sintang (Kapuas 1976-31); BMNH 1982.3.29.204, CAS 49440, 
MZB 3764. USNM 2303 16. 8: 19.0-69.3 mm. Kapuas mamstream near Kampong 
Nibung, about 100 km NE of Smtang (Kapuas 1976-33); MZB 3765. 52.9 mm. 
Kapuas mamstream 53 km W ol'Putussibau (Kapuas 1976-34); MZB 3766. 48.8 
mm. Kapuas basin, Sungai Tawang near Danau Pengunibung (Kapuas 1976-44); 
RMNH 7890. 34.3 mm. Kapuas basm. Smlang. Butlikofer. July 1894; MNHN 
1891 '581-582. 585. 3: 61.7-88.6 mm. Kapuas basm. Knapei, Sebruang. Chaper. 
1890 (identified as Amhassis microlepis by Vaillant); ZMA 112.376, 53.3 mm, 
Kapuas basin. Mendalan River. H. A. Lorentz, 28 June 1908. 

ScALE-EATiNG. — Faradoxodacna piralica feeds predominant- 
ly on scales of Cyprinidae. Stomach contents of one large spec- 
imen from each of four localities are as follows: 

Kapuas 1 976- 1 4, 3 1 .8 mm: 20 scales about 1 .0-3.0 mm long; 
a single aquatic larval insect head capsule about 1.5 mm long. 

Kapuas 1976-33, 69.3 mm: 10 scales about 1.5 mm long, 178 
scales about 2.0 mm long, and 7 scales 3.1-3.3 mm long; one 
small whole chandid (not young off. piratica) 14.4 mm. 

Kapuas 1976-34. 52.9 mm: 88 scales about 2.5-3.5 mm long, 
and 1 1 large, chevron-shaped scales (from Cyclocheilichthysl) 
5.0-5.2 mm wide and 6.5-6.8 mm long; one aquatic larval insect 
head capsule about 1.5 mm long and unidentified soft animal 
remains, possibly all insect, amounting to about 10 per cent of 
total food contents. 

Kapuas 1976-44, 44.8 mm: 16 scales 0.8-2.0 mm long and 
49 scales 2.7-4.4 mm long; no other food items. 

All or almost all of the scales seem to be from Cyprinidae, 
the predominant scaled fishes throughout the Kapuas basin. 
From the differences in size and shapes of scales it is evident 
that all four specimens had taken scales from at least two or 
three different prey, and some from two or three different prey 
species; scales ingested range in size from about that of the scale- 
eater's own scales to several times larger. Since P. piratica has 
relatively small scales, the prey probably ranged in size from 
slightly larger to two or three times the length of the scale-eater. 
In the two smallest specimens collected, 16.5-18.9 mm, the 
dentition is not yet modified for scale-eating; i.e., the crowns in 
the outer row of conical jaw teeth are curved inwards, as in 
juveniles and adults of non-lepidophagous chandids generally. 



rather than outwards as in adult P. piratica. The stomach of the 
16.5 mm specimen was empty except for the head of a minute 
aquatic larval insect, and that of the 18.9 mm specimen entirely 
empty. In a 19.0 mm specimen the dentition is nearly fully 
modified for scale-eating; its stomach contained 25 copepods 
0.7-0.9 mm body length and a single scale 1.0 mm long. For 
discussion of scale-eating fishes see Sazima (1983). 

Distribution. — Known only from the Kapuas. 

Etymology.— The mmt piratica (Latin, a sea-robber) refers 
to the scale-eating behavior. 

Parambassis Bleeker, 1874 

Paramhassis Bleeker. 1874i:86. 102 (type species .J /);/«.» s« apngonoides Bleeker. 

1851. by original designation). 
Pifttdamhassis Bleeker. 1874i:86 (type species Chanda Uila Hamilton-Buchanan, 

1822. by original designation and monotypy). 
Acciiuhnpcrca Castelnau, 1878;44 (type ^x>K\ei, Acanlhopcrcu gullivcri Castelnau. 

1878. by monotypy). 
U'hiilcvina Fowler and Bean. 1930;vii. 163. 334 (type species Amftassis wolffii 

Bleeker. 1850. by monotypy). Misspelled as Whiilcyui. p. 2. 148. 

Remarks.— The species here placed in Parambassis have been 
placed in Chanda Hamilton-Buchanan, 1822 by various au- 
thors, many of whom regarded Clianda lata Hamilton-Buchan- 
an, 1822 as type species of Chanda. The ICZN has ruled that 
Chanda nama Hamilton-Buchanan, 1822 is type species of 
Chanda (Opinion 1121, 1979). Chanda nama is a highly spe- 
cialized scale-eater which I consider to represent a monotypic 
genus and species (see discussion under Paradoxodacna). The 
ne.xt available generic name for the other species formerly placed 
in Chanda is Parambassis Bleeker, 1874. 



Parambassis apogonoides (Bleeker. 1851) 

(Figure 127) 

Amhassis apogonoides Bleeker. 185 1/i:200 (type locality Banjermassing). 
Parambassis apogonoides Bleeker. 1874; 103. 
Chanda apogonoides Fraser-Brunner. 1954;204. 



162 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 127, I'cirainhtissi.s upogonoidcs- Kapuas I'*7b-I4, ■42.6 mm (CAS 4'J442). 



Material Examined— WL'stem Borneo; Kapuas 1^76-14. 16: 22.1-42,6 mm 
(BMNH 1182.3.29.210-211, CAS 49442. FMNH 94248. MZB 3770. RMNH 
28913); Kapuas 1976-15. 2; 37.6-42.3 mm (MNHN 1982-717, MZB 3771); Ka- 
puas 1976-33, 86: 3.3-51.6 mm (CAS 49443. MZB 3772); Kapuas 1976-35. 2: 
42.8-44.3 mm (MZB 3773. USNM 230318); Kapuas 1976-36, 54.5 mm (MZB 
3774); Kapuas 1976-45, 2: 32.2-33.9 mm (MZB 3775, UMMZ 209919); Kapuas 
1976-50. 4; 26.5-35.9 mm {MZB 3776, USNM 2.30319); Sinlang. 9: 20.8-37 8 
mm (RMNH 7890); Raoen embouchure, Kapuas, 58.5 mm (RMNH 7891); Ka- 
puas PoetocsSibau, 42.8 mm(ZMA 1 12.310); Kapuas. Mendalan,62.2 mm(ZMA 
112.309). Easlem Borneo; Mahakam basin, Tcpoc. 2; 56.8-62.3 mm (RMNH 
7889), No locality; Bleeker collection, 3; 40.5-67. 1 mm (RMNH 5558) (note: this 
67.1 mm specimen, now 84 mm total length, with penmarks on the lateral line 
scales presumably made by Bleeker, is probably the holotype [88 mm total length] 
from Bandjermasstng). 

DiAGNosis. — Dorsal profile moderately sloped, gently con- 
cave. Dorsal margin of preorbital bone smooth. Orbital rim 
with 2-3 small serrae dorsoposteriorly, otherwise smooth. Dor- 
sal prcopercular margin serrate. Second anal spine slightly larger 
than third. Gill rakers 21-22. Scales in lateral series 40^7. 
Cheek with 5 scale rows. 

Oral Brooding. — Parental care is unknown in the family 
Chandidae (Breder and Rosen 1966; Blumer 1982). Thus it is 
particularly noteworthy to report that C. apogonouics practices 
oral brooding. The orobranchial cavity of a 51.6 mm male from 
Kapuas 1976-33 contained 81 young of 3.3-7.0 mm. Two size 
classes are present, 78: 3.3-5.5 mm, slender and white, and 3: 
±7.0 mm, much thicker and yellowish, presumably representing 
two broods. The orobranchial cavity also contained one whole 
larval ephemeropteran and a few small pieces of insects. The 
guts of this specimen contained fine black detritus, at least partly 
from insect remains. 

There is no indication of oral brooding in the other Kapuas 
chandid species; a brief survey of freshwater and marine chan- 
dids from other areas in the CAS collections failed to reveal 
additional oral brooding species. 



Distribution. — Sumatra (Muara-Kompeh. Djambi). Borneo 
(Barito, Kahajan, Kapuas, Mahakam. Mendalan [=Barito?] 
rivers). Unknown from northern Borneo. 

Parambassis macrolepis Bleeker, 1857 

(Figure 128 upper) 

Bognla macrolepii Bleeker. 1857(/;13 (type localit) Pontianak, in flumine Ka- 
puas). 

Aniha\si\ {Boi^Oild) nu^inlcpis \on Martens, 1868.9 (Kapuas, unwarranted emen- 
dation of spelling). 

Pcininihassis nucrolepis Bleeker. 1874:104. 

Anihassis nucrolcpis Vaillant. 1893:1 10 (Kapuas). 

Chiincla macrolepis Fraser-Brunner, 1955:205. 

Material Examined. — Western Borneo: Kapuas 1976-19,68.5 mm (MZB 3777); 
Kapuas 1976-20. 15: 65.6-97.5 mm (BMNH 1982. 3. 29.212-213, CAS 49444, 
FMNH 94249. MZB 3778. UMMZ 209864); Kapuas 1976-33, 4: 65.2-82.3 mm 
(MZB 3779. LISNM 230320); Kapuas 1976-49, 2: 57.7-88.0 mm (MZB 3780, 
RMNM 28914); Kapuas basin, Knapei, Scbruang, 2: 57. .3-65. 8 mm (MNHN 
I S91 583-584); Kapuas basin. Sintang, 4: 77,0-98,9 mm (RMNH 7892); Kapuas 
basin, Boenoet, 3: 66.7-72,4 mm (ZMA I 12 337), 

Diagnosis. — Body moderately deep. Dorsal profile strongly 
concave. Second anal spine larger than third. Gill rakers 17, 5 
on upper limit; first 4 rakers on upper limit mound-shaped. 
Scales in lateral series 50-59. Cheek with 9-10 scale rows. Dorsal 
prcopercular margin entire. Dorsal margin of preorbital bone 
smooth. Orbital rim non-serrate. 

Distribution. — Sumatra (Batang Hari). Borneo (Kapuas 
only?). 



Parambassis wolffii (Bleeker, 1850) 

(Figure 1 28 lower) 

Amhasiis li'ollhi Bleeker. I 850:9 (type locality Banjermassin). 
Aiiihcnsi'i rohnsliis Gunlher, 1859:222 (type locality Borneo), 



ROBERTS-WESTERN BORNEO FISHES 



163 




Figure 128. Parambassis. Above, P. macrolepis. Kapuas 1976-33, 82.3 mm {USNM 230320); below, P nvlffii. Kapuas 1976-14, 81.4 mm {CAS 49445). 



164 



CALIFORNIA ACADEMY OF SCIENCES 



Ambassisbouiengeri Volz, 1903:553 (type locality "Banju asm. dcm grossen Astuar 

an der Nordostkiiste der Resident Palcmhang. Sumatra"). 
Acainhoperca wolffii Fowler. 1937:230, 
Chanda wolffii Smith, 1945:482. 
Paramhassis wolffi Allen, 1982:166. 

Material Examined. — Western Borneo: Kapuas 1976-14, 2: 24.8-81.4 mm 
(CAS49445. MZB 37811; Kapuas 1976-19. 99.6 mm (MZB 3782): K.apuas 1976- 
44, 3: 97.4-1 18 mm (CAS 49446. MZB 3783. I.ISNM 23032 1 ): Kapuas 1976-52. 
58.7 mm (MZB 3784). 

Diagnosis. — Dorsal profile ver> steep, almost straight. Depth 
about twice in standard length. Second dorsal, pelvic, and sec- 
ond anal spines greatly enlarged. Gill rakers 29, 10 on upper 
limb. Scales in lateral series 43-46, Cheek with 7-8 scale rows. 
Dorsal prcopercular margin smooth. Posterior margin of orbital 
rims and dorsal margin of preorbital bone with small posteriorly 
directed serrae, 

Parambassis wolffii differs strikingly from all of the marine 
Chandidae and from the other freshwater chandids in India and 
southeast Asia, It is perhaps closely related to two New Guinean 
freshwater species currently known as PciniDihassis gigas and 
P. confinis. 

Distribution.— Thailand (Chao Phrya; Mekong), Sumatra 
(Palembang). Borneo (Baram?, Barito, Kapuas, Kahajan). (Ba- 
ram record of Weber and de Beaufort [1929:402] doubtful- 
source unknown.) 



tong, Kwantan R,, Djambi). Borneo (Baram, Kapuas, Pamang- 
kat, Bankayan), (Not known from Java,) 



Datnioididae 

Daimoides comprises three or four species of morphologically 
generalized, deep-bodied percoids with vertically barred color 
patterns found in fresh and brackish water in southeast Asia 
and the tropical Australian region (especially New Guinea), The 
genus has been placed in the otherwise monotypic family Lo- 
botidae by Jordan (1919), Weber and de Beaufort ( 1936), Nelson 
(19841, and others but there is little or no evidence in favor of 
this relationship and it seems untenable. Fowler (1931) recog- 
nized Datnioides as forming a subfamily Datnioidinae 
of Theraponidae, but Van (1978:193) noted that Daimoides 
does not share diagnostic theraponid osteological characters and 
excluded it from that family, Johnson (1984), unaware of Fow- 
ler's use of the family-group name Datnioidinae, placed Dat- 
nioides along with 1 1 other genera as incertae sedis in an al- 
phabetical list of 80 percoid families. Convinced that Datnioides 
does not belong in Lobotidae or Theraponidae, and without any 
solid evidence indicating its relationship to Nandidae or any 
other percoid family, I hereby recognized Datnioides as forming 
Its own monotypic family. 



Nandidae 

Nandidae. as herein understood, is a tropical primary fresh- 
water family comprising the traditional leaf-fish genera: Mono- 
cirrhus and Polycentnism South America, Afroiiandus and Poly- 
centropsis in west Africa, and Nandiis in south and southeast 
Asia. 

Nandus Valenciennes, 1831 

.Vum/jK Valenciennes in Cu\ier and Valenciennes, 1831:481 (type species Coins 
iiaiuhis Hamilton-Buchanan. 1822. by absolute laulonymy). 

Bcdiila Gray. I83.V34 (type species Bcdiila iwbidosa Gray, 1833, by monotypy''). 
(Reference not seen.) 

Nandus comprises two species: N. nandus in India, Burma, 
and Thailand and A', nehulosus in southeast Asia including the 
Greater Sunda Islands. 

Nandus nebulosus (Gray. 1833) 

Bedida nebulosus Gray, 1833, pi. 88, fig. 2 (no locality). 

Nandus marmoralus Valenciennes //; Cuvier and Valenciennes, 1831:482 (re- 
placement name for Coius nandus Hamilton-Buchanan. 1822). 
\iindiis nebulosus Bleeker, 1852i!:92. 
Nandus bornecnsts Steindachner. 1901422 (type locality Baram River). 

Material Examined. — Western Borneo: Kapuas 1976-8. 2: 16.2-37.8 mm (CAS 
44449. MZB 3792): Kapuas 1976-13. 55.7 mm (MZB 3793); Kapuas 1976-16. 8: 
9,6-78.8 mm (CAS 49450. MZB 3794); Kapuas 1976-17. 57.3 mm (MZB 3795); 
Kapuas 1976-39, 4: 37.3^8 mm (MZB 3796. USNM 230323); Kapuas 1976- 
43. 3: 24.3-69.6 mm (BMNH 1982.3.29.2 14-21 5. MZB 3797); Kapuas 1976-46. 
5: 30.0-42.6 mm (CAS 49451. MZB 3798); Kapuas 1976-47, 71.7 mm (MZB 
3799); Kapuas 1976-51. 4: 27 5-31.5 mm (MZB 3800. USNM 230324), 

Distribution.— Thailand (Chao Phrya, Poon Duang River, 
Nakon Srilamarat, Chantabun, Ban Thung Luang), Malay Pen- 
insula (Johore, Singapore), Sumatra (Lahat, Indragiri, Danau 
Sialong. Taluk, Gunung Sahilan, Sungei Mahe, Laut Tador, La- 



Datnioides Bleeker, 1853 

Datnioides Bleeker. I853r:440 (type species Coius polola Hamilton-Buchanan. 
[^22 = Chaelodon qiiadrdaseialus Scvaihanow 1809. by monotypy. 

In the original description of Datniodes Bleeker (1853:442) 
indicated Coins bmotatus Gray. 1834 and Lohotes he.\a:ona 
Bleeker. 1851 as junior synonyms of Datnioides polota. As 
Bleeker clearly recognized D. polota as the only valid species of 
Datnioides it is the generic type species by monotypy, 

Datnioides comprises at least three species: D. quadrifasciatus. 
supposed to be widely distributed in south and southeast Asia, 
tropical Australia and southern New Guinea; D. microlepis from 
southeast Asia; and D camphelli Whitley, known only from the 
upper Sepik River of northern New Guinea (Munro 1967). 

Key to Datnioides of southeast Asia (from Smith 1945) 

la Scales in lateral line about 70; branched dorsal rays 13- 
14; body and head with 8-10 dark brown cross bands 
some of which become confluent with age; a dark round 
spot on opercle D. quadrifasciatus 

lb Scales in lateral line about 105; branched dorsal rays 15- 
16; body and head with black cross bands, which remain 

distinct at all ages; no round dark spot on opercle 

D. microlepis 

Datnioides microlepis Bleeker, 1853 

Dalnoides murolepis Bleeker, I853i':442 (type locality ■■Pontianak. in llumine 
Kapuas") 

Material Examined -Western Borneo: Kapuas 1^76-14, 245 mm (MZB 3785); 
Kapuas 1976-20.2: I s 1-2 N mm (CAS 40447. MZB 3786). 

Distribution. — Kampuchea. Thailand (Mekong, Chao 
Phrya). Borneo (Kapuas), This species apparently is restricted 



ROBERTS-WESTERN BORNEO FISHES 



165 



to fresh water, and is generally recorded much farther inland 
than D. quadrifasciatus. 

Datnioides quadrifasciatus (Sevastianov. 1809) 

Chaciodoii quadnfascialus Sevastianov, 1809:448 (reference not seen). 

Corns polota Hamilton-Buchanan. 1822:95, 370 (type locality Ganges). 

Loboles hexazona Bleeker. 1851^:9 (type locality Bandermassing. in fluviis). 

Dalmoidcs polota Bleeker, 1853<':441. 

Dalnioides quadnfasaatus Bleeker. 1876-77.32. 

For additional synonymy see Weber and de Beaufort (1936:462). 

Material Examined. — None. 

Distribution. — India. Burma. Thailand. Vietnam. Sumatra. 
Borneo (Sarawak, Sambas, Kapuas, Kukajan, Barito). New 
Guinea. In brackish water in mouths of large rivers and coastal 
lagoons as well as in freshwater lakes and rivers above tidal 
influence. 

Pristolepidae 

As here understood, Pristolepidae includes only Pristolepis. 
Relationships of this deep-bodied and relatively generalized 
pcrchlike genus have not been satisfactorily investigated. It is 
perhaps related to the anabantoids, or to the so-called "pro- 
toanabantoid stock." Pristolepis was excluded from Badidae, 
which may also be related to the anabantoids or protoanaban- 
toids, by Barlow et al. (1968). Pristolepis has been placed in 
Nandidae (e.g., by Greenwood et al. 1966) but is more gener- 
alized than typical nandids and probably is not closely related 
to them. It differs from nandids in having very short jaws with 
ascending premaxillary process not notably elongate; branchi- 
ostegal membranes united very far posteriorly and covered with 
scales; scales with ctenii in multiple rows instead of uniserial; 
and pore-bearing scales of lower lateral line series extending to 
middle of caudal fin. Pristolepis agrees with Nandidae in having 
the lateral line divided into separate upper and lower parts, but 
this character is found also in some anabantoids and most Cich- 
lidae. Pristolepis has a distinctive vertebral formula. 13+12=25. 

Pristolepis Jerdon, 1848 

Pristolepis Jerdon, 1848:141 (type species Pristolepis margiimla Jerdon, 1848) 

(reference not seen). 
Calopra Bleeker, 1 85 1 /■:65 (type species Catoprafasciata Bleeker, 1 85 1 , by mono- 

typy)- 

Paranaiuliis Day, 1865:130 (type species Calopra malabarica Gunther, 1864, by 
monotypy). 

The genus comprises three or four species and is found in 
southern India, Burma, Thailand, Kampuchea, the Malay Pen- 
insula, and Asian continental islands of Indonesia; one or pos- 
sibly two species in western Borneo. 

Pristolepis fasciata (Bleeker, 1851) 

Calopra fasciata Bleeker, 185iy':61. 65 (type locality Bandjcrniassing, in fiuviis). 
Calopra nandoides Bleeker, 1851/:61 (nomen nudum): Bleeker, 1851:172 (type 

locality presumably Java). 
''Calopra Groom Bleeker. 1852a:90 (type locality Billiton). 
Pristolepis fasaatus Bleeker, 1876-78:80, pi. 391, fig, 4. 
''Pristolepis groom Bleeker, 1876-78, pi. 391. fig. 2. 
Calopra nandoides von Martens, 1876:394 (Kapuas), 
Calopra fasciata von Martens, 1876:394 (Kapuas). 
''Catopra grooti von Martens, 1876:394 (Kapuas). 



Pristolepis fasciams Vaillant, 1893:109 (Kapuas); 1902:32 (Kapuas). 
Pristolepis fasciata Mora. 1923:177. 

Material Examined.— Western Borneo: Kapuas 1976-6, 9: 16.7-24.7 mm (CAS 
49452, MZB 3801, USNM 230325); Kapuas 1976-8. 47.0 mm (MZB 3802); 
Kapuas 1976-19, 2: 98.0-105 mm (CAS 49453. MZB 3803); Kapuas 1976-24. 
49.6 mm (MZB 3804); Kapuas 1976-27, 59,9 mm (MZB 3805); Kapuas 1976- 
28. 71.0 mm (MZB 3806); Kapuas 1976-37. 61.2 mm (MZB 3807); Kapuas 1976- 
43, 2: 50.8-89.5 mm (MZB 3808, USNM 230326); Kapuas 1976-47. 92,0 mm 
(MZB 3809). 

It is possible that my Kapuas material includes two species, 
Pristolepis fasciata and Pristolepis grootii, both of which have 
been reported from the Kapuas, but I have been unable to 
determine whether these really are distinct species. For discus- 
sion of supposed difl^erences between them and additional syn- 
onymy see Weber and de Beaufort (1936:478-48 1 ). The account 
of the genus Pristolepis in Bleeker's Atlas is incomplete, but his 
figures off. fasciata and P. grootii should be consulted (Bleeker 
1877:80, pi. 391). Note that Weber and de Beaufort's figure of 
Pristolepis fasciata (Weber and de Beaufort 1936, fig. 95) re- 
sembles not P. fasciata but rather P. grootii as figured by Bleeker. 

Distribution. — Widely distributed in fresh water in Thai- 
land, Malay Peninsula, Sumatra, Borneo (western, southern, and 
eastern), Java. Burma? 

Toxotidae 

The tropical Indo-West Pacific Toxotidae or archerfishes, re- 
nowned for the ability to knock insects off" overhanging vege- 
tation onto the water by accurately aimed jets of water forcefully 
ejected from their mouth, comprise a single genus with six species 
(Allen 1978). One species, Toxotes jaculator. generally occurs 
in brackish habitats (mainly mangrove creeks) but is sometimes 
encountered in pure fresh water in the lower parts of rivers and 
coastal streams. The other species generally go through their 
entire life cycle in fresh water, sometimes hundreds of miles 
inland. 

Toxotes Cuvier, 1817 

Toxotes Cmvkt. 1817:338 (type ipeQ\e% Sciaena jaculator Pallas, 1767, by mono- 
typy). 

Toxotes is readily distinguished from all other freshwater per- 
ciform fishes in southeast Asia by its characteristic body form 
and color, with a very sharp snout, single dorsal fin with 4-6 
large spines and 11-14 rays lying entirely on posterior half of 
body, and sides with several large dark vertical bars or roundish 
spots on a yellowish background. The Indonesian or Malay 
name for Toxotes is ikan sumpit (blowgun fish). Three species 
occur in western Borneo, but only T. microlepis is known from 
fresh water in western Borneo. 

Toxotes microlepis Giinther, 1 860 

Toxotes microlepis Gunther, 1860:68 (type locality Siam). 

Material Examined — Western Borneo: Kapuas 1976-14, 78.2 mm (MZB 3787); 
Kapuas 1976-15. 3: 22,5-40.3 mm (MZB 3788, USNM 230322); Kapuas 1976- 
19, 121 mm (MZB 3789); Kapuas 1976-33. 74,2 mm (MZB 3790); Kapuas 1976- 
45, 2: 70,9-76 2 mm (CAS 49448. MZB 3791), 

This species is distinguished by 5 dorsal-fin spines, 6-7 bars 
on side of body, and scales in lateral series 42-45. 



166 



CALIFORNIA ACADEMY OF SCIENCES 



Distribution.— Thailand (Chao Phr>a, Bangpakong). Su- 
matra (Palembang). Borneo (Kapuas. Barito. Mahakam). 

Polynemidae 

The tropical and subtropical family Polynemidae or thread- 
tins, with seven genera and about 35 species, occurs mainly in 
marine and brackish habitats but several species apparently are 
endemic to fresh water. In southeast Asia, although other genera 
may enter river mouths and the lower portions of rivers, the 
only genus with freshwater species is Polyncnuis. 

Polynemus Linnaeus, 1758 

Po/yiu'iiius Linnaeus. 1758:317 (type species I'nlvncmii^ paraJi^ciii Linnaeus. 

1758, by Opinion 93. IQ26. ICZN). 
Polislonemus Gill, 1861:277 (type species I'lilvnciiiiis nnillililis Schlegel, 18-15. by 

monol\p\ ) 

Polynemus are readily distinguished from all other freshwater 
fishes in southeast Asia by having the pectoral fin divided into 
two discrete portions, an upper portion with 12-17 rays united 
by fin-membrane and a ventral portion with 7-16 extremely 
elongate free filamentous rays. When the fish swim slowly in 
open water the free lower rays can be widely spread apart like 
the ribs of an umbrella. 

There seem to be more freshwater polynemids in Borneo than 
anywhere else. Polynemus macrophlhalnius and P. nndtifilts oc- 
cur in the Kapuas in western Borneo and probably both occur 
in other Bornean rivers; they are also known from rivers in 
Sumatra. At least two other species are known only from rivers 
in Borneo: P. hornadayi Myers, 1936 from the Sadang R. in 
Sarawak, and P. hillen (Fowler, 1905) from the Baram R. in 
Sarawak. 

For keys to genera of Polynemidae and species of Polynemus. 
references to earlier literature, and additional information see 
Myers (1936). 

Key to freshwater Polynennts of western Borneo 

la Lower portion of pectoral fin with 7 free rays 

P. macrophihalmus 
lb Lower portion of pectoral fin with 14-16 free rays 

P multifilis 

Polynemus macrophthalmus Bleeker, 1858-59 

Polynemus imicrophlluilmin BIccker. 1858-5^:111 (type locality Palembang, in 

osliis fluminis Mussi) 
Pnlvncmw: hurnccnus Vaillanl. I8'J3:1IW (Kapuas). 

M.'\TER1AL Examined —Sumalra. Balang Han at Djamhi. 3: 140-223 min(ZM.A 
I 14.431). Western Borneo: Kapuas R. at Bunut. 164 mm (ZMA 1 14.432); Kapuas 
1976-19. 258 mm (MZB 3810) 

Color in life of the single Kapuas specimen (obtained in very 
fresh condition in fish market at Sintang) overall pale greyish 
white or dull silvery. I believe that this species may sometimes 
be brilliant orange or orangish-gold in life. A half-dozen fresh 
specimens of Polynemus with head, body, and fins entirely bnght 
orange were seen by me in the market at Pontianak but were 
not preserved or positively identified to species. The three ZMA 
specimens from Sumatra, although preserved long ago, still show 
such coloration; I suspect this is not an artifact of preservation 
but due to their retention of pigmentation from life. 



Distribution. — Sumatra (Palembang, Mussi R., Djambi, Ba- 
tang Hari). Borneo (Kapuas). 

Polynemus multifilis Schlegel, 1843 

Po/yiwmus nuillihlis Schlegel. 1843 in Sicbold. 1842-50:29 (type locality "Zu- 
idkust van Borneo bij Banjermassing" l=Banto River?]). See also Schlegel ( 1 848). 

Pnlvncnuis qualcrdccimtiln Pel. 1851:10 (type locality ■■lilloribus meridionalibus 
insulac Borneo"). 

Pntislonemus miiltitilis Gill. 1861:277. 

Tnchidioii muhifilis Bleeker. 1865/': 1 74. 

Material Examined. — Sumatra: Djambi. 149 mm (ZMA 114.430). Wcslem 
Borneo: Kapuas R. at Bunut. 4: 156-183 mm (ZMA 1 14 403); Kapuas 1976-19. 
6: 111-184 mm (CA 47198. MZB 3811. RMNH 28839); Kapuas 1976-33. 5: 
73.5-1 19 mm (CAS 47199. MZB 3812); Kapuas 1976-44, 6: 88.9-138 mm (MZB 
3813. USNM 230327); Kapuas 1976-48. 12: 27.0-76.7mm (BMNH 1982. 3. 29. 216- 
218. CAS 49454. MZB 3814). 

The number of free pectoral-fin rays reported for this species 
is 14. Most of my Kapuas specimens have 15 or 16. Color in 
life dull white or faintly silvery. 

Distribution. — Sumatra (Djambi). Borneo (Kapuas, Barito). 



Eleotridae 

The Eleotridae or sleepers comprise some 40 genera and 1 50 
species of mostly tropical, marine, brackish, and freshwater fish- 
es. Two genera and four species occur in the fresh waters of 
western Borneo. 

Eleotris Schneider m Bloch, 1801 

Eleolns Schneider in Bloch. 1801:65 (type species Uobius pisonis Gmelin, 1789, 
b\ subsequent designation of ICZN. 1958. Opinion 93). 

Eleotris melanosoma Bleeker, 1852 

Eleotris )nelano\oiihi Bleeker. lS52c:705 (type locality Wahai. Ceram and Su- 
matra occidenlalis. in man). 

M.\TERiAL E.\,\MiNED, — Western Borneo: Kapuas 1976-1 7, 1 1 1 mm (MZB 3816). 

For a tentative synonymy of E. melanosoma see Koumans 
(1953:297-298). It apparently is very widely introduced in es- 
tuaries and rivers in the East Indies and Australian region; it 
also has been reported from Guam, Samoa, and Mindi on the 
•Atlantic coast of the Panama Canal (Koumans 1953:298-299). 

Oxyeleotris Bleeker, 1874 

Oweleolris Bleeker. 1874/i:294. 302 (type species Pleolris mannorala Bleeker. 

1852. by original designation and monotypy). 
''Giganlogohiiis Fowler, 1905:511 (type species Uiganlogohius jordani Fowler, 

1905. by original designation and monotypy). .See discussion under Oxyeleolns 

nhlinioruld 

Oxyeleotris is unusual among eleotrid genera in being almost 
entirely restricted to fresh water. Endemic freshwater species of 
the genus occur in southeast Asia and others in the Australian 
region. Three species are known from western Borneo. 

Oxyeleotris marmorata (Bleeker. 1852) 

Eleolns marmonita Bleeker. 1852/i:424 (type localits Bandjermassing et Palem- 
bang. in fluviis). 
Oxyeleotris mannorala Bleeker. l874/i:303. 

''Gigantogohiiis lordani Fowler. 1905:51 I (type locality "Baram. Borneo"). 
Callielcotris plalyLephalus Fowler. 1934/': 156 (type locality Bangkok). 



ROBERTS-WESTERN BORNEO FISHES 



167 



Material Examined.— Western Borneo; Kapuas 1976-16, 5: 12.2-105 mm 
(CAS 49455, MZB 3817). Kapuas 1976-17, 2: 52.1-54.5 mm (BMNH 
1982.3.29.219, MZB 38181; Kapuas 1976-19. 2; 187-189 mm (MZB 3819, RMNH 
28915); Kapuas 1976-44, 137 mm (MZB 3820); Kapuas 1976-47. 4; 44.1-148 
mm (MZB 3821, USNM 230328). 

As pointed out by Bleeker (1877:23), O. mannoratus is the 
largest gobioid in the East Indies; it may well be the largest 
gobioid of all. So far as I have been able to determine it is the 
only freshwater gobioid in Sundaland for which specimens over 
300 mm have been reported. Bleeker had specimens to 430 mm 
(total length) but it apparently grows even larger. Although the 
holotype oi Gigantogohnis jordani is still lost (Bohlke 1 984: 107), 
there seems no reason to doubt Koumans's tentative conclusion 
that G. Jordan! is a synonym of (9. inarmorata (Koumans 1940: 
134, 1954:354). The holotype, "26 inches long" (total length?) 
seems to be the largest specimen of gobioid on record. Presuming 
that the length of 26 in. refers to total length, its standard length 
would have been about 540 mm (caudal fin about 17 percent 
of total length). The next largest gobioid specimen known to me 
is a 437 mm Bimaka henverdemi (Weber, 1910) from the Fly 
River, New Guinea (Roberts 1978:1 1, 65), 

Distribution, — Widely distributed in fresh water in Thailand 
(Smith 1945:509). Malay Peninsula. Sumatra, Western and 
northwestern Borneo, 

Koumans (1953:355) reported the occurrence of O. mar- 
morata in estuaries as well as rivers and included the Philip- 
pines, Fiji, and Celebes within its range. This information is 
doubtful. O. mannorata may occur in brackish habitats within 
its range, but all of the well documented locality records appear 
to be from fresh water. Occurrence in the Philippines apparently 
is based on the doubtful record by Meyer ( 1 885:30) from Laguna 
de Bay, Luzon (see Herre 1927:75), Herre( 1953:365, 1954:729) 
never questioned Meyer's identification, and supposed that O. 
mannorata "no doubt is in river mouths in Balabac, Palawan, 
Tawi Tawi, and Cotabato Province, Mindanao," So far as I 
have been able to determine Herre never collected a specimen 
of this species in the Philippines, I suspect that reports of O. 
inarmorata from Celebes and Fiji are based either on erroneous 
locality data or misidentification. 

Oxyeleotris urophthalmus (Bleeker, 1851) 

Eleolns urophthalmus Bleeker, 185W(;202 (type locality "Bandjermassing, in flu- 

viis"), 
Oxyeleotris urophthalmus Bleeker, 1877;23, 

Material E.xamined— Borneo; Sarawak, Kuching, 5; 112-131 mm (CAS-SU 
33135). 

Distribution,— Thailand? Malay Peninsula, Borneo (Sara- 
wak, Kapuas, Kahajan, Mahakam), Celebes? New Guinea? (see 
Koumans 1953:356), 



Oxyeleotris urophthalmoides (Bleeker, 1853) 

Eleotns urophthalmoides Bleeker, l853/);273 (type locality Palembang; Lacus 

Meninju; Sambas). 
Oxyeleotris urophthalmaoides Bleeker, I877;21, 25. 

Material Examined —Western Borneo; Kapuas 1976-51, 2; 56.5-114 mm 
(CAS 49456, MZB 3822). 

Distribution, — Sumatra, Borneo, Nias, Known only from 
fresh water. 



Gobiidae 

Gobiidae, probably the most speciose of all living fish fam- 
ilies, is rather poorly represented in the fresh waters of western 
Borneo, Only eight species were encountered during the Kapuas 
survey of 1976. All of these were taken in freshwater habitats, 
but only two species, Brachygobius xanthomelas and Cala- 
miana sp, undet,, were found in the interior of the Kapuas basin 
living in association with a rich assortment of primary and 
secondary freshwater fish species. The other six were found only 
near the coast and in habitats greatly altered by human activity 
(such as canals or streams flowing through plantations) in which 
primary freshwater fishes were poorly represented. It is note- 
worthy that gobiids were found at freshwater localities in the 
mainstream of the Kapuas River, The number of Gobiidae 
could be considerably increased here by addition of species pres- 
ent in estuarine or brackish waters in western Borneo and likely 
to enter fresh water for short distances. 

It is possible that some of the undetermined species referred 
to Mugilogobius, Psetidogobiopsis, Pseudogobius, and Stigma- 
togobius are undescribed. Their generic identification, for help 
with which I am indebted to Douglas Hoese, should be regarded 
as tentative. 



Brachygobius Bleeker, 1874 

Brachygobius Bleeker, l874/r3l5 (type species Gobiiis doriae GUnlher, 1868, by 
original designation and monotypy). 

Brachygobius (bumble-bee gobies) are short, heavy-bodied 
gobies with moderately large eyes; broad interorbital space; no 
head pores; an upturned mouth with prominent lower jaw; con- 
ical jaw teeth in 2-3 rows: scales in midlateral series 21-50; and 
coloration consisting of a few well defined black and yellow 
(sometimes orangish or cream-colored) vertical bands, Brachy- 
gobius differs from all or almost all other gobiid genera in having 
simple first ray of second dorsal fin segmented rather than un- 
segmented and in having first and second anal-fin rays instead 
of only first anal-fin ray without a medial pterygiophore (D, 
Hoese, pers. comm,, October 1983), There are at least eight 
species (Inger 1958), some found in estuaries, mangroves, and 
tidal creeks but others (perhaps the majority) restricted to fresh 
water, A key to the species is provided by Inger (1958), 



Brachygobius doriae (Gunther, 1868) 

Gobius Doriae Gunther. 1868/);265 (type locality Sarawak). 

Brachygobius doriae Hardenbcrg, 1936;252 (Pontianak, Padang Tikarbay, Kubu 

Terentang, Telok Pekadai). 
Brachygobius intnus Koumans. 1941;269, 1953:194 (in part). 

Material Examined— Western Borneo; Kapuas 1976-2, 4; 23.2-26.5 mm (CAS 
49457, MZB 3823); Kapuas 1976-3, 6; 19.9-30.7 mm (BMNH 1982.3.29.220- 
222. MZB 3824, USNM 230329). 

Brachygobius doriae is a relatively large and particularly heavy- 
bodied species typically found in tidal creeks and disturbed 
habitats such as man-made canals. It has three light-colored 
vertical bands. In numerous live fish observed during the Ka- 
puas survey of 1976 the anteriormost light-colored band was 
orangish and the posteriormost two bands of a decidedly lighter 
canary yellow. According to Hardenberg (1936:252) it feeds 
mainly on mosquito larvae. 



168 



CALIFORNIA ACADEMY OF SCIENCES 



Distribution. — Borneo (Sarawak, Kapuas, coastal streams 
and tidal creeks in western Borneo). Perhaps much more widely 
distributed. 

Brachygobius xanthomelas Herrc, 1937 

Brachv,i;<ihnis xciiilhnniclus Herrc, l<>37:4.1 (type locality Mawai District, Johore). 
Brachy^ohnis niiiiu^ Roumans, 1^41:264, 195-VI^4 (in part). 

Material Examined. — Malay Peninsula: Mawai Dislricl, .lohore. 1 1: 8.4-27.6 
mm (CAS-SU 30<^53. I6'^64. holotypcand paratypcsof B .xunllioiiicias). Western 
Borneo: Kapuas 1976-7, 4: 8.3-1. V6 mm (CAS 444.^8. MZB .■(825); Kapuas 1976- 
14. 14.0 mm (MZB 3826); Kapuas 1976-16. 9: 1 1 .8-16.0 mm (MZB 3827, USNM 
2303.30); Kapuas 1976-17. 16; 12.2-14.9 mm (BMNH 1982.3.29.223-228. FMNH 
94250. MZB 3828); Kapuas 1976-47, 23: 13.2-17.3 mm (CAS 49459, KUMN 
2859. MZB 3829) 

Brachygobius xanthomelas is a relatively small species, ap- 
parently restricted to fresh water, previously known only from 
the Malay Peninsula. In live specimens observed in 1976 the 
light bands were all pale yellow. 

Calamiana Herre, 1945 

Calaiuuina Herrc, 1945:80 (t\pe species Calannana niui^nons Herre. 1945. by 
original designation and monotypy). 

Calamiana comprises several relatively small species of sex- 
ually dimorphic gobies in which the males, growing larger than 
females, develop extremely elongate jaws and expanded or bul- 
late opercular regions. The systematics of the species are poorly 
known and it is likely that some are undescribed. 

Calamiana sp. undet. 

Material Examined. — Western Borneo: Kapuas 1976-13. 2: 15 5-16.0 mm 
(CAS49461. MZB 3833); Kapuas 1976-16, 15.2 mm (MZB 3834); Kapuas 1976- 
39. 84: 8.1-27.8 mm (CAS 49462. MZB 3835); Kapuas 1976-42, 8: 14.0-28.5 
mm (MZB 3836, USNM 23033 1 ); Kapuas 1 976-46. 4: 1 7.8-24.0 mm (MZB 3837. 
rSNM 230332); Kapuas 1976-47. 10: 14.5-20.2 mm (MZB 3838, USNM 230333). 

Calamiana sp. differs from C. magnoris (Herre 1935), the 
only other species of Calamiana exammed by me, in squa- 
niation and coloration. Calamiana sp. has scales in lateral series 
about 27, predorsal scales about 7, scales on abdomen as large 
as those on side on body, C magnoris and C. kabilia have scales 
in lateral series about 30-32, predorsal scales 1 1-12 (C. kabilia) 
or 1 5 (C. magnoris). scales on abdomen much smaller than those 
on sides. Calamiana sp. has posterior half of dorsal fin with 
solid black pigment and body with 4-5 broad, broken, vertical 
dark marks. Calamiana magnoris has fins weakly spotted and 
body more or less uniformly covered with fine mclanophores 
of variable size. Calamiana kabilia has median fins with well 
developed small spots in parallel rows and entire side of body 
with vertical streak in middle of each scale. Judging from size 
of specimens examined and degree of sexual dimorphism, C. 
sp. may be a smaller species. The holotype of C magnoris is a 
male 29.7 mm with sexual dimorphism very weakly developed 
compared to that of my largest specimens of C. sp. The two 
type specimens of C. kabilia are females 32.0-36.0 mm and 
therefore also far larger than my largest (20.1 mm) female C 
sp. 

Calamiana sp. exhibits pronounced sexual dimorphism of 
the sort found in many small Indo-Pacific gobiids in which males 
have greatly enlarged mouths. In males of this species the jaws 
posteriorly extend well beyond the eye, which is rather large. 



while in females the jaws do not extend posteriorly even as far 
as to below its anterior margin. In addition, the cheeks are 
greatly swollen and heavily ossified or bullate, and lower jaw is 
very heavily ossified in males. The behavioral significance of 
this dimorphism, found in many species albeit usually less pro- 
nounced, seems to be unknown. As is often the case in such 
sexually dimorphic gobies, the more extreme examples of which 
live in fresh water, males are larger than females. 

The CAS subset of 59 specimens from Kapuas 1976-39 (84: 
8.1-27.8 mm) includes 14 males with more or less marked 
sexual dimorphism, 18.4-27.2 mm; 7 gravid females 16.2-19.6 
(with oval eggs 0.7-1.5 mm long); 12 ripening females 15.0- 
20. 1 mm (with eggs to 0.5 mm long); and 26 sex undetermined 
8.1-18.4 mm. The largest female, 20.1 mm, has eggs 0.25 mm 
long. The largest gravid female, 19.6 mm has exactly 30 eggs, 
all 1.5 mm long. Observation of this and other females indicates 
that ovulation is synchronous; the full complement of eggs prob- 
ably is spawned at once or over a short penod of time; i.e., there 
is total spawning on an individual basis although by no means 
for populations as a whole. 

Mugilogobius Smitt, 1900 

.\tugilogohiiis Smut. 1900:522 (type species Clenogotviis abei Jordan and Snyder. 
1901. by subsequent designation of Jordan et al., 1913:345). 

Three small collections of subadult gobies were obtained in 
fresh water near the coast during the Kapuas survey of 1976 
representing two unidentified species of Mugilogobius. The 
species of this genus arc very poorly known (D. Hoese, pers. 
comm., 1984). 

Mugilogobius sp. undet. 

Material Examined —Western Borneo: Kapuas 1976-2. 18.1 mm (MZB 3830). 

Mugilogobius sp. undet. 

Material Examined —Western Borneo: Kapuas 1976-2. 5: 7.8-16.0 mm (CAS 
49460. MZB 3831); Kapuas 1976-3. 20 mm (MZB 3832). 



Pseudogobiopsis Koumans. 1935 

Pseudogobiopsts Koumans. 1935:131 (t>pe species Uohiop.sis oligacns Sleeker, 
1875. by original designation). 

The Kapuas survey of 1 976 obtained a small sample of gobies 
living in fresh water near the coast identified by D. Hoese as 
Pseudogobiopsis. 

Pseudogobiopsis jurongensis (Herre. 1940) new combination 

\\iii}]o\a !nn>ngt'ii\i\ Herrc, 1940ij:18 (t\pc locali[\ brook al Jurong, Singapore 
Island) 

Material Examined. — Malay Peninsula: Singapore Island, brook at Jurong, 
23: 18.7-35.9 mm (CAS-SLI 32982-3, holotype and paratypes). Western Borneo: 
Kapuas 1976-3. 6: 15.0-21 6 mm (CAS 49463. MZB 3839). 

The Kapuas specimens are very similar to the type specimens 
from Singapore in almost all respects but a substantial difference 
in size at sexual maturity should be noted. The Kapuas speci- 
mens include a sexually mature male of 21.6 mm and a gravid 
female of 20.0 mm. The male has checks markedly expanded 
and jaws markedly elongate, extending posteriorly to below pos- 



ROBERTS- WESTERN BORNEO FISHES 



169 



terior margin of eye. The female, with numerous eggs about 
0.20 mm diameter, has cheeks not expanded and jaws relatively 
short. The type series from Singapore comprises 16 mature or 
maturing males, 26.9-35.9 mm. 3 gravid females. 22.9-24.3 
mm; and 3 apparently immature specimens (possibly including 
spent females) 18.7-23.2 mm. In the males the jaws are more 
or less elongate, but only specimens of about 30 mm or more 
have jaws as elongate and cheeks almost as swollen as in the 
21.6 mm Kapuas male. The three gravid females all have nu- 
merous eggs about 0.15 mm diameter, noticeably smaller than 
in the gravid Kapuas female. 

Pseudogobius Aurich, 1938 

Pseudogobius Aunch, 1938:158 (type species Gobnis javaniciis Sleeker, 1856, by 
original designalion). 

The Kapuas survey of 1976 obtained a single small goby in 
fresh water well inland, kindly identified by D. Hoese as a Pseu- 
dogobius. 

Pseudogobius sp. undet. 

Material Examined. — Western Borneo: Kapuas 1976-51. 16,8 mm (MZB 3840). 

Stigmatogobius Bleeker, 1874 

Stigmalogobius Bleeker, 1874/):323 (type species Oobius picurosugma Bleeker. 
lS49=Gobius sadanimdio Hamilton-Buchanan. 1822. by original designation 
and monotypy). 

Stigmatogobius brocki (Herre, 1936) 

Vaimosa brocki Herre, 1936:9 (type locality Singapore harbor). 

Material Examined —Malay Peninsula: Singapore harbor (CAS-SU 30965, 
holotype). Western Borneo: Kapuas 1976-3. 18: 13.-3-34.2 mm (CAS 49464. MZB 
3841, USNM 230334); Kapuas 1976-5. 3: 10.4-27.8 mm (CAS 49465. MZB 
3842). 

Stigmatogobius sp. undet. 

Material E.xamined. — Western Borneo: Kapuas 1976-1. 3: 23.1-26.7 mm (CAS 
49466. MZB 3843), 

In morphology and coloration this species strongly resembles 
Stigmalogobius brocki. but there are obvious differences. Both 
species have the snout relatively pointed and jaws elongate and 
nearly horizontal, but in 5. sp. the snout is moderately pointed 
and the jaws extend posteriorly to below anterior third of eye, 
whereas in 5. brocki the snout is more strongly pointed and the 
jaws extend posteriorly almost to below posterior border of eye. 
In both species a vertical dark bar extends ventrally from dorsal 
fin onto body and there is a triangular peduncular spot (base of 
triangle vertical, apex directed posterioriy), but in 5. sp. the 
vertical dark bar extends well onto abdomen and the pendun- 
cular spot is very well defined and restricted to ventral half of 
caudal peduncle whereas in 5". brocki the vertical bar is confined 
to dorsal half of body and peduncular spot less well defined, in 
middle of peduncle adjacent to or overlapping caudal-fin base. 
Finally, 5. sp. has a dark spot in middle of anal-fin base lacking 
in 5. brocki but itself lacks fine melanophores evenly covering 
most of body characteristic of S. brocki. 

The small East Indian species Vaimosa (=Stigmatogobiusl) 
balleata Herre, 1935 (type locality Waigeu) with a similar dark 



vertical bar originating on the dorsal fin, differs from S. brocki 
and 5. sp. in having a very short truncate snout, and a large 
diagonal dark bar extending posteroventrally from hind border 
of eye across cheek and gill cover. 

Channidae 

Channidae or snakeheads are distinguished from other ana- 
bantoids by their elongate cylindrical bodies with long, entirely 
soft-rayed dorsal and anal fins, large mouth with well toothed 
jaws and palate, and relatively simple labyrinth organ. There 
are perhaps about 25 species, three in tropical Africa (perhaps 
constituting an endemic African genus, Paraphiocephalus Sen- 
na, 1924, as yet poorly characterized), the rest in Asia. All of 
the Asian species are placed at present in Channa. Some species 
are widely distributed, from the Indian subcontinent to south- 
east Asia and China, but the majority are endemic to one of 
these three areas. Southeast Asia has more species than any 
other area, and the Kapuas with nine species has more than any 
other river. 

Apparently most snakeheads produce pelagic eggs. A nest is 
made in a swampy area or backwater by making a clearing 
amidst dense grassy vegetation. The eggs are spawned in the 
cleared area, float to the surface, and are guarded by one or both 
parents. A more or less complex system of pathways may be 
cleared in the grass between the nest and open water. The parents 
may continue to guard the young until they are 50 mm or longer, 
and at least in some species the young are highly gregarious and 
remain in close proximity. In several Asian species juveniles 
are brightly colored, usually with red or orange on the side of 
the body, sometimes set off above and below by thin longitu- 
dinal black stripes. The size at which this juvenile coloration is 
lost ranges from about 50 to 450 mm, depending on the species. 
In at least some species these brightly colored young are highly 
gregarious and apparently form a dense school or ball if threat- 
ened (Breder and Rosen 1966; pers. obs.). See additional re- 
marks under C. micropelfcs. At least one species {Channa ori- 
entalis) practices oral brooding (Ettrich 1982). 

Channa Scopoli, 1777 

Channa Gronovius. 1763:135 (non-binomial; not available for zoological no- 
menclature according to Opinion 86, ICZN). 

Channa Scopoli, 1777:459 (type species Channa onentalis Bloch and Schneider, 
1 80 1 , by subsequent designation or subsequent monotypy of Bloch and Schnei- 
der, 180l:lvi, 496). 

OphiLcphalus Bloch 1 794: 1 37 (type species Ophicephalus punctams Bloch. 1 794). 

Ophiocephahis Hamilton-Buchanan, 1822:59 (emended spelling). 

Phdvpnoides Bleeker, 1849:19 (type species Phdypnoides siirakaricnsis Bleeker. 
li^9=Channa onentahs Bloch and Schneider. 1801. by monotypy). 

For nomenclatural discussion and other information on Ophi- 
cephalus see Myers and Shapovalov (1931). 

Channa bankanensis (Bleeker, 1852) 

Ophicephaliis bankanensis Bleeker. l852/:726 (type locality Banka). 
Ophicephalus bankanensis Vaillant. 1902:13, 16 (Pontianak, Sintang). 

Material Examined -Western Borneo: Pontianak, 197 mm (RMNH 7869); 
Sintang. 2: 104-123 mm (RMNH 7870), 

The 197 mm specimen from Pontianak has dorsal-fin rays 
39, anal-fin rays 29, lateral line scales 69, and rakers on first gill 



170 



CALIFORNIA ACADEMY OF SCIENCES 



arch 9; head convex dorsally; an oval black spot on opercle; 
pectoral fin with round white spots; dorsal surface of head with 
small round black spots, ventral surface of head and body with 
even smaller black spots; body with a faint irregular lateral band. 
It generally resembles the figure of C. hankanensis in Bleeker 
(1875-78, pi. 397. fig. 1). 
Distribution. — Borneo (Kapuas. Barito). Banka. 

Channa lucius Cuvier. 1831 

Ophicephahii lucnis Cuvier iii Cu\Ler and Valenciennes, 1831:416 (type locality 

Java). 
Ophioccphalus polylepis Bleeker. 18.'>2i":578 (type locality "Solok, Suniatrae oc- 

cidentalis, in fluviis"). See Weber and de Beaufort (l<)22:.327). 
Ophioccphalus hislriiitus Weber and de Beaufort. I422:.322 (type locality Sungei 

Manggar. Balikpapan. Sarawak). See Alfred (1964/i). 

Material E.XAMiNED. -Western Borneo; Kapuas l')76-l, 105 mm(MZB3909); 
Kapuas 1976-8. 22,1 mm (MZB 3910), Kapuas 1976-13, 108: 31,6-39.0 mm 
(CAS 49488, MZB 3911, LISNM 230348): Kapuas 1976-19,198 mm (MZB 39 1 2): 
Kapuas 1976-39, 129 mm (MZB 3913); Kapuas 1976-51, 120 mm (MZB 3914), 

Channa marulioides (Bleeker, 1851) 

Ophiccphahis mandinulcs Bleeker, 185 1:424 (t\pe locality "Sambas, in fluviis"). 
Material Examined. -Western Borneo: Kapuas 1476-20, 172 mm (MZB 3915), 

Distribution. — Sumatra (Deli, Djambi). Borneo (Sambas, 
Bankayan, Kapuas). Banka. Billiton. 



Material Examined, — Western Borneo: Kapuas near Putus Genteng. 183 mm 
(ZMA 114,391); Kapuas near Sanggau, 130mm(RMNH 26932), 

Channa niicropeltes. attaining nearly a meter in length and 
more than 20 kg, is the largest member of the family Channidae. 
.According to Smith (1945:474) it is very savage in guarding its 
nest and eggs and will even attack human beings if its nest is 
approached. It is also highly destructive of other fishes, con- 
suming fishes of all kinds and sizes and killing far in excess of 
Its actual needs. As first noted by Day ( 1 876), this species under- 
goes one of the most profound color transformations of any 
Asian freshwater fish. Young from about 30 mm to as large as 
450 mm have a pair of dark longitudinal stripes extending the 
length of the head and body to the end of the caudal fin. These 
bands arc usually reported as black but in some 50-60 mm fish 
observed by me in Thailand they were vivid ultramarine blue. 
The interspace between these lines is scarlet or bright orangish- 
red. In larger individuals this coloration is entirely replaced. A 
basically similar but less colorful color change occurs in C. stria- 
la but at much smaller sizes. 

Distribution. — West coast of southern India (Malabar, Ca- 
nara, Trichoor, mouth of Cochin R.). Burma?. Thailand. Malay 
Peninsula. Sumatra (Palembang, Lahat, Muara Kompeh, Gu- 
nung Sahilan, Djambi, Laut Tador. Danau Kota. Danau baru. 
Lake Toba). Borneo (Montrado. Sambas, Kapuas. Barito, Pra- 
bukarto). .lava (Batavia. Tjibiliong. Lelles, Samarang). Banka. 
Bilhlon. 



Channa melanoptera (Bleeker, 1855) 

Ophiccphahis mckm>plcrii\ Bleeker, l855i/:420 (type locality '■Pontianak, in flu- 
mine Kapuas"). 

Material Examined —None, 

Distribution. — Sumatra (Palembang. Indragiri, Gunung 
Sahilan, Djambi). Borneo (Kapuas). 

Channa melasoma (Bleeker, 1851) 

Ophiccphahis melasoma Bleeker. 185W;424 (type locality Sambas), 
Ophiccphahis rhodotacnia Bleeker, I85li:425 (type locality Sambas), See Weber 

and de Beaufort ( 1922:319-320), 
Ophiccphahis mystax Bleeker, 1853(/:I88 (type locality Banka) 
Ophioccphalus nwlaiiosoma Giinther, 1861:473, 
Ophioccphahis haramciisn Steindachner, 1901:3435 (type locality Baram), 

MaterialExamined— Western Borneo: Raocn, Kapuas basin, 5: 105-1 78 mm 
(RMNH 7862). 

Distribution,— Thailand (Menam Mun). Sumatra (Palem- 
bang, Labat. Gunung Sahilan. Ringgat, Laut Tador. Langkat). 
Borneo (Baram, Gambas, Sadong, Kapuas. Jesselton. Tempa- 
suk. Kinabatangan). Banka, Palawan Archipelago, 

Channa micropeUes (Cuvier, 1831) 

Ophiccphahis micropcltC' Cuxier in Cuvier and Valenciennes, 1831.427 (type 

locality Java), 
Ophiccphahis serpcnlinii\ Cuvier in Cuvier and Valenciennes, 1831:429 (type 

locality Siam), 
Ophiccphahis bnillaliis Bleeker, 1845:519 (type locality Batavia), 
Ophiccphahis Slercnsii Bleeker, I853i':444 (type locality Pontianak and Moara 

Kempeh) 
Ophioccphalus diplograminc Day, I865ij;36 (type locality Cochin); Day, 1865/': 

147, pi, 10, 
Ophioccphalus sludcri VoU, 1903tj:553 (type locality Palembang), 



Channa orientalis Bloch and Schneider, 1801 

Channa oncnlalis Bloch and Schneider, 1801:496, pi, 90, fig, 2 (type locality 

"India orientah"), 
Ophioccphalus gachiia Hamilton-Buchanan, 1822:68, 367. pi, 21, fig. 21 (type 

locality ponds and ditches of Bengal), 
For additional synonymy see Weber and de Beaufort (1922) and De Witt (1960), 

Pectoral fins with a series of curved vertical bars, typically 
SIX in number, bar nearest fin-base often expanded into a large 
basal blotch. Dorsal, anal, and caudal fins sometimes dusky or 
dark (even black) with a thin white margin; dark portion of 
median fins sometimes with faint vertical bars. 

Channa pleurophthalnius (Bleeker, 1851) 

Ophiccphahis pleiirophlhalmin Bleeker. 1851i:27() (type locality "Bandjermass- 

ing. in fluMis"), 
Ophiccphaliis urophthahmis Bleeker, 1852(/:578 (type locality "Palembang, in 

fluvus"). See Bleeker (1879:49). 
''Ophiccphahis spinlalis Fowler, 1904:530 (type locality Padang), Sec Weber and 

dc Beaufort (1922:324-325), 

Materlm-Exampmed,- Western Borneo; Kapuas 1976-20.4: 159-263 mm (CAS 
49489, MZB 3916, L'SNM 230349); Kapuas l«76-32, 127 mm (MZB 3917), 

Distribution. — Sumatra (Palembang, Djambi, Padang?). 
Borneo (Kapuas, Barito). 

Channa striata (Bloch, 1793) 

Ophiccphahis slrialiis Bloch, 1793:141, pi 359 (i\pe locality Malabar), 
For additional synonymy see Weber and de Beaufort (1922:317), 

Material Examined —Western Borneo: Kapuas 1976-2, 44.8 mm (MZB 3918); 
Kapuas 1476-43, 103 mm (MZB 3919). 

Channa striata is a very important food fish and its range 
undoubtedly has been expanded by introductions. It is now 



ROBERTS-WESTERN BORNEO FISHES 



171 



found throughout most of the East Indies, the Malay Peninsula, 
and Thailand, and in India. Sri Lanka, and the Philippines. 

Anabantidae 

The anabantoid family Anabantidae or climbing perches 
comprises three genera. Ctenopoma and Sandelia in Africa and 
Anabas in Asia. 

Anabas Cuvier. 1816 

Anahas Cuvier, 1 8 1 6:339 (type species .Uilluas lesluclinciis Bloch. 1 795, by mono- 
typy). 

Anabas testudineus (Bloch, 1795) 

Anthms lestudmeus Bloch, 1795:121 (type locality Japan, southern India). 

Perca scandens Daldorff. 1797:62 (not seen by me), 

Anahas lesludmeus Cuvier, 1816:339. 

Anahas scandens Vaillant, 1902:13 (Pontianak). 

For additional synonymy see Weber and de Beaufort (1922:334). 

Material Examined. — Western Borneo: Pontianal<. 88.8 mm (RMNH 7873). 

Anabas. the climbing perch, now generally regarded as com- 
prising a single species, is exceptionally variable. Body form 
varies from oval and compressed to elongate and subcylindrical. 
Meristic variation has also been noted (e.g.. Weber and de Beau- 
fort 1922:336 on geographical variation in number of dorsal- 
fin spines). Further study may reveal more than one species (see 
Dutt and Ramaseshaiah 1980, 1983). 

Anabas are among the hardiest of fishes. The suprabranchial 
cavity is exceptionally large and the labyrinth it contains is 
complexly folded. In some of the more elongate subcylindrical 
morphs the paired suprabranchial cavities are so greatly en- 
larged that they are externally evident as bulla-like swellings on 
the dorsal surface of the skull. 

Belontiidae 

The family Belontiidae. although currently recognized, prob- 
ably IS not distinct from the next family. Osphronemidae. As 
currently understood. Belontiidae is the largest and most diverse 
family of anabantoids. with 1 1 genera and more than 30 species, 
and is distributed throughout the Indian subcontinent, southeast 
Asia, and much of China. Five genera and 1 1 species are known 
from western Borneo, 

The belontiid Trichopsis vittata (Cuvier in Cuvier and Va- 
lenciennes. 1831) is recorded from Sintang by Weber and de 
Beaufort (1922:351). Presumably this is based on the listing of 
its junior synonym Osphwmenm sirialus (Bleeker. 1850) from 
"le Kapoeas de Sintang jusque Boenoet" by Popta (1906:255). 
I know of no other records of this otherwise common and rather 
widely distributed southeast Asian species from western Borneo, 
and doubt that it occurs there. Popta's record presumably would 
have been based upon material collected by the Dutch expe- 
ditions to central Borneo of 1896-1900 and deposited in the 
RMNH. A recent search for this material has been unsuccessful 
(M. S. Hoogmoed, pers, comm.. August 1984). 

Belontia Myers, 1923 

Belontia Myers. 1923:63 (type species Pulyacanthus hassclti Cuvier in Cuvier and 
Valenciennes, 1831, by original designation). 



This genus includes two species and has a disjunct distribu- 
tion, with B. hassclli in the East Indies and Thailand and B. 
signata (Giinther. 1861) in Sri Lanka. B. signata is recorded 
from Java by Day (1878:371) and Jayaram (1982:382) but the 
basis of this record is not stated; the species was not mentioned 
from Java by Bleeker or by Weber and de Beaufort (1922). 

Belontia hasseiti (Cuvier. 1831) 

Polyacamhus Hassclli Cuvier in Cuvier and Valenciennes, 1831:353 (type locality 

Java). 
Polyacantluis Kiihli Bleeker, 1845:520 (nomen nudum). 
Polvacanlhiis Einlhorcnii Bleeker. 185l/:423 (type locality "Sambas, Borneo oc- 

cidentahs, in fluviis"). 
Polvacamhus Helfrichii Bleeker. 1855^:162 (type locality "Bandjermasin. in tlu- 

\'11S")- 

Bclonlia hassclli Myers. 1923:63. 

Material Examined. — Kapuas 1976-4, 62,5 mm (MZB 3844): Rapuas 1976- 
16. 3: 40.1-67.1 mm (C.^S 49467. MZB 3845). 

Distribution. — Malay Peninsula (Perak. Johore. Singapore). 
Sumatra (numerous localities cited by Weber and de Beaufort 
1922:339). Borneo (Sambas. Kapuas. Kahajan. Barito). Java. 



Betta Bleeker, 1850 

Bella Bleeker. 1850:12 (type species Bella Infasciala Bleeker, \i'iO=Panchax 
picliim Valenciennes in Cuvier and Valenciennes. 1846, by monotypy), 

Micracanthus Sauvage, 1879:95 (type species Micracanlhiis marchei Sauvage. 
\%l%=Bella splcndcns Regan, 1910, by monotypy). See Roberts (1981), 

Parophiocephaliis Popta, 1905:184 (type species Parophwceplialiis iinimaculaius 
Popta, 1905, by monotypy). 

Oshimia Jordan. 1919:342 (unncedcd replacement name for Mieraeanlhus Sau- 
vage. 1878. supposed by Jordan to be preoccupied b\ Microcanlhus Swainson. 
1839). 

Psciidohella Richter. 1981:273 (type species Bella piignax Cantor. 1849. by orig- 
inal designation). 

It has long been recognized that some species of Bella are 
oral brooders while others are bubble nesters. Whether all species 
fall neatly into one of these two categories is not yet known, 
however. Recently, Richter has proposed that the bubble nesting 
species should be left in the genus Bclta and proposed a new 
genus, Pseudobelia. for the oral brooders. This is contraindi- 
cated because the type species of Betta. B. picta. is itself an oral 
brooder (Vierke 1981; Richter 1983). Another problem which 
argues against the designation of a separate genus for the sub- 
strate spawners is the probability that oral brooding and sub- 
strate spawning are not evolutionary "fixed" or stable characters 
in Betta but have been selected for repeatedly in different species. 
In this case generic division of Betta based on brood care would 
inevitably result in polyphyletic taxa and an unnatural classi- 
fication. At least for the time being it seems best to recognize 
only a single genus. Betta. of which there seem to be about 1 5 
species, some oral brooders and others substrate spawners. 

With the notable exception of the long-jawed and very dis- 
tinctively colored B. macrosloma. preserved specimens of 5f //a 
are notoriously difficult to identify to species. Some have dis- 
tinctive coloration in life, but color is difficult to use because it 
is so changeable depending on sex, maturity, reproductive con- 
dition or "emotional" state, perhaps geographical vanation, and 
certainly variation related to method of preservation. Meristic 
characters are of little help because the genus is morphologically 
relatively conservative, and probably because species vary geo- 



172 



CALIFORNIA ACADEMY OF SCIENCES 



Table 14. Frequencies of Vertebral Counts in Betta. For definition of vertebral counts sec p. 22 



Abdominal 



Id II 



Caudal 



Total 



18 l>) 20 21 



1-) T? 1J TO 1 



23 24 



■^ 30 31 32 33 34 35 



akarcnsii Regan, l'*IO(Akar R., holotype) 
iiiiuhutoiiics Bleeker, l850(Kapuas 1976-23) 
inccina Vicrke. 1979 (Djambi, types) 
dimiilmla new species (Kapuas. types) 
foerschi Vierke, 1979 (Mentaya R., types) 
macwstoma Regan. 1910 (Brunei near Labi) 
/i».?nii-\ Cantor. l850(Penang) 
pui^nax Cantor. 1850 (Kapuas) 
cf. iM'iiiula Regan, 1910 (Kapuas 1976-1) 
uninmciiUild (Popta. 1906) (Balung R.) 



57 
3 

15 
10 
II 



II 47 



13 3 

2 

II 



1 


") 


9 


47 




14 


7 


2 




II 



graphically more than generally recognized. Thus the largest and 
smallest species oi Betta have almost identical counts of fin rays 
and scales. Species distinctions based on presence or absence of 
small spines in the dorsal and anal fins and the number and 
thickness of such spines are not so reliable as previously sup- 
posed by some workers. Differences in morphometric features 
such as body proportions (particularly head length and body 
depth) and fin shapes seem to distinguish certain species but are 
difficult to define objectively and may vary geographically. Ver- 
tebral counts have not been used previously in defining species, 
and I have gathered some data to see whether they might be 
helpful (Table 14). The frequencies of vertebral counts seem to 
be very distinctive for B. unimaculala. and help to distinguish 
B. dimidiala new species. In the present state of knowledge it 
is not possible to discuss with confidence the geographical dis- 
tribution of the commoner and more widely distributed species 
oi Bctta. On the other hand, within single river basins, where 
four or more species may occur, they can usually be more or 
less readily distinguished by differences in color and morphol- 
ogy, although the species names which should be applied may 
be in doubt. There are four species in western Borneo. 

Betta anabatoides Bleeker, 1851 

(Figure 129a) 

Bella anahalindc Bleeker, 1 85 1 r: 269 (type locality "liandjermassing, in fluviis"). 

Material Examined.— Western Borneo; Kapuas 1976-1, 2: 46.8-59.6 mm 
(BMNH 1982.3,29,229, MZB 3846), Kapuas 1976-23. 8: 23.3-43.4 mm (CAS 
49468. MZB 3847. RMNH 28916. HSNM 230335); Kapuas 1976-30. 59.2 mm 
(MZB 3848). 



Betta dimidiata new species 

(Figure 130) 

Bella akarcnsi\ Vierke, 1983; 166 (Kapuas), 

Holotype, — MZB 3849, 31,7 mm (ripening female), Kapuas basin, Sungai 
Seriang, 37 km W of Putussibau (Kapuas 1976-42). 

Paratypes.-CAS 49469 (9), MZB 3850 (3), 12; 22,4-30,2 mm, same locality 
as holotype; MZB 3851 (2), USNM 230336 (7), 9: 16,8-28.9 mm, Kapuas hasm, 
Sungai Tekam on right side of Kapuas mainstream 5-6 km upstream from Sanggau 
(Kapuas 1976-16); CAS 49470 (3), MZB 3852 (1), 4; 20,9-34,3 mm, Kapuas 
basin, small streams flowing into Kapuas mainstream wnhin 10 km of Sanggau 
(Kapuas 1976-17); BMNH 1982,3,29,230-235 (6). FMNH 94251 (6). MNHN 
1982-718 (5). MZB 3853 (7), 24; 9,5-23,3 mm, Kapuas basin, Danau Piani near 
Ketungau. 38 km NNE of Sintang (Kapuas 1976-32); CAS 49471 (2). MZB 3854 
( I ), 3; 23.6-35,0 mm, Kapuas basin, Sungai Mandai Kctchil near confluence with 
Kapuas mainstream 18 km WSW of Putussibau (Kapuas 1976-39); MZB 3855 
(1), RMNH 28917(2), 3; 27.0-29.0 mm, Kapuas basin, small oxbow lake opposite 



Empangau, 124 km NE of Sintang (Kapuas 1976-43); MZB 3856 (1), USNM 
230337 (5), 6: 23.0-28.5 mm. Kapuas basin. Sungai Genlu where it flows into 
Kapuas mainstream 55 km NE of Sintang (Kapuas 1976-46); CAS 49472 (6), 
MZB 3857 (I), 7; 20.7-24.8 mm. Kapuas basin, small forest stream flowing into 
Kapuas mainstream NE of Gunung Setunggul, 53 km NW of Sintang (Kapuas 
1976-47); MZB 3858 (1), UMMZ 209927 (5), 6; 14.3-27.4 mm. Kapuas basin, 
small forest stream near Tajan. 87 km E of Pontianak (Kapuas 1976-51). 

Diagnosis. — Srt/a dinudiala is possibly the smallest species 
of Bctta (largest of 75 specimens from nine samples only 35.0 
mm); scales in lateral series 27-29; transverse scale rows 8-9, 
apparently fewer than in any other species (with no "half scales" 
or with at most one interposed scale on dorsal-fin base and one 
on anal-fin base); dorsal-fin origin opposite 1 1th- 14th scale in 
lateral series; scales between eyes 2 or 3; dorsal fin with 1-3 
(usually 2) and anal fin with 1-2 (usually 1) spines; total dorsal- 
fin elements 9-1 l.anal 23-27; pectoral-fin rays 12-15; vertebrae 
10 (rarely 1 1)-H 9-2 1=29-3 1, with a strong mode at 30 (Table 
14). 

Nearly all Bctta species have at least 1 1 transverse scale rows. 
The species nearest B. dimidiata in this character apparently is 
Bctta akarcnsis Regan. 1910, which may also be its closest 
relative. The holotype of 5. akareiisis has 10 transverse scale 
rows on the body proper plus a row of sheath scales along the 
dorsal fin and the anal fin giving a total count of 12 if these are 
included; the number of scales from the gill opening to below 
the dorsal fin seems to be 1 2 or 13 (as in B. dimidiata), but 
these arc difficult to count because some scales are missing on 
both sides (I am indebted to O. Crimmen and A. Wheeler for 
scale counts on the holotype of B. akarcnsis). In fin formulae 
and vertebral counts B. akarcnsis again difliers from B. dimi- 
diata. As indicated by Regan (1910:779), and confirmed by 
examination of radiographs, the holotype of B. akarcnsis has 
no dorsal-fin spines, and only a single anal-fin spine; the total 
number of dorsal-fin supporting elements is 8 (one less than in 
any specimens of 5. dimidiata) and of anal-fin elements 28 (one 
more than in B. dimidiata); this is correlated with the scale 
count of 31 reported by Regan, also higher than observed in B. 
dimidiata. The number of scales between the eyes is also fewer 
in B. dimidiata than in other Bctta I have examined, but com- 
parative data for this character are not available for B. akarcnsis 
and most other species, Bctta akarcnsis probably attains a larger 
size than B. dimidiata; the holotype and only known specimen 
is 37.0 mm. 

Frequencies of vertebral counts in B dimidiata are presented 
in Table 14, Frequencies of dorsal-fin elements are 9(25), 10(36), 
11(4) and of anal-fin elements 23(3), 24(11), 25(31), 26(23), 



ROBERTS-WESTERN BORNEO FISHES 



173 




Figure 129. Bella, a, B anahmoides. Kapuas 1V76-I. 59.6 mm (MZB 3846); b, B pugna.\. Kapuas 1976-51, 46. b mm (MZB 3866). 



27(2). Head length 2.7-2.9; eye diameter 9.7-10.9; bony inter- 
orbital width 11.7-12.6; snout 11.6-14.1, mouth e.xtending pos- 
teriorly to a point below and slightly in front of anterior margin 
of eye; body depth at dorsal-fin origin 4.2-4.7. Filamentous ray 
of pelvic fin in both sexes extending posteriorly about one-third 
or at most one-half of length of anal-fin base. Anal fin with 
filamentous termination extending to below about middle of 
caudal fin in both sexes. Caudal fin not notably rounded or 
expanded, upper rays extending slightly beyond lower rays in 
some specimens. 

Body usually with one or sometimes two broad longitudinal 
dusky stripes or in some specimens (males?) with three some- 
what narrower and darker longitudinal stripes. Female holotype 
with a single broad stripe commencing below snout-tip (contin- 
uous around chin), extending across opercular region and on 
ventral half of body to caudal-fin base; this stripe is narrower 
but better defined on head. Some specimens have a second broad 
longitudinal stripe beginning behind head and extending length 
of body; this is usually somewhat less well defined than stripe 
on ventral half of body. Some specimens have a longitudinal 
stripe on head below eye which does not extend onto body, 
while in a few this extends as a third narrow stripe for entire 
length of body. Dorsal-, anal-, and caudal-fin rays and inter- 
radial membranes with more or less numerous, uniformly dis- 
tributed (crowded or contacting) and very numerous (crowding 
or contacting each other) chromatophores (reddish or reddish- 
brown in life?). Mode of brood care unknown. 



Etymology.— The name dimidiata (Latin, halved or dimin- 
ished) refers to the small size of the species. 



Betta pugnax (Cantor, 1850) 

(Figure 129b) 

Macropodus pugnax Cantor, 1850:1066 (type locality Pinang, Malay Peninsula). 
Bella pugnax Regan, 1910:779, 
Pseudohella pugnax Richtcr. 1981:273. 

Material Examined. — Malay Peninsula: Pinang, 18: 27.4-63,0 mm (CAS-SU 
33181). Western Borneo: Kapuas 1976-6, 2: 32.4-46,1 mm (CAS 49473, MZB 
3859); Kapuas 1976-13, 32,8 mm (MZB 3860); Kapuas 1976-16. 2: 14,6-18.6 
mm (CAS 49474, MZB 3861); Kapuas 1976-17, 3: 25,3-56,8 mm (BMNH 
1982.3,29,236-237, MZB 3862); Kapuas 1976-37, 5: 12,7-»6.4 mm (MZB 3863, 
USNM 230338); Kapuas 1976-39, 10: 39,2-59,8 mm (MNHN 1982-719, MZB 
3864. RMNH 28918); Kapuas 1976-42. 2: 41.5-50.0 mm (MZB 3865, UMMZ 
209908); Kapuas 1976-51. 2: 37.6-46.8 mm (MZB 3866, ZMA 1 16.547). 

Comparison of the well preserved Kapuas and Pinang samples 
reveals their close similarity in all respects and no reason to 
doubt their conspecificity. In large males of this oral brooding 
species from Penang and from Kapuas the gill cover and entire 
ventral portion of head (including branchiostegal membranes) 
is very darkly pigmented. 



Betta cf taeniata Regan, 1910 

Bella laemala Regan. 1910:781 (type locality River Senah, Sarawak). 
':Beua edillhie Vierke. 1984:60 (type locality Barito Delia bei Banjarmasin). 



174 



CALIFORNIA ACADEM'i OF SCIENCES 




Figure 130. Bctlii diniidiaia Rapuas 1976-42. 31.7 mm female (MZB 3849, holotype). 



Material Examined. — Western Borneo: Kapuas 1976-1, 31: 22.5— 16. 7 mm 
(BMNH 1982. 3. 29. 238-241, CAS 49475, MNHN 1982-720, MZB 3867, RMNH 
28919, USNM 230339); Kapuas 1976-4, 5: 17.7-31.8 mm (MZB 3868, UMMZ 
209852); Kapuas 1976-5. 34.9 mm (MZB 3869); Kapuas 1976-7, 35.5 mm (MZB 
3870); Kapuas 1976-8, 12: 17.3-34.2 mm (C\S 49476. MZB 3871). 

This species is (distinguished from other Kapuas Bctta by 
having regularly punctuated dark marks on the interradial mem- 
branes of the dorsal and caudal fins (particularly marked on the 
latter). Most specimens have up to five interrupted longitudinal 
stripes on the body, the lowermost two characteristicallyjoining 
or fusing together on about the posterior one-third of the body. 
Scales between eyes about 5. 

Betta edithae is distinguished from B. taeniata mainly by 
having its anal fin with ii27-28 instead of only ii20-22 rays. 
My material from Kapuas 1 976- 1 has anal-fin rays ii26(2), ii27(3), 
ii28(3). ii29(3) and thus in this respect agrees with the descrip- 
tion of B. edithae better than with that of B. taeniata. Clearly 
more information is needed on geographical distribution and 
meristic variation in these species, one or both of which must 
be very widely distributed. 

Parosphromenus Blecker, 1879 

Pannphronieniis Blocker, 1879:19 (type species Osphroinenus dassnen Sleeker, 
1859, by monoKpy). 

Parosphromenus includes four species: P. deissneri from the 
Malay Peninsula, Banka, and western Borneo; P. filainentosiis 



Vierke, 1981 (closely related to P. deissneri). from southeastern 
Bprneo; P. paludicola, from the Malay Peninsula at Trengganu; 
and P. parvuhts from western and southeastern Borneo. The 
genus presumably occurs in Sumatra but has not been reported 
there, presumably due to insufficient collecting of small fishes 
in suitable habitats. Meristic data on the species are presented 
in Table 15. 

Parosphromenus deissneri (Bleeker. 1859) 

Osphronrcniis Dcnsncn Blecker, 1859^:377 (type locality Banka). 
Parosphromenus Deissneri Bleeker. 1879:20. 

Material Examined —Malay Peninsula: Johore. 2: 20.7-26.4 mm (CAS-SU 
34833); .lohore. Kulai, 2; 19.8-19.9 mm (CAS-SU 39449). Western Borneo: Ka- 
puas 1976-4. 24.7 mm (MZB 3872); Kapuas 1976-5. 12: 12.4-24.5 mm (CAS 
49477. MZB 3873, USNM 230340); Kapuas 1976-7, 4: 9.3-23.0 mm (MZB 3874, 
RMNH 28920); Kapuas 1976-8, 3: 16.9-25,6 mm (BMNH 1982.3.29.242-243); 
Kapuas 1976-51. 22.6 mm (MZB 3875). 

Distribution. — Malay Peninsula (Malacca, Johore). Western 
Borneo (Kapuas). Banka. 

Parosphromenus parvulus Vierke, 1979 

(Figure 131) 

Parosphromenus parvulus Vierke. 1979:247 (type locality "Mentaya-Fluss-Sys- 
tem, 250 km nordwestlich von Banjarmasin"). 

Material E.xamined. — Western Borneo: Kapuas 1976-4. 16.8 mm (MZB 3876); 
Kapuas 1976-5, 2: 18.9-19.6 mm (CAS 49478, MZB 3877). 



Table 15, Meristic Characters in P^ROS/'HfiOAjfA't'.s. 



P deissneri 
Johore (CAS 34833) 
Kapuas 
Bangka (holotype, after Bleeker) 

P filamemosus 

SE Borneo (types, after Vierke) 
P. paludicola 

Trengganu (types, after Tweedie) 
P parvulus 

Kapuas 

Mentaya (types, after Vierke) 



Dorsal-hn spines -frays 



11-12 + 7=18-19 

12 + 6-7=18-19 

13 + 7 = 20 



12-13 + 6-7=19-20 
17-18 + 6-7 



10 + 6=16 
10-11 + 7=I7-U 



.Anal-hn spines + rays 



12+10=22 

11-12 + 9-11=21-23 

13 + 9 = 22 



11-12+10 = 21-22 



15 + 7 = 22 



7+12-13=19-20 
8-9+10-11 = 18-20 



Scales in lateral 
series 



27-28 

28-29 

30 



29-30 
30 



26-27 
26-27 



ROBERTS-WESTERN BORNEO FISHES 



175 




Figure 131. Parosphromenus- Above. P pamiluy Kapuas 1976-4. 16.8 mm (MZB 3876); below, P dcissiieri. Kapuas 1976-5. 21.9 mm (CAS 49477), 



Distribution. — Known only from Borneo (Kapuas, Men- 
taya). 

Sphaerichthys Canestrini, 1860 

Sphacnchihvs Canestrmi. 1860:707 (t\pe spccrcs Sphucnchllivi oiphroDwnoidi's 
Canestnni, 1860, by monotypy). 

Small, body deep, compressed. Jaws with very small, fixed 
teeth. Dorsal-fin spines 6-10; anal-fin spines 7-1 1. Pectoral-fin 
rays 10-12. Scales in lateral series 28-30. Preorbital and pre- 
opercle serrate, other head bones non-serrate. Lateral line ves- 
tigial or absent. Swim bladder extending almost to caudal fin, 
obliquely truncate posteriorly. Pelvic fin with a spine and five 
well developed rays; first ray sometimes very elongate. Caudal 
fin obliquely truncate, caudal-fin rays 7-8 + 7-8=14-16. Total 
vertebrae 25-28. 

This distinctive anabantoid genus comprises four species (all 
present in Borneo); 5. osphromenoides, S. vaillanli. S. acrostoma 
Vierke, 1979, and S. selantanensis Vierke, 1979 (originally de- 



scribed as a subspecies of 5. osphromenoides but here treated 
as a full species). S. osphromenoides occurs in Malay Peninsula, 
Sumatra, and Borneo; the other three are known only from 
Borneo. 5. osphromenoides and 5. vaillanli occur in western 
Borneo (Kapuas) and are treated in more detail in the following 
pages. The key provided here includes all of the species but is 
based on relatively few specimens. 

Key to Sphaerichthys 

la Body relatively deep; snout short; dorsal-fin origin almost 
directly above anal-fin origin; total dorsal-fin elements 
15-20; anal-fin spines 7-9; total vertebrae 25-26 2 

1 b Body relatively elongate; snout elongate; dorsal-fin origin 
considerably posterior to a vertical through anal-fin ori- 
gin; total dorsal-fin elements 13-15; anal-fin spines 10- 
1 1; total vertebrae 27-28 3 

2a Dorsal-fin spines 8-11; no pale vertical bar extending 
from dorsal-fin origin to just behind pelvic-fin origin; no 



176 



CALIFORNIA ACADEMY OF SCIENCES 



pale longitudinal stripe extending length of body 

5. osphromenoides 

2b Dorsal-fin spines 7; pale vertical bar extending from dor- 
sal-fin origin to just behind pelvic-fin origin; pale longi- 
tudinal line extending length of body (known only from 
Barito) S. selateuensis 

Ventral two-thirds of body with 6-8 vertical or slightly 
oblique dark bars; anal-fin rays 16-18 5. vaillanli 

Body without 6-8 dark bars; anal-fin rays 20-21 (known 
only from Mentaya) S. acrostoma 



3a 



3b 



Sphaerichthys osphromenoides sensu Regan, 1910 

'!Sphaenchlhys osphromcnoklcs Canestnni. 1 860:707 (lype locality "aus Indien"). 
Osphromemis malayamis Duncker, 1904:163, lable I. Iig. 8 (type locality Kuala 

Lumpur). 
Sphaerichthys osphronwiionlt's Regan, l'-^10:776. 

Material Examined. — Malay Peninsula: Kuala Lumpur, 3: 19.4-33.3 mm 
(HZM 8310. syntypes of O nwlciyanii!.): Simpang Rengani. Johore. 42.2 mm 
(CAS-SU 3945 1 ); Kulia. Johore, 18: 14.4-37.4 mm (CAS-SLI 39450); Kota Tinggi, 
Johore, 2: 24.9-25.4 mm (CAS-SU 33864); Singapore, 22: 13.0-32.3 (CAS-SU 
19993). Sumatra: Taluk, 2: 28.7-30.6 mm (ZMA 114,386); Bagan Api Api, 14: 
19.9-34.9 mm (ZMA 1 14.385); GunungSahilan, 13: 10.4-28.2 mm (ZMA 114.384); 
DjamhI. Batang Han, 30,8 mm (ZMA 114.387). Borneo: Kapuas 1976-4, 23,9 
mm (MZB 3878); Kapuas 1976-5. 5: 12,1-29,9 mm (CAS 49479, MZB 3879, 
USNM 230341), 

Having made direct comparisons of specimens of this species 
with the types of .S', osphromcnnidcs sclalancnsis. I am inclined 
to regard the latter as a distinct species. In addition to the char- 
acters indicated in the key above, 5. selatancmis has a thicker 
head, wider mouth, and blunter snout than 5, osphromenoides 
of the same size. 

Sphaerichthys osphromenoides carries the eggs in its mouth, 
as I myself once observed during fieldwork in Johore. The re- 
productive and parental behavior of its congeners is unknown. 

Important Note. — It is conceivable that Regan's identifi- 
cation of Sphaerichthys osphromenoides Cancstrini, 1860 is in- 
correct, and that the species under consideration here should 
be known as Sphaerichthys malayanus (Duncker, 1904). Ca- 
nestrini's original description may be translated as follows: 

Sphaet'ichihys mihi no\'. gen moderately long dorsal fin originating 
over the origin of the anal fin: anal fin longer [than dorsal) and joined 
with the caudal fin. First soft pelvic-fin ray only slightly elongated, the 
others normally developed, not rudimentary. Pectoral fins normally 
developed, caudal fin pointed (acuminate), Opercle entire, preopercle 
with spines at angle and on its horizontal margin. Subopercle and 
interopercle margins entire, preorbital spiny. Mouth opening small, 
mouth protrusible. Jaw teeth uncommonly small, barely visible; vomer 
and palate smooth (toothless). Head and nape covered with scales, 
scales of nape finely ciliate (ctenoid). Small pores on head. Lateral line 
consisting of a row of fine pores barely visible to the naked eye. Six 
gill rakers. Two nostrils on each side, nostrils separated from each 
other by less than their diameter Labynnth organ originating from 
some of the anterior gill arches, with dorsoantenorly projecting leaflets, 
(laminae). 

Swim bladder ("shining through") extending up to tail (tail fin''). 
Dorsal profile of head (stimprofil) slightly concave. A true intermediate 
genus (connecting link) between Ospliromvinn and irhhopviliis 

Sphaerichthys osphromenoides mihi 

Head 2-1/2 in body length (without caudal). Eye diameter 3-1-2 in 
head length and greatest depth of body 1-2/5 in length without caudal. 
Number of scales in a longitudinal row from opercle to caudal about 
26. in a transverse row at the greatest depth about 12. 

The dorsal and anal spines are laterally compressed and increasingly 



longer posteriorly. Judging from the strongly stained alcohol specimens 
the ground color is dark brown (castanienbraun). the fins e.vcept the 
pectoral are dark. Dorsal half of body at caudal-fin base with a dark 
spot. Two white cross-bars (vertical bars) on posterior half of body 
extending between the dorsal and anal fins, of which the first ends at 
the base of last three anal-fin spines and the second at base of 7th- 
1 0th anal-fin soft-rays. The lower half of the second white bar is banded 
(eingefasst) by two dark marks, which barely extend dorsally to the 
lateral line. 

The two specimens of this species at hand come from the Indies 
and their length without caudal measures not quite 1-1/2 Viennese 
inches and their greatest height almost 10 lines, 

Regan (1909:776), without comment and without examining 
Cancstrini's types, placed O. malayanus in the synonymy of 5, 
osphromenoides. Duncker (1904:164) suggested that the diflTer- 
ences between his description of Osphromemis malayamis and 
Cancstrini's of Sphaerichthys osphromenoides might be due to 
racial differentiation or inaccurate observation; this probably 
influenced Regan. I am inclined to credit the accuracy of Ca- 
ncstrini's observations; now that several species of Sphae- 
richthys are known it seems likely that Cancstrini's and Dunc- 
ker's specimens belong to difltrent species. The original description 
does not permit identification of Sphaerichthys osphromenoides 
Cancstrini, 1860 with any known species. Although some parts 
of the description agree with Sphaerichthys and 5, osphrome- 
noides of Regan et seq., other points contradict this conclusion. 
The strongest of these are: 

1. Dorsal-fin spines 12 (vs. 7-10). 

2. Depth of body 1-2/5 in length of body without caudal (vs. 
more than 2). 

3. Transverse scale rows 12 (vs. 14-15). 

Some other seeming discrepancies should be commented upon. 
The expanded caudal fin is obliquely truncate, but when the 
caudal-fin rays are adpressed may appear to be acuminate. Ca- 
ncstrini's remarks seem to imply that his fish has a complete 
lateral line; if so. it is unlike any known species of Sphaerichthys 
sensu Regan, 1909, in which lateral line is absent or is repre- 
sented by a few pored scales on the posterior half of the body. 

Cancstrini remarked on the possible identity of his Sphae- 
richthys osphromenoides with the poorly known "Osphromemis 
notatiis K,v,H." and Ctenops nobilis McClelland, specimens of 
which were unavailable for him to examine. The former is prob- 
ably based on young Osphromemis goramy. while the latter is 
a valid genus and species diflcring in numerous respects from 
the fishes under consideration (see Liem 1965), But the com- 
parison with "O. notatiis" with "dorsal fin 13/1 1" strongly sug- 
gests that the notation "D 1 2/7-8" for S. osphromenoides is not 
a typographical error. 

The two type specimens (syntypes) of 5, osphromenoides pre- 
sumably are deposited in the Naturhistorisches Museum Wien, 
the ichthyological types and other material of which are tem- 
porarily unavailable for study, Mr. Harald Anhelt searched un- 
successfully in the NMW for the syntypes of S. osphromenoides 
in June 1984. 

Distribution. — Malay Peninsula (Negri Sembilan, Johore). 
Sumatra (Bagan Api Api, Batang Han, Djambi, Taluk, Gunung 
Sahilan). Borneo (Kapuas). I have seen specimens from Billiton 
(2: 22.4-27.5 mm, ZMA 114.388) which may also belong to 
this species but did not study them sutficiently to be sure of 
their identification. So far as I have been able to determine this 
is the only Sphaerichthys occurring outside Borneo. 



ROBERTS-WESTERN BORNEO FISHES 



177 



Sphaerichthys vaillanti Pellegrin, 1930 

OsphromeniLS nobdis VaiUant. I8<)3;102 (not of McClelland. 1844; specimens 

from Kapuas). 
Sphaerichlhrs vatllami Pellegrin. 1>J30:243 (type locality Kapuas). 

Material Examined. -Western Borneo: Kapuas, 2: 27.7-29.2 mm (MNHN 
1891-510-11, syntypes); Kapuas 1976-16, .30,3 mm (MZB 3880); Kapuas 1976- 
32, 22,9 mm (MZB 388 1 ); Kapuas 1 976-43, 2; 27,5-27,9 mm (CAS 49480, MZB 
3882), 

This species is known only from the Kapuas, It is closely 
related to S, acrostoma from the Mentaya. 

Trichogaster Bloch and Schneider, 1801 

Tnchogasler Bloch and Schneider, 1801:164 (type species Lcihnis irichoptenis 
Pallas, 1770, by subsequent designation of Cuvier in Cuvier and Valenciennes, 
1831:388), 

For additional synonymy see Myers (1923/i), 

This southeast Asian genus comprises four species, two of 
which occur in western Borneo, 

Trichogaster leerii (Bleeker, 1852) 

Tnchopus Leerii Bleeker, I852(/:577 (type locality Palembang, in fluviis). 
Trichopodus leeri Regan, 1909:783, 
Trichogaster leeri Myers |923/>:l 12. 

Material Examined. — Western Borneo: Kapuas 1976-16, 4: 39,0-69,3 mm 
(CAS 49481, MZB 3883); Kapuas 1976-17, 2: 40,5-57,7 mm (BMNH 
1 982,3,29.244, MZB 3884); Kapuas 1 976- 1 9, 8; 63,4-72,3 mm (CAS 49482, MZB 
3885); Kapuas 1976-20, 70.7 mm (MZB 3886); Kapuas 1976-32, 3: 57.5-71.2 
mm (MZB 3887, RMNH 28921); Kapuas 1976-33, 2: 57.8-63.4 mm (CAS 49483, 
MZB 3888); Kapuas 1976-42, 3: 51.0-69.5 mm (KUMF 2860. MZB 3889); Ka- 
puas 1976-43. 6: 36.9-75.8 mm (MZB 3890. USNM 230342); Kapuas 1976-44, 
5: 55,6-73,0 mm (MZB 389 1 . RMNH 28922); Kapuas 1 976-5 1 , 4; 47.0-7 1 .0 mm 
(MZB 3892, UMMZ 209928). 

Distribution. — Thailand (Chao Phrya), Malay Peninsula. 
Sumatra. Borneo (Kapuas, Barito). 

Trichogaster trichopterus Pallas, 1770 

Lahrus trichopterus Pallas, 1770:45 (type locality not stated), 
Trichogaster trichopterus Bloch and Schneider, 1801:165, 
Tnchopus sepat Bleeker, 1845:520 (nomcn nudum), 
Osphromenus siamensis Giinther, 1 86 1 :385 (type locality Siam), 

MATERiALExAMrNED— Western Borneo: Kapuas 1976-16, 59,1 mm (MZB 3893). 

Distribution.— Thailand (Salwin, Chao Phrya, Mekong). 
Laos. Kampuchea, Vietnam, Malay Peninsula, Sumatra, Borneo 
(western, southern, and eastern), Java, Madura, Commonly 
transported alive in the general area of its distribution, so that 
its natural distribution is impossible to determine. Introduced 
into Philippines and presumably into Bali, 



Osphronemidae 

Osphronemidae, as currently understood, comprises a single 
species, the giant gouramy. Although its suprabranchial organ 
is especially elaborate, it probably should not be placed in a 
monotypic family (see remarks under Belontiidae), 



Cuvier and Valenciennes, 1831 =Osphronemus goramv Lacep^de, 1 802, by sub- 
sequent designation of Bleeker, 1879:17). 
Osphromenus Cuvier m Cuvier and Valenciennes. 1831:377. 

Osphronemus goramy Lacepdde, 1 802 

Osphromenus olfax Commerson. ca. 1777 (unpublished manuscript name, cited 
by Lacep^de, Cuvier, and many others). 

Osphronemus goramy LacepMe, 1 802: 1 16 (type locality I'lsle de France=Mauntius, 
where it was introduced pnor to 1770). 

Osphromemis olfax Cuvier m Cuvier and Valenciennes, 1831:377 (unwarranted 
replacement name for Osphronemus goramy Lacepdde, 1802). 

Osphromenus notatus Cuvkt m Cuvier and Valenciennes, 1831:377 (type locality 
Java). See Bleeker (1879:3). 

Osphromenus satyrus Bleeker, 1845:519. 

Osphromenus gourami Regan. 1910:774 (unwarranted spelling emendation, in- 
correctly attributed to Lacfpede, 1802). 

Materjal Examined. — Western Borneo: Kapuas 1976-14, 26,4 mm (MZB 3897); 
Kapuas 1976-15, 3: 23,4-29.1 mm (CAS 49485, MZB 3898); Kapuas 1976-19, 
92.4 mm (MZB 3899); Kapuas 1976-24, 44,9 mm (MZB 3900); Kapuas 1976- 
29. 2: 34.6-39.4 mm (MZB 3901. USNM 230345); Kapuas 1976-33. 23.6 mm 
(MZB 3902); Kapuas 1976-37. 34.3 mm (MZB 3903). 

The scientific name of the giant gouramy has an interesting 
history. As pointed out by Cuvier, Commerson originally named 
the species Osphromenus olfax (from the Greek, osphromenai. 
and Latin olfax) because of his belief that the suprabranchial 
organ is an olfactory organ. Its laminated structure superficially 
resembles that of an olfactory organ, although its primary func- 
tion undoubtedly is air-breathing. For whatever reason, La- 
cep^de changed the name to Osphroncimis goranty. This spelling 
is clearly intentional, i,e,, not a typographical error or lapsus 
calami, and thus must stand according to the ICZN, notwith- 
standing Cuvier's objection to it. So far as I am aware, aerial 
olfaction has not been demonstrated in teleosts but it should be 
looked for especially in large, air-breathing species such as the 
giant gouramy that feed on terrestrial vegetation. 

Distribution,— Before it was widely transported and intro- 
duced in many new lands by man, Osphronemits goramy may 
have been restricted to Sumatra, Borneo, and Java, One of the 
most celebrated of Asian food fishes, and an important species 
in freshwater aquaculture in tropical Asia, the giant gouramy is 
found in Thailand; India (introduced by Francis Day in the 
second half of nineteenth century); Sri Lanka, Reunion, Mau- 
ritius, and Madagascar (introductions in eighteenth century); 
.Australia, etc, etc. Cuvier recommended its introduction into 
France and French colonies, 

Helostomatldae 

Helostomatidae, endemic to southeast Asia, comprises a sin- 
gle species, the kissing gouramy. The kissing gouramy is prob- 
ably the most highly specialized filter-feeding freshwater fish in 
Asia, with extraordinarily fine gill rakers and highly modified 
soft structures on its gill arches, Liem (1967/)) described the 
functional anatomy of the head, including the neurocranium, 
jaws and suspensorium. opercular apparatus, hyoid apparatus, 
and pectoral girdle, but inexplicably omitted the most highly 
specialized and interesting functional unit, the branchial arches. 



Osphronemus Lacepede, 1802 

Osphromenus Commerson, 1777? (unpublished manuscript). 

Osphronemus LacepMe, 1802:116 (type species Osphromenus olfax Cuvier m 



Helostoma Cuvier, 1829 

Helostoina Cuvier, 1829:228 (type species Helostoma temmmckii Cuvier, 1829, 
by monotypy). 



178 



CALIFORNIA ACADEMY OF SCIENCES 



Helostoma temniinckii Cuvier, 1829 

Hc/osroma Temminkn Cuvier 1829:228 (type loeality "Moluqucs." erroneous, 
incorrect spelling of species name attributable to typographic error). 

Helosioma Temminckii Cuvier in Cu\ier and Valenciennes. I83I;.W2 (type lo- 
cality indicated as .lava) 

M.^TERiAL Examined, — Western Borneo: Kapuas 1976-16. 4: 98.3-112 mm 
(MZB 3894. LISNM 230343); Kapuas 1976-33. 74.0 mm (MZB 3895): Kapuas 
1976-43. 8: 75.5-120 mm (CAS 49484, MZB 3896. USNM 230344). 

Distribution. — Central Thailand (Chao Phrva)- Malay Pen- 
insula. Sumatra. Borneo (western and southern). Java. 

Luciocephalidae 

Overall appearance pike-like; head and body elongate, sub- 
cylindrical, dorsal and anal fins placed far back; jaws extremely 
elongate and protrusible. Ascending premaxillary processes ex- 
tending posteriorly to behind eyes. Gular bone present (absent 
in all other anabantoids). Dorsal fin with 9-11 rays, anal 18- 
19, pectoral 15-16, pelvic i5, caudal 2 + 6/6 + 2. Anal fin bilo- 
bate. caudal rounded. First pelvic-fin ray prolonged (filamen- 
tous). Scales ctenoid; lateral scale series 40-42. Branchiostegal 
rays 5. Rakers on first gill arch 1 1. Vertebrae 19 + 21-22=40(3) 
or 41(4) (CAS 19998, Singapore). A single species, 

Luciocephalus Bleeker, 1851 

Diplnpicnii Gray. 1830. lig. I (type species Diplnpicnn piik hrr Grny. 1830. by 

monotypy: preoccupied by Diploptcrus Bote. 1826, Aves), 
Z.»i.(iitT/i/ia/».'; Bleeker. 1 85 lr:274 (replacement name for D//'/i'/i/i'/;n Gray, 1830). 

Luciocephalus pulcher (Gray, 1830) 

Diploptcrus pukiicr Grzy. 1830. pi 87. lig. 1 (type locality none). 
Liivioccphaliis piiliiicy. 1851<:274 (Bandjermassing. in fluviis). 

Material Examined. — Malay Peninsula: Singapore and immediate vicinity. 8: 
33.2-121 mm (CAS 19998). Western Borneo: Kapuas 1976-8. 27.6 mm (MZB 
3904): Kapuas 1976-11. 53.7 mm (MZB 3905); Kapuas 1976-13, 3: 17.7-65.8 
mm (CAS 49486. MZB 3906); Kapuas 1976-17, 2: 76.3-1 15 mm (MZB 3907, 
IISNM 230346). 

Distribution, — Malay Peninsula (Singapore, .lohore). Su- 
matra (Lahat, Palembang, Musi ilir, Benakat, Sungi Surikaka, 
Sungi Mahe. Indragiri, Laut Tador. Taluk. Gunung Sahilan. 
Djambi), Borneo (Baram, Bankayan, Kapuas, Kahajan, Ban- 
jarmasin). Banka. Billiton. 

Mastacenibelidae 

Mastacembelidae (spiny eels) and the related Synbranchidae 
(swamp eels) are perhaps the most highly modified percomorph 
families which are entirely or almost entirely restricted to fresh 
water. Their relationships to other percomorphs are unknown. 
Mastacembelids occur only in tropical Africa, where there are 
two genera and perhaps 50 species (most in the Zaire basin 
including Lake Tanganyika), and in Asia. In Asia there are four 
genera and some 24 species. 

Macrognatlius and Mastaccnibclm. the genera with most of 
the Asian species, were revised by Sufi (1956). This otherwise 
very good revision is marred by lumping of data and failure to 
indicate specimens or localities from which data were obtained 
for certain species. Several valid species are lumped under 
Macrogiiathus aculcaliis (Bloch, 1786) and Mastaccmhclus ar- 
iiialus (Lacepede, 1 800) by Sufi. MMidgnathus already has been 



revised by me (Roberts 1980); the results of this study are in- 
corporated herein. In the present account the apparently valid 
species lumped under Af. armatus are identified: .1/. notoph- 
tluiliniis new species, from the Malay Peninsula, Sumatra, 
and Borneo; M. favus Hora, 1923, from the Malay Peninsula 
and Thailand; M. undulatus McClelland, 1844, from south- 
eastern China and northern Vietnam; and the true M. armatus. 
from the Indian subcontinent including Sri Lanka and Burma. 
Original observations on systematics and distribution are also 
presented for Maslaccmhchis inacidatus and other species found 
in western Borneo. Meristic data on Mastacembelits are pre- 
sented in Table 1 6. Two genera and five species occur in western 
Borneo. 

Key to Mastacembelidae of Borneo 

I a Rim of anterior nostril with six fingerlike fimbriae; dorsal- 
fin spines 14-31; dorsal- and anal-fin rays 48-70; total 
vertebrae 70-82; maximum adult size about 300 mm 2 

lb Rim of anterior nostril with two fimbriae and two broad- 
based flaps; dorsal-fin spines 33—40; dorsal- and anal-fin 
rays 68-85; total vertebrae 84-95; all species probably 
attain 400 mm or more 4 

2a Fleshy rostrum greatly enlarged, its concave ventral sur- 
face with a large series of paired bony toothplates bearing 
numerous fine teeth; dorsal-fin spines 14-22; body with 
14-17 oblique dark bars Macrogiiathus aculeatus 

2b Fleshy rostrum not greatly enlarged, round in cross sec- 
tion, without toothplates; dorsal-fin spines 27-31; body 
without oblique dark bars 3 

3a Dorsal-fin spines 29-31; scales relatively large, readily 
visible to naked eye when skin is dry, about 14-17 above 
lateral line and less than 200 in lateral series; sexually 
mature females with an elongate genital papilla 

Mastaccmbelus maculatus 

3b Dorsal-fin spines 26-28; scales relatively small, about 22- 
25 above lateral line and 300 in lateral series; females 
without elongate genital papilla (northeastern Borneo) 

Mastaccmbelus kcithi 

4a Dorsal-fin spines 33-35, vertebrae 84-92; no vertical black 
bar below eye 5 

4b Dorsal-fin spines 37-39. vertebrae 90-96; a vertical black 
bar usually present below eye 

Mastaccmbelus notophthalmus 

5a Caudal fin with 14-15 rays, its upper and lower rays 
broadly attached to dorsal and anal fins; adults with blood- 
red horizontal bars of variable length and distribution 

Mastaccmbelus crythrotacnia 

5b Caudal fin with 19-21 rays, more or less separated from 
dorsal and anal fins; no blood-red bars 

Mastaccmbelus umcolor 



Macrognathus Lacepede, 1 800 

Macri'xnullnis Lacepede. 1800:9 (type species Opiiiciiuni aculcaluni Bloch. 1786. 

by subsequent designation ol' Jordan, 1917:56). 
Rvihliohdclla Bloch and Schneider, I80l:xliv (spelling mistake for Rhynchoh- 

dclUi). 
RhviulinhdcHa Bloch and Schneider, 1801:478 (type species Ophidium aculeatiiin 

Bloch. I 786. by subsequent designation of Jordan. 1917:59). 

For diagnosis, systematics, and discussion of this genus see 



ROBERTS-WESTERN BORNEO FISHES 



179 



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180 



CALIFORNIA ACADEMY OF SCIENCES 




FiGLiRE 132 Maslacemhclidac. a, .Mw'UH'cnihelii; unuflnr Kapuas l^'TfiO*^, I 12 mm (MZB 3^47), b. Maslacfnthclns unicolor. Kapuas 1*^76-27, 172 mm (M/B 
i'^Ab): c. Mci\UicciiibLiii<. maculalm. Kapuas l'^76-24. 137 mm gravid Icmalc {MAB 3435); d. Miicnif;milluis amlccHin. Kapuas 1976-39. 122 mm (CAS 42591). 



key above and Roberts (1980). There are at least three species, 
characterized by ditTerences in coloration and meristics: A/, aral 
(Bloch and Schneider, 1801) from the Indian subcontinent, in- 
cluding Sn Lanka and Burma; M. siamensis (Gunther, 1861) 
from Thailand; and M. aculealus from the Malay Peninsula. 
Sumatra. Borneo, and perhaps Java. Specimens trom Java have 
distinctive coloration and may represent a fourth species. 

Macrognathus aculeatus (Bloch, 1786) 

(Figure 132) 

Ophitimni ikult'iiliini Bloch, 1786:478 (type locality "Siisscn Wasser von Oslin- 

dien"), 
Macrognathus aculealus Laccpedc. 1800:9, 
For additional synonymy sec Roberts ( 1 98U). 

Material Examined, — Western Borneo: Kapuas 1976-8, 5: 101-226 mm (CAS 
49490, MZB 3920); Kapuas 1976-16, 191 mm (MZB 3921); Kapuas 1976-17. 
123 mm (MZB 3922); Kapuas 1976-27. 147 mm (MZB 3923); Kapuas 1976-33, 
3; 133-165 mm (MZB 3924. USNM 230350); Kapuas 1976-39.2: 116-122 mm 
(CAS 42591. MZB 3925); Kapuas 1976-42,2: 107-108 mm (FMNH 94252, MZB 
3926); Kapuas 1976-47, 2 122-175 mm (MNHN 1982-721, MZB 3927); Kapuas 
1976-51, 9: 102-127 mm (CAS 49491, MZB 3928. USNM 230351), 

This species dilTers from all other Macrognathus in having 
14-17 oblique dark bars on body (absent in specimens from 
Java); rostral toothplates 37-55, 2 1 in a single otherwise normal 
specimen (vs. 10-25 in other Macwgnalhus); and 22-29 pec- 
toral-fin rays (18-24 in other species). 

Distribution. — Peninsular Thailand (Trang), Malay Penin- 



sula (Tembling R.). Sumatra (Deli, Batang Hari, Lahat, Gunung 
Sahilan. Palembang). Borneo (Kapuas, Sambas. Kayan (?) R.). 
Java (northern Java east to Brantas R.). 



Mastacembelus Scopoli. 1777 

Masiaccinbclus Gronovius, 1 763: 1 33 (non-bmomial. not available lor zoological 

nomenclature), 
Mas[<)ccinbclusScopo]\, 1 777:453 (misspellingolA/£j,NMt'(V?i/((7».v Gronovius; type 

species Opimlnim inaslaccnihclus Solandcr in Russell, 1794. by absolute tau- 

tonymy). 
Fararfn'nch<>hih'lla Bleeker. 1874i;:368 (type species '^Rhyncltohi/cllu maculata 

Rv/dl "^Masuiccnihclus inaculalus Cuvier //; Cuvier and Valenciennes. 1831, 

by monotypy) 
Bdctlorhynchus Jordan and Tanaka, 1927.391 (type species Mastacembelus mac- 

ulatiis Cuvier in Cuvier and Valenciennes, 1831. b\ onginal designation and 

monotypy) 

As previously pointed out (Roberts 1980), the Asian species 
currently assigned to the genus Mastacembelus can be divided 
into two groups on the basis of morphology of the rim of the 
tubular anterior nostril. Mastacembelus sensu stricto has rim 
with two fimbriae and two flaps; the other species have rim with 
six hmbriae. Si,\ fimbriae also occur on rim of anterior tubular 
nostril in Macrognathus. African mastacembelids apparently all 
have rim with two fimbriae and two flaps. Species with six 
fimbriae have fewer fin spines and rays and vertebrae and small- 
er adults than most species with two fimbriae and two flaps. 



ROBERTS- WESTERN BORNEO FISHES 



181 



The classification of Asian mastacembelids is being studied by 
R. Travers (1984 and continuing). 

Mastacembelus erythrotaenia Bleeker. 1850 

Mastacacemblus eatenatus Meckel. 1846'^:248 (misspnnl for Maslacembclus ca- 
lenatus; type locality Borneo [^Banto?]). I have been unable to consult the full 
text or to confirm the date of this publication. 

Maslaceinhelus en-ihrolaema Bleeker. 1850:6 (type locality Banjermassing). 

Maslacembelus argiis Giinther. 1861:542 (type locality Siam). See Sufi (1956). 

Material Examined. — Sarawak: Kuching. 2: 270-317 mm (CAS-SU 33717). 
Western Borneo: Kapuas 1976-16. 2: 235-252 mm (CAS 49492. MZB 3929); 
Kapuas 1976-33. 299 mm (MZB 3930), 

Mastacembelus eatenatus was indicated by Bleeker (1859a: 
107) as a possible synonym of A/, erythrotaenia. So far as I am 
aware this is the only appearance of the name M. eatenatus in 
print other than in the original description. If the holotype, 
NMW 73350, proves to be the same species as M. erythrotaenia 
the name M. eatenatus should be treated as a nomen oblitum. 
The holotype is part of the Frank material from Borneo acquired 
by Meckel in 1 839 (NMW ace. no. 1 839. 1. 46) which may have 
originally been part of the Barito collection made by Salomon 
Mullerin 1836. 

Distribution. — Kampuchea (Mekong). Thailand (Mekong, 
Chao Phrya, Patalung R., Krabi). Malay Peninsula (Penang, 
Perak). Sumatra (Deli, Palembang, Muara Kompeh, Batang Hari, 
Sungei Mahi, Laut Tador, Indragiri, lower Langkat). Borneo 
(Baram. Kuching, Sambas. Kapuas, Bandjermasin, Balungan). 



Mastacembelus maculatus Cuvier, 1831 

(Figure 132) 

Maslacembelus maculalus Cuvier m Cuvier and Valenciennes. 1831:461 (type 
locality "Moluques." erroneous). 

Mastaeetnbelus maculalus var. chrysogasler Bleeker. 1 852(i:93 (type locality Java 
and Sumatra). 

Maslacembelus maculalus var. diclyogasler Bleeker. 1852a:93 (type locality Bil- 
liton). 

"^Maslacembelus gueni hen Day, 1865/):37 (type locality Malabar). 

Pararhynchobdella maculala Bleeker. I874u:368. 

Maslacembelus vaillanli Fowler, 1905:491. fig, 8 (type locality Baram R., Sara- 
wak). 

Maslacembelus pcrakensis Myers III Herreand Myers. 1937:74. pi, 7 (type locality 
Bukit Merah Reservoir. Perak, Malay Peninsula). 

Ataslacembelus billilonensis de Beaufort. 1939: 194 (type locality Billiton and Wai 
Lima, L^mpong Dist,. Sumatra). 

Material Examined —"Moluques": 2: 182-186 mm (MNHN 5378. syntypes 
.\t. maculalus). Malay Peninsula: Perak, Bukit Merah. 137 mm (CAS-SU 30972. 
holotype .M. perakerisis): Perak, Tapah Fishenes Station. 34 (CAS-SU 39412). 
Sarawak: Baram R.. 143 mm (ANSP 1 14888. holotype ,U, millami). Western 
Borneo: Kapuas 1976-6.2: 33.5-115 mm (MZB 3931. 3951): Kapuas 1976-7, 2: 
32.7-36.9 mm (CAS 49493. MZB 3932): Kapuas 1976-8. 68.8 mm (MZB 3933); 
Kapuas 1 976- 1 7. 4: 85,3-1 78 mm (BMNH 1982,3,29,245-7. MZB 3934); Kapuas 
1976-24, 137 mm (MZB 3935); Kapuas 1976-25. 2: 77.7-233 mm (CAS 49494. 
MZB 3936); Kapuas 1976-28. 2: 141-178 mm (FMNH 94253. MZB 3937); Ka- 
puas 1976-30. 5; 74.9-227 mm (MNHN 1982-722. MZB 3938. RMNH 28923); 
Kapuas 1 976-42. 8 1 .8 mm (MZB 3939); Kapuas 1 976-46. 5: 68,2-1 50 mm (MZB 
3940, USNM 230352); Kapuas 1976-42. 13:92.3-172 mm (IRSNB 19750. CAS 
49495, MZB 3941. UMMZ 209924): Kapuas 1976-51. 2: 92,6-176 mm (MZB 
3942, ZMA 116.548). 



This species is distinguished from all other mastacembelids 
except the south Indian Mastacembelus guentheri Day, 1865 in 
having sexually mature females with an elongate tubular genital 
papilla (possibly an ovipositor). Juveniles to 70 mm have an 



exceptionally broad pale dorsal longitudinal stripe and dorsal 
and anal fins clear except for a basal row of numerous round 
dark spots. Adults are relatively uniformly dark or somber on 
the sides of the body, abdomen uniformly pale or dusky or with 
transverse bands, fins with highly variable bands or marks. 

Mastacembelus perakensts is undoubtedly a synonym of M. 
maculatus. I have compared the holotype with other specimens 
of this species and find them identical. Contrary to the original 
description, the holotype of M. perakensis has a subocular or 
preorbital spine. I have also examined the holotype of y\/. vail- 
lanti and agree with Sufi (1956:1 13) that it is a synonym of A/. 
maculatus. Meristic data from the type specimens of M. mac- 
ulatus. M. perakensis. and M. vaillanli are presented in Table 
16. Mastacembelus guentheri from southern India (and Assam?) 
IS closely related to M. maculalus: perhaps they are conspecific. 
Sufi (1951:123) reported a 247 mm female M. guentheri with a 
large genital papilla. 

Distribution.— Thailand (without specific locality; Bleeker, 
1865:174). Malay Peninsula (Perak, Singapore). Sumatra (Deli, 
Langkat [R.?], Lematang [R.?], [ilir], Tagora R., Petok, L. Ko- 
rinchi. Lahat, Solok, Kaju Tanam, Pajakumbuk, Wai Lima, 
Sungei Penoh, Palembang, Kipahiang). Borneo (Kina Balu, Ka- 
hajan, Seminis, Kapuas). Java (Lcbak, Bogor, Tjipanas, Tjand- 
jur, Djasinga, Tji Barangbang, Palabuan, Kepong). Billiton. 

Mastacembelus notophthalmus new species 

1. Maslacembelus armalus Vob. l903/':380 (Deli. Sumatra); Volz. 1904:492 (Su- 
matra). 

.\faslacembelus armalus Weber. 1912:20 (Taluk, Sumatra); Herre. 1940:54 (Ma- 
wai Distnct); Sufi. 1956:134 et scq, (in part; Malay Peninsula and Sumatra, fig. 
31. 32); de Beaufort and Bnggs. 1962:433 (in part). 

Holotype— CAS-SU 39403. 248 mm. Tapah Fishenes Station. Perak. west 
Malaysia. A. W. Herre. 24 October 1940. 

Paratypes.- CAS-SU 39404. 11: 54.4-251 mm. same data as holotype; CAS- 
SU 39402. 119 mm. Batang Padang near Tapah Fishenes Station. Perak. west 
Malaysia. A, W. Herre. 23 October 1940; CAS-SU 35660. 3: 55.1-78.3 mm. 
Mawai Distnct. Johore. west Malaysia. A. W, Herre. February 1937; CAS-SU 
39406. 257 mm. Jalong. Perak. west Malaysia, A. W. Herte. October 1940; MZB 
3948. 59,1 mm. Kapuas basin. Sungai Engkonis. 13 km WNW of Sanggau (Kapuas 
1976-13); CAS 49497. MZB 3949. 2: 43,6-389 mm. Kapuas basm. Sungai Bc- 
hmbing. 46 km SSE of Sinlang (Kapuas 1 976-2 1 ); MZB 3950, 265 mm. Kapuas 
basin, small forested stream flowing into Sungai Pinoh near Ribang-Rabing (Ka- 
puas 1976-28), 

DiAGT^osis. — Mastacembelus notophthalmus differs from all 
other species closely related to or previously referred to M. 
armalus in having a dark vertical bar below eye (present and 
well developed in all but one or two of the smaller specimens 
in the type series) and in having the reticulate pattern absent or 
very much broken up and more evident on the ventral half of 
the body than on the dorsal half The dorsal half of the body is 
dark, with some irregular small light marks; the ventral half of 
the body is light with irregular small dark marks which some- 
times constitute an irregular and much broken reticulate pattern. 
The boundary between the dark upper and lighter lower colored 
areas is straight and sharp. Such sharply marked "color pattern 
reversal" between upper and lower halves of the body occurs 
in some samples of M. armalus from southern India but is 
otherwise very unusual among Asian mastacembelids. The re- 
versal does not include coloration of the fins. The dorsal and 
anal fins have a more or less bold pattern of sharply contrasted 
light and dark marks; in young the dark marks are restricted to 



182 



CALIFORNIA ACADEMY OF SCIENCES 



the base of dorsal and anal fins, but larger ones may have a 
continuous submarginal longitudinal black stnpe connected by 
regularly spaced vertical bars to dark marks along the fin bases. 

Distribution— Malay Peninsula (Perak, Johore). Sumatra. 
Western Borneo (Kapuas). 

Discussion. — In order to describe M. notophlhalnius n. sp. 
it has been necessary to look into the status of M. annatus 
Lacepede, 1 800 and its synonyms. As recognized by Sufi (1956). 
M. annatus is the most widely distributed and most variable 
species in the family Mastacembelidae, and has numerous syn- 
onyms. Most of the supposed synonyms are from the Indian 
subcontinent, but there are also some from China and Thailand. 
I tentatively consider that there are four valid species, all with 
entirely or almost entirely allopatric distributions. Mastaccm- 
beliis notoplithalnnis is the only member of the M. annatus 
species complex occurring in Indonesia. Its range extends into 
the Malay Peninsula, where it approaches and possibly over- 
laps the southern part of the range o^M./avus. M. notophthalmus 
and M. favus are sharply distinguished by meristic characters 
(Table 16) as well as coloration. 

Etymology. — The name notophthalmus (Latin, eye-mark) 
refers to the dark bar extending below the eye in almost all 
specimens of this species. 

The holotype of A/, annatus. which should be in the MNHN, 
was without type locality and is now lost (M. L. Bauchot, pers. 
comm., lOJanuary 1983). Hamilton-Buchanan (1822), the next 
author to deal with the species, reported it from the Ganges. 
Then. Cuvier in Cuvier and Valenciennes (1831), reported it 
from the Ganges and Bengal. Hamilton-Buchanan did not pre- 
serve specimens, but the Gangetic and Bengal specimens ex- 
amined by Cuvier are preserved in Pans (MNHN 5695, 5697- 
8, Table 16). At the time Cuvier had Lacepede's dried holotype 
on hand, but based his account on the well preserved alcoholic 
specimens with good locality data. This species has numerous 
supposed synonyms from the Indian subcontinent (Sufi 1956: 
134-135), some of which may represent valid species. 

Mastaccmbclus uudulalus (McClelland, 1844; type locality 
Chusan) seems closely related to M. armatus but is restricted 
to southern China (including Hainan) and the Red River. It is 
characterized by a relatively short and narrow head, small eye, 
scale rows above lateral line about 40 (vs. about 25 in other 
members of A/, armatus complex), 5-6 predorsal vertebrae, and 
only 87-89 total vertebrae (Table 16). 

Mastaccmbclus favus Hora, 1923, from Thailand and the Ma- 
lay Peninsula, characterized by a relatively large fleshy head and 
deep body, and very bold color pattern of large reticulations 
equally well developed over entire body; usually 4 predorsal 
vertebrae and 87-90 total vertebrae. 

Like A/, armatus. M. undulatus has a variably developed 
reticulate color pattern which tends to be more evident or more 
regularly developed on the dorsal half of the body. 

Mastacembelus unicolor Cuvier, 1831 

{Figure 132) 

Mustacembetus unuolor Cuvier m Cuvier and Valenciennes, 1 831:453 (type lo- 
cality Java). 

Material Examined, -Sunialra: Balu Sanghar, 2: 153-172 mm (CAS-SU 8667). 
Western Borneo: Kapuas 1976-21,41.8 mm (MZB 3<»43); Kapuas 1976-24,98.5 
mm (MZB 3944); Kapuas 1976-25, 3: 82.2-104 mm (CAS 49496. MZB 39451: 
Kapuas 1976-27. 172 mm (MZB 3946); Kapuas 1976-29, 9: 64.2-135 mm (BMNH 



1982.3.29.248-249. MNHN 1982-723. MZB 3947, USNM 230353). Java: 137 
mm (MNHN 5693, holotype). 

The elements of color pattern are completely obscured in the 
holotype of M. unicolor. which is uniformly darkly discolored; 
hence my identification of it with the present Kapuas material 
is based on meristics (Table 16) and morphology. The species 
has a distinctive color pattern, including two (sometimes three) 
variably developed rows of pale elongate spots on side of body, 
a large black spot on pectoral-fin base and usually a dark sub- 
marginal band on the pectoral fin, and a pale marginal band on 
the otherwise uniformly dark median fins. In juveniles dark 
coloration of the median fins is incomplete and restricted to 
large marks along the fin bases. Caudal fin narrowly joined to 
base of dorsal and anal fins; caudal-fin rays 19-21. 

Comparison with Mastacembelv.s d,4)7. — Sufi (1956:128- 
1 30) placed Mastaccmbclus dayi Boulenger, 1912 (type locality 
Irrawaddy) as a junior synonym of M. unicolor. Having ex- 
amined three specimens of A/, dari from Rangoon (NRM NNN/ 
19369895379, 159-204 mm, and 19389995376, 306 mm) I 
consider it a \alid species notably diflx^ring in caudal-fin ray 
counts (Table 16) and coloration from M. unicolor. The caudal 
fin in these three specimens is entirely or almost entirely separate 
from dorsal and anal fins (caudal fin joined to dorsal and anal 
fins near base in .M. untcolor) and has more rays than any other 
Mastaccmbclus known to me, 13-15+12=25(1), 26(1), 27(1). 
They also have slightly fewer vertebrae than A/, unicolor ex- 
amined (Table 16), but this distinction needs confirmation by 
examination of more specimens. The difference in coloration 
involves distribution and orientation of body spots and color 
pattern of median fins. M. dayi has 3 or 4 (rather than only 2 
or 3) main rows of large pale spots on side of body; the spots 
exhibit a strong tendency to vertical (rather than horizontal) 
elongation and fusion; dorsal body surface (almost or quite im- 
maculate in .\f. unicolor) covered with pale round spots. Median 
fins entirely or almost entirely covered with pale spots similar 
to those on body, particularly large on dorsal fin; on dorsal and 
anal fins spots tend to oblique elongation and fusion; spots on 
caudal fin round, well defined and rather numerous. Although 
smaller juveniles of A/, dayi have not been observed it is clear 
from the pattern present in adults that the fins do not undergo 
color change similar to that occurring in A/, unicolor. 

Distribution. — Malay Peninsula (Temheling R., Telom R., 
Pabang). Sumatra (Deli; Batu Sanghar; Laut Tador, upper Lang- 
kat, Siboga, Pajakombo, Taluk, Solok, Lake of Singkarah, Pa- 
dang, Batang, Bingin Telok, Rawa R.). Borneo (Baram, Kapuas, 
Mahakam). Java (.lakarta. Bogor, Lebak, Tjandjur, Garut, Pur- 
woredjo Modjokerta, Surakarta, Surabaya, Brantas R.). Banka. 



Synbranchidae 

Synbranchidae are elongate and eel-like; without paired fins; 
dorsal and anal fins absent or reduced to median folds, caudal 
fin reduced or absent; gill membranes united, continuous around 
isthmus, with a single ventral gill opening. Pantropical, usually 
restricted to fresh water, about four genera and 1 5 species. 

Osteology, relationships and systematics of Synbranchidae 
are treated by Rosen and Greenwood ( 1 976), but some system- 
atic problems remain. The Old World species are very poorly 



ROBERTS-WESTERN BORNEO FISHES 



183 



known. Synbranchidae are thought to be closely related to Mas- 
tacembelidae (Travers 1984). 

Monopterus Lacepede, 1 800 

Monoplcnis Lacepede. 1 SOOT 38, 139 (i>pc species Monopterus javanemis La- 

ccpdde, 1 800, by monotypy). 
Fluia Bloch and Schneider, 180I:56.S (unnecessary replacement name for .A/o- 

nopto'HS LacepMe, 1800), 
For additional synonymy see Rosen and Greenwood (1976:57). 

Monopterus albus Zuiew, 1793 

Muraena alba Zuiew, 1793:299 (type locality unknown; "presumed to be Asiatic 

Russia" according to Rosen and Greenwood 1976:57). 
Monopterus javanensis LacepMe, 1800:176 (type locality "detroit de la Sonde, 

auprcs des cotes de I'isle de Java"). 
For additional synonymy see Okada (1960:718): Rosen and Greenwood (1976: 

571). 

Material Examined. — Western Borneo: Kapuas 1976-16, 4: 36.0-37.6 mm 
( AMNH 4 1 578, MZB 3760); Kapuas 1 976- 1 7. 6: 40.7-485? mm (AMNH 41579, 
MZB 3761). 

Synbranchidae obtained by the Kapuas survey of 1976 were 
kindly examined for me by D. E. Rosen and M. N. Feinberg 
and identified by them as M. albus. 

Distribution. — A/o«op/fn« albus reportedly is widely dis- 
tributed in southeast Asia (including Burma, Thailand, Sumatra, 
Borneo, Java, Celebes). Northern and southern China. Japan. 
Okinawa. Rosen and Greenwood (1976:58) suggest it may also 
be native to Queensland, Australia. 

Soleidae 

This flatfish family seems to be represented by only a single 
genus. Achiroides, in the fresh waters of western Borneo. 

A doubtful exception is Liachinis nwlanospilus (Bleeker, 1 854), 
a widely distributed southeast Asian marine species reported 
from Sintang by Weber and de Beaufort (1929:429). I have 
examined the specimens upon which this record is based (ZMA 
1 14.433, 2: 63.3-69.9 mm, supposedly collected at Sintang by 
Lorentz in 1909). These were originally identified as L. mela- 
nospilus by Chabanaud in 1928 and their identity is not in 
question; the Sintang locality is probably erroneous. 

Achiroides Bleeker, 1851 



Achiroides Bleeker, 1851t:262 (name only: type species Flagusia iiuianorhynehus 
Bleeker, 1851, by monotypy); 1851^:402, 408 (generic diagnosis). 

Right-handed soleids with scales feebly ctenoid on both sides; 
one straight lateral line on both sides; fleshy tentacles around 
corner of mouth and on chin on both sides (more numerous on 
blind side); snout not forming prominent hook; dorsal and anal 
fins broadly joined to caudal fin; dorsal-fin rays 49-64, anal 35- 
47; pectoral fin absent; pelvic fins short, rather broad-based, 
with 3-5 rays; caudal fin large, broadly rounded, with 16 rays, 
all branched; vertebrae 28-32, Largely or entirely confined to 
fresh water (also in brackish water according to Weber and de 
Beaufort 1929:181). 

Achiroides leucorhynchus Bleeker, 1851 

Achiroides leucorhynchus Bleeker, 1851</:41 1 (type locality Surakarta, Javae cen- 
tralis, in fluviis). 



A small species of Achiroides. largest known specimen posi- 
tively identified about 50 mm; no dark pigmentation on mouth 
and surrounding area on blind side; dorsal-fin rays 49-53; anal- 
fin rays 35-37; vertebrae 28-29 (counts based on Kapuas spec- 
imens). 

Distribution. — Borneo (Kapuas.) Java (Solo basin). 

Identifications of .-). leucorhynchus from peninsular Thailand, 
based on a 77 mm specimen with 42 anal-fin rays (Smith 1945: 
440) and from the coast of Sumatra 1 00 miles west of Singapore, 
based on a 68 mm eyeless specimen (Herre and Myers 1937: 
51) require confirmation. I have not located the specimens in 
question. 

Achiroides melanorhynchus (Bleeker, 1851) 

Plagusia nielcinorhynchus Bleeker. 185 la: 15 (type locality Banjermassing. in flu- 

vus). 
.Achiroides melanorhynchus Bleeker, 1851c:262 (name only). 
.-ichiroides melanorhynchos Bleeker, 1851(^:402, 
Eurypleura melanorhyncha Kaup, 1858:100. 
Synapiitra melanorhyncha Gunther, 1862:487. 
Synaplura achira Duncker. 1904:168 (part). 

Material Examined. — Western Borneo: Kapuas 1976-14. 21: 16.7-66.2 mm 
(AMNH 48942. CAS 49499, MZB 3956, USNM 230355); Kapuas 1976-33, 10: 
21.2-112 mm (BMNH 1982.3.29.252-253, CAS 49500. FMNH 94254. MZB 
3957); Kapuas 1976-37, 2: 55.3-66.1 mm (IRSNB 19751. MZB 3958); Kapuas 
1 976-40. 40.2 mm (MZB 3959); Kapuas 1976-44. 90.4 mm (MZB 3960); Kapuas 
1976-47. 2: 34.4-39.4 mm (MNHN 1982-724, MZB 3961); Kapuas 1976-48.95.9 
mm (MZB 3962); Kapuas 1976-50.4: 25.6-44.4 mm (MZB 3963. USNM 230356); 
Kapuas 1976-54. 14: 32.4-49.1 mm (CAS 49501, MZB 3964, UMMZ 209929, 
ZMA 116.549); Sintang, 10: 31.7-61.9 mm (RMNH 7908); Boenoet. 2: 46.6-65.8 
mm (ZMA 1 14.358): Sintang. 5: 30.1-57.3 mm (ZMA 1 14.359); Poeloes Sibau, 
2: 49.3-53.1 mm (ZMA 1 14.357). 

A relatively large species of .Achiroides. attaining well over 
100 mm; dark pigmentation on mouth and surrounding area 
on blind side usually (always?) present; dorsal-fin rays 54-61; 
anal-fin rays 39^7; vertebrae 31-32 (counts based on Kapuas 
specimens). 

Distribution. — Malay Peninsula (Singapore, Pahang). Su- 
matra (Palembang). Borneo (Kapuas, Barito, Mahakam). 

Achiroides sp. undet. 

Material Examined. — Western Borneo: Kapuas 1976-16.61.8 mm (MZB 3965). 

No dark pigmentation on mouth and surrounding area on 
blind side; dorsal-fin rays 64; anal-fin rays 47; vertebrae 31. 

Possibly this is simply a specimen of .4. melanorhynchus lack- 
ing the characteristic oral pigmentation and with an unusually 
high anal-fin ray count. 



Cynoglossidae 

The predominantly marine family Cynoglossidae (tongue soles) 
comprises three or four genera and about 140 species. A number 
of species of the genus Cynoglossus apparently occur exclusively 
in fresh water, and these have not been well collected. They 
occur mainly in west Africa and southeast Asia. In the intro- 
duction to his revision of Cynoglossus. Menon (1977) stated the 
species are marine or estuarine shallow-water burrowing forms 
with a few species— C. microlepis (Bleeker). C. heterolepis We- 
ber, and C. kapuasensis Fowler— entering freshwater streams. 
This statement is misleading. Cynoglossus heterolepis. C. ka- 



184 



CALIFORNIA ACADEMY OF SCIENCES 




Figure 133. Cyiiogloistis. Upper, C. kupuasenns. Kapuas 1976-54, 199 mm (MZB 3968; lower. C n-andcrsi. Kapuas 1976-48, 107 mm (MZB 3971). 



piiascnsis. and C. waandcrsi. and possibly C. fcid ma nni and C. 
microlepis are known only from fresh water, and al least the 
first three species probably do not occur in marine habitats. C. 
hctcrolcpis is known only from large rivers in southern New 
Guinea (flowing into the shallow Gulf of Papua) which formed 
a single freshwater drainage during Pleistocene sea-level minima 
(Roberts 1978). The central or north Sunda drainage appears 
to have been the center of distribution for as many as four 
freshwater species of Cynoglossiis. According to Mcnon (1977: 
87) these four species are closely related to each other and to 
C. luieralcpis; together with the marine species C. scmilaevis, 
C. ahlvvriatiis. and C. gracilis they form his "hcicrolcpis com- 
plex." This complex is characterized by a much elongated, 
pointed snout, with angle of mouth nearer branchial opening 
than snout tip; both nostrils present; interlinear scales 15-25; 
eyes very small, interorbital space about equal to or somewhat 
greater than eye diameter; three lateral lines on eyed side, none 
on blind side. 

Cynoglossus Hamilton-Buchanan. 1822 

Cynoglossiis Hamillon-Buchanan, 1822:32, 365 (type species Cvniiglossiis lingua 

Hamilton-Buchanan, 1822, by monotypy). 
For generic synonymy see Menon (1977:16). 



Cynoglossus murolcpis Vaillant, 1893:101 (nee Bleeker, 1851; "Knapei, Se- 

bruang," Kapuas). 
Cynoglossus kapuascnsis Fowler, 1905:519 (lype locality Rapuas). 
''Cynoglossus kapwasensis Scale, 1910:288 (mispnnt for kapuasensis, Sandakan; 

manne?; probable misidcntificaiion). 



90° 



100" 



110° 



120° 



130° 




-ao" 



-10" 



-0° 



-10° 



Cynoglossus kapuasensis Fowler, 1905 

(Figure 1 33 upper) 



Figure 135. Clumfrliinos Cieographical distribution. 



ROBERTS-WESTERN BORNEO FISHES 



185 




Figure \i4. Choncrhinos amabitis. a, Kapuas 1976-36, 45.2 mm (MZB 3972. hololypc); b, Kapuas 1976-9, 48.7 mm (MZB 3973, paratype). 



Material Examined. — Western Borneo: Kapuas 1976-14. 52.9 mm {MZB 3966); 
Kapuas 1976-19,4: 169-240 mm (ANSP 1 52022, CAS49502, MZB 3967, USNM 
230357): Kapuas 1976-54, 199 mm (MZB 3968); Kapuas, Knapei. Sebruang. 142 
mm (MNHN 1891-489, formerly identified as C imcrolepis). 

Distribution. — Borneo (Kapuas). 

Radiographs of two of my specimens of C kapuascnsis reveal 
dorsal-fin rays 108-1 10, anal-fin rays 85-86. caudal-fin rays 10, 
and vertebrae 9 + 42=51. 

Cynoglossus waandersi (Bleeker, 1854) 

(Figure 133 lower) 

Plagusia waandersi Bleeker. 1854<j:98 (type locality Palembang, ubi conlluunt 

flumina Lamatang et Enim). 
Arclia waandersi Bleeker. 1859(j:l85. 
Cynoglossus waandersi Bleeker. 1866-72:31. 

Material Examined. -Kapuas 1976-32, 188 mm (MZB 3969); Kapuas 1976- 
34, 3: 102-168 mm (CAS 49503. MZB 3970. USNM 230358); Kapuas 1976-48, 
107 mm (MZB 3971). 

A radiograph of one of my Kapuas specimens of C waandersi 
reveals dorsal-fin rays 95, anal-fin rays 70, caudal-fin rays 10, 
and vertebrae 9 + 34=43. 

Distribution. — Sumatra (Batang Hari, Palembang, Taluk, 
Gunung Sahilan, Moesi R.). Borneo (Kapuas). 



Tetraodontidae 

The freshwater pufferfishes of western Borneo belong to two 
very different genera, Chonerhinos and Tetraodon. The five 
species of Chonerhinos. exclusively freshwater and restricted to 
southeast Asia, have been revised by me (Roberts 1982e): all 
five occur in Borneo, three in western Borneo. The Asian fresh- 
water species of the widely distributed Tetraodon. which in- 
cludes numerous marine species as well, have been revised by 
Dekkers (1975). Three species occur in fresh water in western 
Borneo (Fig. 135). These revisions should be consulted for ad- 
ditional information. 

Key to Freshwater Tetraodontidae of Western Borneo 

la Nasal organ confined to depression on snout; three lateral 
line canals on side of body; dorsal-fin rays 23-28; anal- 
fin rays 18-22; pectoral-fin rays 13-17; vertebrae 24-28; 
color in life almost uniformly bluish or greenish dorsally, 

white ventrally {Chonerhinos) 2 

lb Nasal organ an elevated tube divided into two lips or 
lobes; two lateral line canals on side of body; dorsal-fin 
rays 12-15; anal-fin rays 10-12; pectoral-fin rays 18-24; 
vertebrae 19; color in life highly disruptive, with reticu- 



186 



CALIFORNIA ACADEMY OF SCIENCES 




FiGURL 13b. i^'lhincrlunos modcslui Rapuas 1476-35, 54.6 mm (CAS 44505). 



lated, spotted or occllaied pattern predominantly brown- 
ish above. olT-white, grayish, yellowish, or brownish ven- 
trally (Tclraodon) 4 

2a Seales on side of body anterior and ventral to peetoral tin 
with spines directed posteriorly 3 

2b Scales on side of body anterior and ventral to pectoral fin 
with spines directed dorsally C. modcsnis 

3a Caudal peduncle relatively deep, with a roundish dark 
spot in its middle; dorsal and anal fins with angulated 
margins; upper lip not projecting beyond lower lip; ex- 
posed portion of eye round; length of nasal organ 10.1- 
17.9 ('. anuiNlis 

3b Caudal peduncle relatively elongate, immaculate; dorsal 
and anal fins usually with rounded margins; upper lip 
usually projecting; exposed portion of eye usually hori- 
zontally oval; length of nasal organ 17.3-27.4 

C. nejasius 

4a Head with flat, straight or slightly concave dorsal profile; 
opposed surfaces of nasal lobes never with spongy tissue; 
side of body with many polygonal dark spots leaving only 



lighter network or with wide-meshed dark network en- 
closing some dark rounded spots 5 

4b Head with convex or rounded dorsal profile; opposed 
surfaces of nasal lobes often with spongy tissue in larger 
specimens; side of body with many dark spots and round- 
ed blotches T. nigroviridis 

5a Sides and entire abdomen with very well defined dark 
reticulations surrounding pale areas; center of sides often 
with numerous ocelli in pale areas with large oval dark 
spots; body spines often in deep dermal pits hidden under 
dermal papillae T. palembangensis 

5b Dorsal surface and sides of body with a loose network or 
more open reticulated pattern; abdomen plain; ocelli ab- 
sent or few on sides; body spines mostly in superficial 
dermal pits without papillae T. leiunts 

Chonerhinos Bleaker, 1854 

C7?()H<"r/;/)iiM Bleekcr, 1854c 2 54-260 (type species Tflraodon «ar((H.v Richardson. 

1848; Bleeker, 1866c: 14), 
C/u'iurhinus Bleeker. 1865:213 (unjustified spelling change). 



ROBERTS-WESTERN BORNEO FISHES 



187 




Figure 137. Chonerliinos nefaslus. a. Kapuas 1976-40, 47.0 mm (MZB .W80, holotypel; b. Kapuas 1976-35, 40,7 mm (MZB 3983, paralype). 



Chonerhinos amabilis Roberts. 1982 

(Figure 134) 

Chonerhinus nantus Weber and de Beaufort. 1962:374 (specimens from Lahang 

Hara, Soengei Serawai, Kapuas basin). 
Chonerhinus modesnis Vaillant, 1893. 1902 (pro pane); Popta, 1906 (pro parte); 

Weber and de Beaufort, 1962 (fig, 84). 
Chonerhinos amabilis Roberts. 1982(>:5 (type locality Kapuas River near Putus- 

sibau). 

Material Examined. — Sumatra: Moesi R. at Mocra Klingi, 2; 36.2-38.3 mm 
(UMMZ 171708, paratypes). Western Borneo: Kapuas 1976-9, 48,7 mm (MZB 
3973. paratype); Kapuas 1976-24. 41.8 mm (MZB 3974, paratype); Kapuas 1976- 
33, 38.3 mm (MZB 3975, paratype); Kapuas 1976-34, 2: 35.9-36.8 mm (MZB 
3976, USNM 230359, paratypes); Kapuas 1976-35, 2: 35.6-46.0 mm (FMNH 
94255. MZB 3977, paratypes); Kapuas 1976-36, 2: 45,0-45.2 mm (CAS 49504, 
paratypes, MZB 3972, holotypc); 36.9 mm (MNHN 1891.216, paratype); Sintang. 
2: 40,9-41,2 mm (RMNH uncat,, paratypes); Raun, 4: 55,5-68.1 mm (RMNH 
7935. paratypes); Soengai Serawai at LebangHara, 3: 56.3-70.4 mm (ZMA 108.912, 
paratypes). 

This species feeds almost exclusively on large aquatic insect 
larvae (Roberts 1982e). 

Distribution. — Sumatra (Moesi). Western Borneo (Kapuas). 

Chonerhinos modestus (Bleaker, 1850) 

(Figure 136) 

Telraodon {Arolhron) modestus Sleeker, 1850:16 (type locality Banjermassin), 
Chonerhinos modestus Bleeker, 1854c:260, 



Material Examined. — Sumatra: Indragin, 3: 44.7-73,5 mm (NHMB 822- 
824); Lahat, 3: 66.6-81.1 mm (RMNH 12004). Western Borneo: Kapuas 1976- 
35. 2: 64.6-106 mm (CAS 49505, MZB 3978); Kapuas 1976-48, 2: 46.8-48.4 
mm (MZB 3979. USNM 230360); Kapuas at Sanggau, 78.7 mm (RMNH 26931, 
lectolype); Kapuas at .Sanggau, 2: 49.2-59.2 mm (RMNH uncat); Kapuas at 
Sintang. 3: 50.0-58.9 mm (RMNH 7934). Southern Borneo: Banlo Rivcral Moara 
Teweh, 2: 90.5-95.3 mm (NMW 64647). Banka Island: Muntok, 4: 52.2-97.0 
mm (NMW 64646). 

Chonerhinos modestus. attaining at least 106 mm and prob- 
ably the largest member of its genus, feeds mainly on terrestrial 
insects, shrimps, seeds, and to a lesser extent on whole fish, fin 
rays or scales (Roberts 1982e). 

Distribution. — Sumatra (Lahat, Palembang), Borneo (Ka- 
puas, Barito). Bangka Island (Muntok). This species may occur 
in Thailand and the Malay Peninsula but all specimens of C/;o- 
nerhinos from these places examined by me are C. nefaslus. The 
records from Barito R. and Banka are based on specimens ex- 
amined in Vienna after publication of my revision. 



Chonerhinos nefastus Roberts, 1982 

(Figure 137) 

Chonerhinos modestus D'Aubenton and Blanc, 1966:561 (Mekong basin, Kam- 
puchea); Taki, 1974:199-200, fig, 187 (Mekong basin, Laos); Imaki et al., 1978: 
29, pi. 18 (Kapuas R. at Sintang). 

Chonerhinos nefastus Roberts, 1 982£>: 1 (type locality Kapuas R. near Putussibau). 



188 



CALIFORNIA ACADEMY OF SCIENCES 



90° 100° 110° 120° 130° 




• leiurus 

* palembangensis 



"^^^ 



C^' 



-10° 



Figure 138. Tctraodon Geographical distribution of two species found in 
western Borneo. 



Material Examined. — Western Borneo; Kapuas 1976-33, 4: 34.3^3 3 mm 
(CASa^SOb. MZB 3481. paratypes); Kapuas 1976-34, 3: 36.6-43.7 mm (BMNH 
1982.3.29.:54-5, MZB 3982, paratypes); Kapuas 1 976-35, 6: 32.9-60.2 mm (IRSNB 
632, MZB 3983, ROM 38601, USNM 230361. paratypes); Kapuas 1976-37. 51.7 
mm (MZB 3984, paratype); Kapuas 1976-39, 64.9 mm (MZB 3985, paralype); 
Kapuas 1 976-44, 4; 36.7-57.8 mm (MZB 3980, hololype, MZB 3986. CAS 49507, 
paratypes). Southern Borneo; Banto R. at Moara Teweh, 39. 1 mm (NMW 64645). 
For matenal examined from Kampuchea, Thailand, Malay Peninsula, Sumatra, 
Sarawak, and additional localities from Borneo sec Roberts (1982i';10-l I). 

This species feeds largely on the lin rays and scales of other 
fishes (Roberts 19820. 

Distribution. — Kampuchea (Mekong basin). Thailand (Me- 
kong. Songkhia Lake). Malay Peninsula (Perak). Sumatra (Moesi 
R). Borneo (Niah, Rejang, Kapuas, Mentaya, Barito, Maha- 
kam). The record from the Barito is based on the 39.1 mm 
specimen from Moara Teweh reported above, examined in Vi- 
enna after publication of Roberts (1982c). 



Tetraodon Linnaeus, 1758 

Ictraodon Linnaeus, 1758;332 (type species Tvtraoiion hncalu^ Linnaeus, 1758, 
by subsequent designation of Lesson, 1830;198-199). 

Tetraodon leiurus Bleeker. 1851 

Tciraitdon Iciiini.s Bleeker, 1851/';97 (type locality Batavia. in man ct ostiis flu- 

\iorum). 
Ictwdon Hilficndorhi Popta, 1905;185-186 (type locahty Boh River, upper Ma- 

hakam). See Dekkers (1975). 
Iclrndoii Bcrgii Popta, 1905; 186 (type locality Bongan River at Bulit, Kapuas). 

See Dekkers (1975). 
iclraodon ram/i()i/,?;c/i.v;.v Chabanaud. 1923:137-140 (type locality Pnom-Penh). 

See Dekkers (1975). 

Material Examined. -Western Borneo; Kapuas 1976-14,42.8 mm (MZB 3987). 

Distribution (Fig. 138). — Kampuchea (Mekong). Thailand 
(Nontaburi, Bangpakong R., Thale Luang, Patalung R.). Malay 
Peninsula (Jahan, Sembeling). Sumatra (Pangian R., Deli, La- 
hat, Palembang, Padang, Solok, Sinkiarak Lake). Borneo (Ka- 
puas, Mahakam). Java. 

Although Dekkers (1 975; 1 12) indicated east Malaysia as part 



of its range, he did not list any specimens examined from there 
or include any locality records from there. 

Tetraodon nigroviridis Marion de Proce, 1822 

Tetraodon nigrovindis Marion de Proce, 1 822; 130 (type locality freshwater pool 

on NE coast of Sumatra). 
Telrodon siimduns Cantor, 1850;1356-1357 (type locality Malay Peninsula). 

Material Examined. — None. 

The inclusion of this species is based on identification of a 
72 mm specimen (ZMA 1 10,21 1) from South Natuna Island by 
Dekkers (1975:125). 

Tetraodon palembangensis Bleeker, 1852 

Tciiuodon pulfiiibangensis Bleeker, 1852(/;21, 25, 26 (type locality Palembang). 
Tetraodon piiigiiis Vaillant, 1902;38 (type locality Mahakam River at Tepu). 

Material Examined. — Western Borneo; Kapuas 1976-19, 8; 54.7-132 mm 
(CAS 49508. MZB 3988); Kapuas 1976-44, 6; 68.0-185 mm (MZB 3989, USNM 
230-362); Kapuas 1976-51. 64.5 mm (MZB 3990). 

Distribution (Fig. 138). — Laos (Mekong). Thailand (Me- 
kong, Chao Phrya, Tale Luang, Tale Noi, Tale Sap). Malay 
Peninsula (Perak. Jahan R.). Sumatra (Palembang, Gunung Sa- 
hilan, Djambi R., Indragin R.). Borneo (Kapuas, Mahakam). 
Java. Billiton. 



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ROBERTS-WESTERN BORNEO FISHES 



189 



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Genensch overzicht der Fauna. Visschen. Nat. Geneesk. Arch. Ned. Ind. 2: 

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. 1846a Overzicht der Siluroidcn welke te Batavia voorkomen. Nat. 

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1849. Bijdrage tot de kennis der ichthyologische fauna van hel eiland 



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. 1850. Bijdrage tot de kennis der ichthyologische fauna van Midden-en 

Oost-Java, met beschnjving van eenige nieuwe species. Verb. Bat. Gen. 23:1- 

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. 18516. Over dne nieuwe soorten van Tclraodon van den Indiscben 



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Nat. Tijds. Ned. Ind. 1:259-275 (35). 
. I851(/. Over eenige nieuwe soorten van Pleuronectoiden van den In- 

dischen Archipel. Nat. Tijds. Ned. Ind. 1:401^16 (38). 

1 85 If. Over eenige nieuwe soorten van Mcgalops. Dussunneria. No- 



loplerus en Aslroneslhes Nat. Tijds. Ned. Ind. 1:417^24 |39). 
. 1851/ Derde bijdrage tot de kennis der ichthyologische fauna van Bor- 



neo, met beschnjving van eenige nieuwe soorten van zoetwaterfvisschen. Nat. 
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. 185lj Nieuwe bijdrage tot de kennis der Pcrcoidei. Scleroparei, Sciae 



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. 1851/) Vierde bijdrage tot de kennis der ichthyologische fauna van 

Borneo, met beschnjving van eenige nieuwe soorten van /oetwatervisschen. 
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. 1851(. Vijfde bijdrage tot de kennis der ichthyologische fauna van Bor- 
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. 1852a. Bijdrage tot de kennis der ichthyologische fauna van Blitong 



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ROBERTS-WESTERN BORNEO HSHES 



195 



Index to Genera and Species 



abbreviatus, Cynoglossus 1 84 
abei. Ctenogobius 168 
Acanthonolus 45 
Acanthonotus argenteus 45 
Acanthoperca 161 

gullivcn 161 

woimi 164 
Acanthophthalnius 5. 95 

anguillans 95 

I'asciatus 95 

ja\anicus 97 

lorcnUi 95, 104 

pahangcnsis 106 

semicmctus 18. 96. 98 

shclfordi 13. 14, 16-18. 96, 98 

vermicularis 95 
Acanthopsis 5, 99 

choirorhynchos 14, 17, 18, 100 

choirorhynchus 8 

dialu/oma 5 
Acanlhopsoides 94 

gracilis 14, 17-20, 94. 96 
-Acantoccras 95 
.Acantophis 95 
Acamophthalmus 5, 6, 95. 97-99. 10.3. 106 

anguillans 17. 95-98 

fasciatus 95 

javancius 95. 97 

muracniformis 96 

mariae 98 

mycrsi 96 

oblongus 17. 19, 96-98 

pangia 96. 98 

pcrakcnsis 97 

semicmctus 16. 19, 20. 96. 98 

shellbrdi 16. 18. 96. 98 

supcrbus 16. 20, 96, 98, 99 

\crmicularis 97 
Acamopsis 5, 94, 95. 99, 100 

choirorhynchos 18, 99 

choirorhynchus 94, 96 

dialu/ona 99 

lachnosloma 96. 99 
acerus. Paracrossocheilus 17, 57 
achira, Synaptura 185 
Achiroides 183 

leucorhynchus 18, 19. 183 

melanorhynchos 183 

melanorhynchus 16. 18-20. 183 

sp. undet, 16, 183 
Acrochordonichthys 136-138. 141. 142 

buettikofcn 138 

chaniacleon 137 

melanogaster 137. 138 

cf. melanogaster 14. 136. 138 

obscurus 1 38 

pachyderma 137. 138 

platycephalus 1 37 

rugosa 137 

rugosus 137 

varius 138 
acrosloma. Sphaerichthys 175-177 
aculeatum, Ophidium 178, 180 
aculeatus, Macrognalhus4, 14, 16, 18-20, 178, 180 
Adamacypris 59 
Afronandus 164 

agilis, Rasbora 13, 14, 16. 17, 19, 20, 66-68 
agrcnsis, Plalacanthus 103 
akarensis. Betta 172 
Akysis 134. 136, 138-141 

armalus 117. 137. 138 



baramensis 134. 136. 138 

hendncksoni 136 

hutchinsoni 142 

leucorhynchus 140 

macronema 1 36 

cf. macronema 1 36 

major 1 34 

pseudobagarius 18, 136. 138-140 

variegalus 136, 142 
Albulichthys 27, 30, 40 

albuloides 28 

krempfi 28 
albuloides. Albulichthys 28 
albuloides. Systomus 27. 28 
albus. Monopterus 16. 183 
alcinaces, Chela 31 
Allochela 3 1 
Allodanio 30 
almorhae. Botia 101 
altus. .^mhlyrhynchichthys 41 
altus. Puntius 62 

amabilis, Chonerhinos 9, 14, 17, 18, 184, 186-187 
Ambassis apogonoidcs 161 

(Bogoda) microlepis 162 

houlengen 164 

microlepis 160, 162 

robustus 162 

wolffii 161. 162 
Amblyrhynchichthys 27-29. 40, 41 

altus 29. 41 

truncatus, 6. 16. 18-20, 28, 29 
amboinensis, Cryptopterus 146. 147 
Amphilus atesuensis 12 
.^nabas 1 7 1 

oligolepis 8 

seandens 171 

testudincus 8, 171 
anableps, Cobitis 95 
anchisporus. Barbus 60 
anchisporus. Puntius 14, 16, 18, 19, 60 
anabatoides, Betta 13, 17, 18. 172. 173 
Anematichthys 33 
anguillans, .Acanthophthalmis 95 
anguillaris, Acanthophthalmus 17, 95, 96 
anguillaris, Acamophthalmus 95-98 
anguillans, Cobitophis 97 
anguillans. Silurus 127 
angusticeps. Pseudotylosurus 1 54 
anomaloptenx. Parakysis 18. 19, 142. 143 
anomalurus. Chela 56 
Anomalura 55 

anomalura. Oxygaster 16. 18. 55. 56 
Anthias testudineus 1 7 1 
Aoria 120 
.Aonchthys 120 
Aperioptus mcgalomyeler 103 
aphelocheilus. Protomy/on 84 
Aplocheilus 77 
apogon. Barbus 33. 35 
apogon. Cryptopterus 16. 145. 150 
apogon, Cyclocheilichthys 8. 14. 16-19. 33-35 
apogon. Cyclocheilichthys (Anematichthys) 35 
apogon. (Cryptopterus 145. 146 
apogon. Silurus 145 
apogon. Systomus 35 
apogonoidcs. Ambassis 161 
apogonoides. Chanda 16. 19, 161 
apogonoidcs. Channa 1 8 
apogonoides, Cyclocheilichthys (Anematichthys) 

35 
apogonoides, Parambassis 1, 11, 159. 161. 162 



apogonoides. Systomus 35 

aral, Macrognathus 180 

Arapaima 23 

Arelia waandersi 185 

argenteus, Acanthonotus 45 

argus, Mastacembelus 181 

argus, Scatophagus 6 

argyropleuron. Anus 1 10 

argyrotaenia, Leuciscus 66, 67 

argyrotaenia, Rasbora 8, 16, 19, 20, 66-68, 72, 74 

Anus 21, 110, 111 

argyropleuron 1 10 

anus 1 10 

cf. stirlingi 1 10 

damelsi 1 10 

gagora 1 1 

(Hemianus) damelsi 110 

leptaspis 6 

melanochir 16, 18, 19, 21. 110. 11 1 

octik 110 

sp. undet. 126 

stirlingi 1 10 

stormii 16, 21, 110, 111 
anus. Anus 1 10 
anus, Pimelodus 1 10 
armatus, Akysis 117, 137, 138 
armatus. Barbus 35 

armatus. Cyclocheilichthys 13. 14. 16-19, 33-35 
armatus, Leiocassis 17, 18, 116, 117 
armatus. Mastacembelus 178. 179, 181, 182 
.Aspidobagrus 4, 120 

gulio 4 
atesuensis, Amphilius 12 
attu, Silurus 151 
axelrodi, Rasbora 2, 11, 13, 21, 44, 66-68, 70 



Baganus2, 12, 133. 134. 136. 140 

hagarius 136 

lica 22, 133. 134 

nieuwenhuisi 1 33 

yarrelli 16. 18. 133. 134, 140 
bagarius. Baganus 136 
bagarius, Pimelodus 133 
Bagnchthys 12, 111-115. 123, 124 

hypselopterus 8, 16, 19, 111-113. 123 

macracanthus 111-113 

macropterus 16. 19. 112-115 

micranodus 19. 112-115 
Bagroides 111. 115 

macropterus 111. 114 

mclanopterus 1 1 5 

melaplerus 8. 16. 19. 115 
Bagrus 

gulio 120 

heterurus 122 

hoevenu 121 

hypselopterus 111. 112 

lamarii 120 

macronema 120 

macronemus 122 

micracanthus 120 

micropogon 1 1 7 

nemurus 120. 121 

nignceps 120. 122 

poeeilopterus 1 16 

sieboldii 122 

singanngan 122 

stenomus 1 16 

sundaicus 1 10 

wolffii 126 



196 



CALIFORNIA ACADEMY OF SCIENCES 



w^ckii 126 

yarrelli 133 
balaena, Pcctenocypns 1. 16. 18. 58. 59 
Balantiochcilos 29 

Balantiocheilos melanopterus 8. 17. 19. 29 
baheata. Vaimosa 169 
bankancnsis, Chaca 8. 16. 19. 143 
bankanensis. Channa 169. 170 
bankanensis. Leuciscus 70 
bankanensis. Ophicephalus 169 
bankanensis. Rasbora II. 14, 16-20. 66, 68. 70. 

71. 74 
banksi. Puntius binolatus 61 
baramcnsis. Akysis 134. 136. 138 
baramensis, Glyptostcrnum 138 
baramcnsis. Liocassis 1 17 
baramcnsis. Myslus 122-124 
baramensis, Ophloccphalus 170 
barbatuloides, Cobitichthys 103 
barbaluloides, Cobitis 103, 104 
barbatuloides. Misgumus 8, 103 
barbatulus. Cobitis 95 
Barbichlhys 29 

lacvis8, 16-19. 30 

laevis var. sumatranus 30 

laevis nitidus 30 

nitidus 30 
Barbodcs 60 

lascialus 64 

hemictcnus 61 

hexa7ona 65 

lateristriga 65 

sealci 6 1 
Barbucca 94. 110 

dialolica 18. 19. 96. 100. 101 
Barbus 38 

anchisporus 60 

apogon 33. 35 

armalus 35 

binotalus 60 

binotatus var. palavancnsis 61 

blitoncnsis 60 

boulcngcn 62 

brachyncmus 30 

bramoides 6 1 

biilu 60 

carassioidcs 6 

collingwoodii 62 

elongalus 6 1 

enoploides 35 

cnoplos 35 

enoplus 33 

cxerelln 64 

fascialus 64 

fascialus var. chaperi 64 

Fowlerii 35 

gobioides 30 

gomosoma 61 

hampal var. bimaculala 40 

heleronema 33, 36 

hexa/ona 65 

hoevenii 41 

ivis 61 

kahajani 65 

kalopterus 38. 39 

kusancnsis 60 

laevis 29. 30 

laevis var. sumatranus 30 

latenstriga 65 

lawak40. 41 

lincatus 35. 65 

macracanthus 35 

maculalus 60 



maculatus var, hagenii 61 
marginatus 45 
melanopterus 29 
microlepis 78 
obtusirostns 45 
orcsigenes 60 
palavanensis 61 
penta/ona 65 
podonemus 43 
pohspilos 61 
procto/ysron 60 
quinquemaculatus 61 
repasson 37 
Schwanenl'eldii 66 
schwanefcldi var. rubra 66 
sctigerus 42. 43 
stngatus 62 
sumatranus 8. 60 
lamhra 80 
telra/ona 64, 65 
Iruncatus 28. 29 
\alenciennesii 35 
waandersi 60 
(Puniius) sumatranus 61 
barila, Cyprinus (Banlius) 30 
barilla, Cypnnus 30 
Barilius 21. 30. 44 
borneensis 17. 30, 31 
nanensis 31 
ponticulus 31 
salmolucius 30 
Barynotus microlepis 8, 36 
Batrochocephalus mino 1 10 
batrachus, Clanas 127 
cf batrachus, Clanas 127 
batrachus, Silurus 127 
Bdellorhynchus 180 
bcaulbrti, Rasbora 10, 66, 71 
Beaurortia91, 92 

Icveretti 85 
bcavcni. Ncmachcilus 107 
Bedula 

nebulosis 164 
hcldti, Cryplopterella 145. 148 
belinka. Systomus (Barbodes) 60 
Belodontichthys 2. 144 

dinema 16. 144 
Belodonlichthys macrochir 144 
Bclone 
cancila 152. 153 
canciloidcs 153 
caudimaculala 27 
Bclonion I 52 
Belontia 171 
hasselti 171 
hasscltii 13. 16 
signata 1 71 
Bendehsis 30 

bcndelisis. Cyprinus (Banlius) 30 
bcrdmorci. Syncrossus 101 
bergii. Tetrodon 188 
Betta 171-174 
akarensis 172 

anabatoides 13. 17. 18. 172. 173 
cf. taeniata 1 72. 173 
cocci na I 72 

dimidiata II, 16. 18-20, 172-174 
cdithae 173. 174 
I'ocrschi 172 
macrosloma 171. 172 
picta 171 

pugnax 8. 10. 13. 16, 18-20, 171-173 
splendens 1 7 1 



taeniata 13, 14, 173, 174 

trifasciata 1 7 1 

unimaculata I 72 
bicirrhis. Cryptopterus 148 
bicirrhis. Kryptoptenchthys 147 
bicin-his. Kryptopterus 16. 18. 19, 146-148, 150 
bicirrhis, Silurus 145, 147 
bicolor, Labeo 39 

bicolor, Leiocassis (Pseudomystus) 1 16 
bicomis, Paracrossocheilus 57 
hilmcata, Homaloptera 88, 89 
bilineatus. Doryichlhys 158 
billitoncnsis, Maslacembelus 181 
bimaculala, Barbus hampal var, 40 
bimaculata, Hampala 17, 40 
bimaculatus. Callichrous 4 
bimaculatus. Mystus 121 
bimaculatus. Silurus 150 
binotatus. Barbus 60 
binotatus. Coius 164 
binotatus. Puntius 13. 14. 17, 60, 61 
binotatus banksi, Puntius 61 
bistnatus. Ophiocephalus 170 
biviiiatus. Ophicephalus 170 
bleeken. Clupeichthys 14. 16-18. 24 
blitonensis. Barbus 60 
bo. Lobocheilus 41 
bo. Macrones 124 
bo. Mystus 124 
bo. Tylognathus 41 

boaja. Doryichthys 8, 16. 18. 158. 159 
boaja. Syngnathus 158. 159 
bocourti. Heterobagrus 120. 123. 124 
bocourti, Mystus 124 
hoddaerti, Boleophlhalmus 8 
Bogoda macrolcpis 162 
Bola 30 

bola, Cypnnus (Banlius) 30 
Boleopthalmus boddacrti 8 
bongan. Macrones 122 
bongan, Mystus 124 
borapctensis. Rasbora 68 
borneensis. Banlius 17. 30. 31 
borneensis, Clupeoidcs 25 
borneensis, Discognathus 40 
borneensis, Gana 17, 40 
borneensis, Gastromy/on 9, 79, 84. 85, 87, 93 
borneensis, Hemipimclodus 110, III 
borneensis, Nandus 164 
borneensis, Notopterus 17, 18, 19, 24 
borneensis, Osteochilus 16, 18, 46, 47 
borneensis, Pimelodus 110, 111 
borneensis, Polynemus 166 
borneensis. Prolomy70n 84 
borneensis. Rasbora 16. 18. 19. 20. 68. 71. 72 
borneensis. Rohita 47 
Botia 101. 102 

almorhae 101 

helodes 102 

hymenophysa 8. 18. 96. 101-103 

macracantha 8, 16-19, 9h, 102 

revcrsa 17, 96, 101-103 
boulcngcn, Ambassis 164 
boiilengeri, Barbus 62 
Brachydanio reno 70 
Brachygobius 167 

donac 1 3, 167 

nunus 167, 168 

xanthomelas 14, 16, 19, 167. 168 
brachyncmus. Barbus 30 
brachynotopterus. Osteochilus 52 
hrachynotoptcrus. Rohita 52 
brachypoplerus, Eutropius 128. 129 



ROBERTS-WESTERN BORNEO FISHES 



197 



brachypoplcrus. Pseudeutropius 12. 17-19. 129 
brachyrhynchops. Dor\ichlhys 159 
bramoides. Barbus 61 
bramoides. Punlius 14. 16. 18. 61 
bramoides. Puntlus (Barbodes) 62 
Brcitcnstcinia 136. 140-142 

cf insignis 19. 136, 140 

insignis 8. 140 
breilensleini, Parachela 8 
brevicauda. Oslcochilus 49 
breviceps. Engraulis 26 
breviceps. Leiocassis 1 16 
breviceps. Leiocassis (Pseudom\stus) 1 16 
breviceps. Selipinna 25 
brevipinnis. Glyptothorax 12 
bnttani. Rasbora 14. 18, 19. 66, 68, 72 
brocki, Sligmatogobius 13. 169 
brocki. Vaimosa 169 
Buettikoferi. Acrochordonichthys 138 
bulu. Barbus 60 

bulu. Pumiopliles 16-19. 41. 60 
bulu. Puntius 60 
bulu. Systomus 60 
bulu. Systomus (Puntius) 60 
Bunaka herwerdenii 167 
Butis 

gymnopomus 13 



cachia. Cypnnus (Chela) 55 
Cachius 31 
cachius. Chela 55 
cachius. Cypnnus 31 
Calamiana 168 

kabilia 168 

magnoris 168 

sp. undet. 10. I I. 16. 18. 19. 167. 168 
Callichrous 150 

bimaculatus 4 

eugeneiatus 1 50 

hypophlhalmus 150 

macronema 8 

micronema 1 50 

weberi 1 5 1 
Callieleotris platycephalus 166 
calvanus. Silurus 126 
cambodgiensis. Tetraodon 188 
campbclli. Datnioides 164 
cancila. Belone 151. 153 
cancila. Xenentodon 153. 155 
canciloides, Belone 153 
canciioides. Mastacembclus 153 
canciloides. Xenentodon 14. 17-20. 153. 155 
Capoeta 

deventen 35 

enoplos 35 

macrolepidota 40 

microlepis 33. 36 

padangensis 45 

siaja 33 
carassioidcs. Barbus 6 
catenatus. Mastacembclus 181 
Catopra 165 

fasciata 8, 165 

grootii 8. 165 

malabarica 165 

nandoidcs 165 
caudatus. Microphis 159 
caudimaculata. Belone 27 
caudimaculata. Rasbora 13. 68. 72, 74, 76 
cavasius, Macrones 122 
cavasius. Mystus 122-124 
caveni. Rasbora 68 



Cephalocassis 21. 110 

melanochir 1 10 

stormii 1 10 
ccphalotaenia. Leuciscus 72 
cephalotaenia. Rasbora 13. 14. 66. 68. 72 
Ceratoglanis 144. 145 

scleronema 16. 144 
Chaca 2. 10. 143 

bankanensis 8. 16. 19. 143 

chaca 143 

Gray 143 
chaca. Chaca 143 
chaca. Platyslacus 143 
Chaetodon quadril'asciatus 164. 165 
chamaleon. Acrochordonichthys 137 
chamaleon. Sosia 137 
Chanda 160, 161 

apogonoides 16. 19. 161 

lala 161 

macrolepis 16. 17. 19. 162 

naina 160, 161 

wolffii 16, 19. 20, 164 
Channa4. 20. 169, 170 

apogonoides 18 

bankanensis 169. 170 

cf. striata 4 

luciusS. 13. 14. 16. 18. 20. 170 

macrolepis 18 

marulioides 8. 17. 170 

melanoplera 1 70 

melasoma 1 70 

micropeltes 8. 13. 169. 170 

onenlahs 10. 18. 168. 170 

pleurophthalma 18 

pleurophthalmus 17 

striata 4. 8. 13. 19. 20. 170 
chaperi. Barbus fasciatus 64 
chaperi. Diastatomycter 9. 145 
chapcn. Hemisilurus 145 
chaseni. Leiocassis 1 17 
chedra. Cypnnus (Banlius) 30 
Chedrus 30 

grayii 30 
Chela 31. 55 

alcinaces 3 1 

anomalurus 56 

cachius 55 

fasciatus 3 1 

hypophthalmus 8. 55 

maassi 14. 18. 19, 31 

macrochir 8 

megalolcpis 31. 57 

oxygastcr 56 

oxygastroides 57 

(Allochela) maassi 31 

(Malayochela) maassi 31 
Chitala 24 
chitala, Mystus 24 
Chlonas 127 

choirorhynchos. Acanthopsis 14. 17. 94 
choirorhynchos. Acantopsis 18. 96. 99. 100 
choirorhynchos. Cobitis 99 
Chondrostoma lipocheilos 42 
Chonerhinos 2. 8. 11. 185. 186. 187 

amabilis9. 14. 17. 18, 184, 186, 187 

modestusS. 19. 184. 186. 187 

nefastus 18. 184. 186. 187 

remotus 184 

silus 184 
Chonerhinus 8. 185 

modestus 8. 187 

naritus 187 

nefastus 8. 184. 186-187 



Chopraia 88 

rupicola 88 
chrysogaster. Mastacembclus maculatus var, 181 
Chrysophekadion 

polyporo 45 
chrysophekadion. Labeo 45 
chrysophekadion. Morulius 16. 45 
chrysophekadion. Rohita 45 
chr\sopunctatus. Hcmirhamphodon I 58 
Cirrhinus fasciatus 64 
Clanas4. 127. 128 

batrachus 127 

cf batrachus 127 

jagur 128 

lazera 1 2 

leiacanthus 13. 18. 127. 128 

liacanthus 127 

magur 127 

meladerma 4. 127 

melanoderma 4, 20. 127 

melanosoma 127 

melasoma 127 

nicuhofi 4. 128 

pentaptcrus 128 

teijsmanni 128 

tcysmanni 127. 128 
Clupea 

hypsclosoma 25 

phasa 26 

(Clupeoides) potamophilus 25 

potamophilus 25 

telara 26 
Clupeichthys 24 

bleeken 14. 16-18. 24 

gomognathus 24 
Clupeoides 25 

bomeensis 25 

hypsclosoma 16. 24. 25 

potamophilus 25 

pseudopterus 24 
Cobitis 94. 107 

anableps 95 

barbatuloides 103. 104 

barbatulus 95 

choirorhynchus 99 

fasciatus 6. 106 

fossilis 95 

hasselti 103 

hymenophysa 101 

kuhlii 95 

macracanthus 102 

macrochir 106 

oblonga 97 

octocirrhus 95 

pangia 95 

taenia 95 
cobitis. Crossocheilos (Crossocheilichlhys) 32 
cobitis. Crossocheilus 16. 18. 32. 33 
cobitis. Crossochilus 17. 32 
cobitis. Lobochcilos 32 
Cobitichthys 

barbatuloides 103 
Cobitophis95. 97 

anguillaris 97 

perakensis 96. 97 
coccina. Betta 172 
Coilia 

coomansi 20. 25 

melanochir 26 
Coius 

binotatus 164 

nandus 164 

polota 165 



198 



CALIFORNIA ACADEM\- OF SCIENCES 



LOllingvvoodi. Puntius 17 
collingwoodii. Barbus 62 
collingwoodii, Puntius 62 
confinis, Parambassis !64 
contractus. Gastromy/on 17. 84-87 
coomansi, Coilia 20, 25 
Corica (Clupeichthys) goniognalhus 24 
Corica goniognathus 24 
Cosmanum 59 
Cosmochilus 10. .^1 

falcifcr 3, 16, 20. }\ 

harmandi }l 
crocodilus. Engraulis 26 
crocodilus, Lycolhnssa 16, 18, 19, 25, 26 
Crossocheiios 32 

(Crossochcilichthys) cobttis ^2 
Crossocheilus 32, 39. 40 

cobitis 16-18. 32. 33 

oblongus 13, 16-18, 20, 32 

oblongus var. nigriloba 32 

paracrossocheilus 

sp. undct, 19 
Crossochilus 32 

cohitis 32 

oblongus 32 

vittatus 57 
Crossostoma 83 
Cryplopterella 145 

hcldti 145. 148 
Cryptopterus 145 

amboinensis 146. 147 

apogon 145, 150 

bicirrhis 148 

lais 147 

limpok 147 

macrocephalus 148 

micronema 149, 150 

micropogon 145 

micropus 147 

schiheides 150 
cryptoptcrus, Kryptoplerus 16-18, 146, 147 
cryptopterus, Silurus 145, 147 
ctenoccphalus, Gastromy/on 83, 87 
Clcnogobius abei 168 
Ctenopoma 1 7 1 
Ctcnops nohilis 176 
cyanochloros, Pimelodus 135 
cyanonuias, Rohita 45 
cuvien, Dangila 37, 38 
Cyclochcilichthys 2. 8. 21, 22. 29. 33-39, 78. 161 

apogon 8. 14, 16-19. 33-35 

armatus 13, 14, 16-19. 33-35 

dc7vvaani 35 

dumcrili 35 

cnoplos 3,3-35 

luneronema 14. 16. 18, 19, 33, 34, 36 

janthochir 17-19. 31. 35. 36 

lincatus 35 

niacracanthus 35 

mcgalops 37 

nickongensis 35 

microlcpis 18-20. 33, 35, 36 

rcpasson 17-19. 33. 34, 37 

rubnpinnis 35 

sp 18 

tapicnsis 35 

(Anematichlhys) apogon 35 

(Ancnialichthys) apogonoidcs 35 

( Ancmatichthys) janthochir 36 

(Cyclochcilichthys) armalus 35 

(Cyclochcilichthys) cnoplos 35 

(Cyclochcilichth\s) macracanthus 35 

(Cyclochcilichthys) repasson 37 



(Siaja) devcnten 35 

(Siaja) heteroncma 36 

(Siaja) microlepis 36 

(Siaja) siaja 35 
Cynoglossus 183-185 

abhreviatus 184 

rddmanni 184 

gracilis 184 

hctcrolcpis 183, 184 

kapuascnsis 16, 20. 183-184. 185 

kapwascnsis 184 

lingua 184 

microlepis 183. 184, 185 

scmilacvis 1 86 

waandersi 18, 19. 184. 185 
Cypnnus 31, 55 

banlia 30 

(Banlius) bania 30 

(lianlius) hcndelisis 30 

(Banlius) hola 30 

(Banlius) chedra 30 

(Banlius) shacra 30 

cachius 31 

(Chela) cachia 55 

lampta 40 

laubuca 3 1 

(Morulius) morala 45 

oxygaslcr 56 

puntio 60 

rasbora 66 

sophore 60 
Cyrene 37 

Icstna 6. 38 

ocellata 6, 38 

dadiburjori, Laubuca 31 
dadyburjon. Laubuca 31 
Dangila 6, 30. 37, 38 

cuvieri 37, 38 

fasciata 8, 16, 18, 38 

fcstiva 16. 38 

fcstiva var, stcrcus-muscarum 38 

kocdjcm 38 

Icptocheila 37 

lineata 16. 18. 38 

microlepis 38 

ocellata 17-19. 38 

rosea 38 

spilurus 53 

taeniata 38 
daniconius. Rasbora 66, 68 
danielsi. Anus (Hemiarius) 1 10 
Danio (Allodanio) ponticulus 30 
Dasyatis 22 
Datnioides 21. 164 

campbelli 164 

microlepis 16. 17, 164 

polota 164. 165 

quadrifascialus 164. 165 
dayi. Mastacembelus 179. 182 
deignani. Kryptoplerus 149 
deissnen. Osphromenus 174 
deissncn. Parosphromenus 13. 174, 175 
deissnen. Parosphronemus 13. 14. 20 
Dclsmania 23 

dcokhatoides. Doryichthys 16. 20, 158. 159 
dcokhatoidcs, Syngnathus 158, 159 
doekhatoides, Doryichthys 13 
Deplocheilos 45 
Dcrmogcnys 1 54 

cf, oncntalis 13 

cf pusillus 154 

pusillus 5, 154, 156 



devcnten. Capoeta 35 

devcnten, Cyclochcilichthys (Siaja) 35 

de/waani. Cyclochcilichthys 35 

de/waani, Pangasius 130 

diabohca, Barbucca 18, 19, 96, 100. 101 

dialuzona. Acantopsis 5, 99 

Diastatomycter 145 

chapen 9. 145 
dictyogasler, Mastacembelus maculatus var. 181 
dimidiata. Betta 11. 16, 18-20, 172-174 
dinema. Bclodonlichthys 16, 144 
dmema. Wallago 144 
Dinolopterus 128 
Diplocheilichthys 45. 52 

pleurolaenia 52 
diplogrammc. Ophioccphalus 170 
Diplochcilos 45 
Diplomystes 142 
Diploptcrus 1 78 

pulcher 178 
Discognathichlhys 40 
Discognathus 40 

borncensis 40 

vanabllis 40 
Discolabeo 40 

fishen 40 
disparts. Liniparhomaloptera 85 
docmak. Silunis 120 
doriac. Brachygobius 1 3 
doriac. Eucirrhichthys 95. 97 
doriac. Brachygobius 167 
doriae. Gobius 167 
donae. Liocassis 1 1 7 
dorsimaculata, Rasbora 72 
dorsiocellata, Rasbora 16, 18-20, 68, 73 
Dor>ichthys8, II, 22, 158, 159 

bilinealus 158 

boaja 8, 16, 18, 158, 159 

brachyrhynchops 159 

dcokhatoides 16. 20, 158. 159 

doekhatoides 13 

heterosoma 158, 159 

(Microphis) ignoratus 159 

martensi 1 59 

martcnsii 14, 16, 17, 158, 159 

spinosus 159 
dumenli. Cyclochcilichthys 35 
duostigma, Osteochilus 48 
dusoncnsis, Rasbora 13, 16, 20, 66. 68. 73. 74 

eatenatus, Mastacaccmblus 181 
cdithae, Betta 173, 174 
cinthoveni, Rasbora 13 
eintho\enii, Leuciscus 73 
cinthovemi. Polyacanthus 171 
einthovenii. Rasbora 16. 68. 73 
Eirmotus 38 

oclo/ona 16. 20. 38 
clanga. Rasbora 68 
elegans, Rasbora 17, 68, 73, 74 
Eleotns 166 

marmorata 166 

mclanosoma 1 66 

urophthalmoides 167 

urophthalmus 167 
Ellopostoma 21, 94, 103 

megalomyclcr 16. 17. 18. 94. 96 
clongatus. Barbus 61 
elongatus. Punlius 61 
Encheloclanas 1 28 

tapeinopterus 128 
cndecanahs. Puntius 18. 19, 38, 62. 63. 64 
Engraulis 



ROBERTS-WESTERN BORNEO FISHES 



199 



brcviceps 26 

crocodilus 26 

melanochir 26 
cnnealcpis. Rasbora 17. 66. 68. 73, 74 
cnncaporos. osteochilus 13. 17 
enneaporus. Rohita 47 
cnneaporus. Osteochilus 46—48 
cnoploides. Barbus 35 
enoplos. Barbus 33. 35 
cnoplos. Capocta 35 
enoplos, Cyclocheilichthys 33-35 
Epalzeorhynchos 32. 38^0 

kaloptcrum 8. 16-19. 39 

kallopterus 39 
Epal/eorhynchus 38 

kalliurus 32 

siamensis 32 
enthrorhina. Homaloplera 89 
erythrotaenia, Macrognathus 18 
erythrotaenia. Mastacembelus 1 1. 16. 178, 179. 181 
erythrurus. Labeo 39 
Euastrum 59 
Eucirrhichthys 95 

doriae 95. 97 
cuepiptera. Vaillantella 16-19. 97. 109 
euepiplerus. Nemacheilus 109 
eucpiplerus, Vaillantella 109 
eugeneiatus. Callichrous 1 50 
eugeneiatus. Ompok 18. 19. 150. 151 
eugeneiatus. Silurodes 150 
eugrammis, Puntius 16 
eugrammus, Puntius 16. 17, 64. 65 
Eunotia 59 

Eurypleura melanorhyncha 183 
Eustira maassi 31 

Eutropius braehypopterus 128, 129 
everetti. Barbus 64 
everetti, Puntius 64 

falcalus. Puntioplites 60 

fasciata. Catopra 8. 165 

fasciata. Chela 3 1 

fasciata. Dangila 8. 16. 18. 38 

fasciata. Pnstolepis 7. 14. 16, 17, 19, 165 

fasciatus, Acanthophthalmus 95 

fasciatus. Acantophihalmus 95 

fasciatus. Barbodes 64 

fasciatus, Barbus 64 

fasciatus, Cirrhinus 64 

fasciatus, Cobitis 6, 106 

fasciatus, Gastromyzon 17. 18, 84-87 

fasciatus, Nemacheilus 96, 106-108 

fasciatus, Noemacheilus 107 

fasciatus, Pristolepis 8. 165 

fasciatus. Pseudogastromyzon 85 

fasciatus. Puntius 65 

fasciatus. Puntius (Barhodes) 64 

fasciatus. Systomus (Barbodes) 64 

falcifer. Cosmochilus 3. 16. 20. 31 

falcifer, Labeo 41 

falcifer, Lobocheilos 41 

falcifer, Lobocheilus 41 

favus. Mastacembelus 178, 179, 182 

feldmanni. Cynoglossus 184 

festiva. Cyrene 6. 38 

festiva. Dangila 16. 38 

festiva var. stercus-muscanum 38 

filamentosa. Gymnochanda 14, 17. 19. 159, 160 

filamentosus. Parosphromenus 1 74 

filanientus. Macrones (Hemibagrus) 122 

Filirasbora 41 

rubnpinna 41 
fisheri. Discolabeo 40 



flavofasciata. Vaillantella 109, 110 
fluv'iatilis, Tetraodon 6 
fluvidraco, Pimelodus 1 26 
Ruta 183 
foerschi. Belta 172 
foerschi, Puntius 63, 64 
Formosania 83 

lacustre 85 
formosanus, Hemimyzon 85 
formosum, Osteoglossum 23 
formosus, Scleropages 6. 16. 19, 22-24 
fossilis, Cobitis 95 
fossilis. Heteropneustes 12 
fowlerii. Barbus 35 
frenatus, Labeo 39 
fulvidraco, Pimelodus 126 
fuscus. Leiocassis 1 16 
fuscus. Leiocassis (Pseudomystus) 116 
fuscus. Ophicephalus 

gachua, Ophiocephalus 170 
gagora. Arius 1 10 
galeatus, Trachycorystes 12 
Garra 40. 79 

borneensis 1 7. 40 

golyla 79 

taeniata 40 

Gastromyzon 2, 9. 79. 83. 85-87. 91-93 

borneensis 9. 79. 84. 85. 87. 93 

contractus 17, 84-87 

ctenocephalus 83, 87 

fasciatus 17, 18. 84-87 

lepidogaster 85,87 

megalcpis 87 

monticola 87 

nieuwenhuisi 93 

punctulatus 87 

ridens 17, 84, 85, 87, 88 
gigas. Parambassis 164 
Gitantogobius 166 

jordani 166. 167 
gladius. Psephurus 8 
Glaniopsis 83 

hanilschi 84 

multiradiata 84 
Glyptosternon 

platypogonoides 134. 136 

striatus 134 
Glyptostemum 134, 138 

baramensis 138 

kuekenthali 135 

majus 134 

platypogonoides 136 
Glyptothorax 12. 134. 135, 140 

brevipinnis 12 

laak 135 

major 16, 19, 134, 135 

nieuwenhuisi 134, 135 

platypogon 17, 18, 135 

platypogonoides 8. 17. 134-136. 140 

tiong 134. 135 
Gobio javanicus 42 
gobioides. Barbus 30 
Gobionichthys 41 
Gobiopsis oligactis 168 
Gobius 

doriae 167 

javanicus 169 

pisonis 166 

pleurostigma 169 

sadanundio 169 
goniognathus. Clupeichthys 24 
goniognathus. Corica (Clupeichthys) 24 



gonionolus. Puntius 62 

goniosoma. Barbus 61 

goniosoma, Systomus (Barbodes) 61 

goramy, Osphronemus 176, 177 

goramy, Osphronemus 3. 16-18. 177 

gotyla, Garra 79 

gourami, Osphromenus 177 

gracilis, Acanlhopsoides 14, 17-20, 94, 96 

gracilis, Cynoglossus 184 

Gray, Chaea 143 

grayii, Chedrus 30 

griswoldi, Protomyzon 83, 85 

grootii, Catopra 8, 165 

grootii, Pristolepis 165 

guentheri. Mastacembelus 11. 179. 18! 

gulio. Bagrus 120 

gulio. Mystus 126 

gulliveri. Acanthoperca 161 

guntea. Lepidocephalichthys 96 

Gymnochanda 159 

filamentosa 14. 17. 19. 159. 160 
gymnopomus. Butis 13 
Gymnolus 

notopierus 24 
Gyrinocheilus 81 

pustulosus 17, 81, 82 



hagenii. Barbus maculatus var. 61 
hampal var. bimaculata. Barbus 40 
Kampala 40 

bimaculata 17. 40 

macrolcpidota 13, 17, 18, 40 
hanitschi, Glaniopsis 84 
Haplocheilus panchax 31 
Harengula hypselosoma 25 
harmandi, Cosmochilus 31 
hasselti. Belontia 171 
hasselti. Cobitis 103 

hasselti. Lepidocephalichthys 8. 96, 103. 104 
hasselti. Osteochilus 13. 14. 46-49 
hasselti. Polyacanthus 171 
hasselti. Rohita 48 
hasselti. Silunchthys 13. 17. 151 
hassellii. Belontia 13. 16 
hasseltii. Lepidocephalichthys 103 
hasseltii, Osteochilus 16 
hasseltii, Silunchthys 151. 153 
helfrichii, Leuciscus 77 
hclfrichii, Polyacanthus 171 
helfrichii. Rasborichthys 18, 19. 78 
Helgia 88 
Helicophagus 130 
helodes. Botia 102 
Helostoma 177 

lemmmckii 6. 16. 18. 19. 177. 178 

temminkii 178 
Hemianus 21. 110 

danielsi 1 10 

stormi 1 10 
Hemibagrus 116. 120. 122 

wyckii 126 
hcmictenus. Barbodes 61 
hemictenus. Puntius 61 
Hemimy/on formosanus 85 
Hemipimelodus 1 10 

borneensis 110. Ill 
Hemirhamphodon 1. 10. 154-156, 158 

chrysopunctatus 158 

kuekenthali 155 

phaiosoma 13, 16-18, 155. 158 

pogonognathus 11-14. 16. 18-20. 154-158 

sp. undet. 16. 155. 158 



200 



CALIFORNIA ACADEM\' OF SCIENCES 



Hcmirhamphus 

Phai5oma 154. 155 

pogonognathus 13, 155 
Hcmisilurus 144. 145 

chapen 145 

helcrorhynchos 16. I''. 145 

moolenburghi 16. 145 

schilbeides 1 50 

scleronema 144. 145 
hendricksoni. Akysis 1.36 
henverdenii. Bunaka 167 
Heterobagrus 120. 123. 124 

bocourti 120, 123. 124 
Hctcrobranchus 128 

lapoinoplerus 1 28 
hctorolepis. Cynoglossus 1S3. 184 
hclcromorpha. Rasbora 68 
hcteronema. Barhus 33. 34 

helcroncma. Cyclocheilichlhys 14. 16. 18. 1>3. 36 
hclcronema. Cyclocheilichlhys (Siaja) 33, 36 
hcteronema. Oxybarbus 36 
Hctcropneustcs lossilis 12 
helcrorhynchos. Hemisilurus 16. l") 
helcrorhynchos. Schismatorhynchos 17, 79 
helcrorhynchos, Wallago 145 
hclcrorhynchus, Lobocheilos 7Q 
heterorhynchus, Tylognalhus 79 
helerosoma, Doryichlhys 158, 159 
heterosoma. Syngnalhus 159 
Hetcrothnssa 26 
Hderotis 23 
hetcrurus, Bagrus 122 
hcxanema, Laides 14, 16, 131 
hexanema, Lais 8, 131 
hexanema, Pangasius 131 
Hexanematichlhys 1 10 
hexa/ona. Barbodes 65 
hcxa/ona. Barbus 65 
hcxa/ona, Loboles 164, 165 
hcxa/ona, Puntius 65 
hilgendort'ii, Telrodon 188 
hillcn, Polynemus 166 
Hmiantura 2, 10, 22 

signifer 2. 11, 16, 20, 23 
hirsutus. Leiocassis 1 15 
hispidus. Labeo 41 
hispidus. Lobocheilos 41 
hispidus. Lobocheilus 42 
hispidus. Tylognalhus 42 
Huo 144 

hocksi. Pangasius 10. 132. 133 
hocvcnii, Bagrus 121 
hoe\enii. Barbus 4 1 

hoexenii. Lcplobarbus 6. 8. 16, 17. 19. 41 
hoflmanni. Sinohomaloptera 85 
Honialoplera 83. 85. 88. 89. 90. 93. 94 

hilineata 88. 89 

el" stephcnsoni 85. 90. 91 

erythrorhina 89 

javanica 91 

johorensis 93. 94 

ncbulosa 13. 14. 17. 18. 84. 85. 88 

(Neohonialoptcra) johorensis 93. 94 

nigra 9| 

ocellata 88 

ogilviei 14. 85, 89 

ophiolepis 17, 19, 84, 85, 89, 90 

orlhogoniala 18, 84, 85, 89 

slephensoni 17. 18. 84. 90 

tweediei 14, 16, 18. 19. 84. 85. 90. 94 

"assinkii 88 

/nllingeri 17. 84, 85, 91. 92 
Hiimalopleroides 88 



Homaloptcrula 88 

npleyi 88 
hornadayi. Polynemus 166 
hosu. Lcplobarbus 41 
hosii. Liocassis 1 17 
howong. Macroncs I 24 
howong. Mystus 124 
hubbsi. Rasbora 68 
humeralis. Pangasius 16. 131-133 
humilis. Hypergastromyzon LIS, 83. 84, 91-93 
hulchinsoni. Akysis 142 
Hymenophysa 101 

hymenophysa 102 

MaClelland 101 

Maclellandi 101 

macracanthus 102 
hymenophysa, Bolia 8, 18, 96, 101-103 
hymenophysa, Cobilis 101 
hymenophysa, Hymenophysa 102 
Hypergastromyzon 1, 83, 91, 92 

humilis 1, 18, 83, 84, 91-93 

sp. 84 
Hypolophus sephen 22 
hypophthalmus, Callichrous 150 
hypophthalmus. Chela 8, 55 
hypophthalmus, Ompok 8, 17-19, 150, 151 
hypophthalmus, Oxygaster 14, 16. 18, 19, 56 
hypophthalmus. Silurodes 150 
h\pophthalmus. Silurus 150 
Hypostomus sp. 1 1 
Hypselobagrus 120. 123 

macroncma 122 

mieracanthus 120 

nignceps 122. 124 

woim 126 
h\psclopterus. Bagrichthys8. 16. 19. 1 1 1-1 13. 123 
hypscloptcrus. Bagrus 111. 112 
hypsclosoma. Clupca 25 
hypselosoma. Clupeoides 16. 24. 25 
hypsclosoma. Harengula 25 

ignoratus. Doryichthys (Microphis) 159 

ignoralus. Microphis 159 

Inlecypris 59 

inomatus, Leiocassis (Pseudoniystus) 1 16 

insignis. Brcitensteinia 8. 140 

intcrmcdius. Ostcochilus 16. 19. 46—18. 50 

IMS. Barhus 61 

jacobsoni. Rasbora 68 

jaculator. Toxotes 165 

jaculator. Sciaena 165 

jagur. Clarias 1 28 

janthochir. Cyclocheilichlhys 17-19, 33. 36 

janthochir. Cyclocheilichlhys (.AneniatichthNs) 36 

janthochir. Systomus 35. 36 

javanensis. Monopterus 183 

javanica, Homaloptera 91 

javanicus. Acanlhophthalmus 97 

javanicus, Acantophthalmus 95, 97 

javanicus, Gobio 42 

javanicus, Gobius 169 

javanicus, Lobocheilus (Gobionichlhys) 42 

ja\anicus. Oryvias 1 

jentinkii, Osteochilus 52 

johorensis, Homaloptera (Neohonialoptcra) 93, 94 

johorensis, Mystus 122 

johorensis, Neohomaloptera 13, 84, 85, 94 

johorensis, Puntius pcnta/ona 65 

jordani, Gigantogobius 166, 167 

juaro, Pangasius 133 

julheni, .Syngnathus 1 59 

liirongcnsis, Pseudogobiopsis 168 



jurongensis, Vaimosa 168 

kabilia. Calamiana 168 

kahajanensis. Osteochilus 16, 18. 46. 47, 49. 50 

kahajanensis. Rohita 49 

kahajani, Barbus 65 

kajan, Macrones 124 

kajan, Mystus 124 

Kalimantama 27, 29, 40 

lawak 18, 40, 41 
kalliurus. Epal/eorhynchus 32 
kalloptcrus. Epalzeorhynchos 39 
kaloehroma. Leuciscus 74 
kalochroma. Rasbora 17. 68. 74 
kalopterum. Epalzeorhynchos 8. 16-19. 39 
kalopterus. Barbus 38. 39 
kappenu. Osteochilus 17-19. 45-47. 49. 51 
kappcnii. Rohita 49 

kapuasensis. Cynoglossus 16. 20. 183-184 
kapuasensis. Nemacheilus 17. 94. 97. 107 
kapwasensis. Cynoglossus 184 
Kaupia 1 58 

keithi. Mastacembelus 178. 179. 181 
kilogcneion. Rohita 45 
kocdjem. Dangila 38 
korlhausac. Pectenocypris 57. 58 
krcmpfi. .Albulichthys 28 
Kryptoplcrichthys 145 

bieirrhis 147 

lais 147 

macrocephalus 145. 147 

palcmbangensis 145 
Kryptoplerus22. 145-148. 150. 152 

apogon 16. 145. 146 

bicirrhis 16. 18. 19. 146-148. 150 

cryptoptcrus 16-18. 146. 147 

deignani 149 

lais 18. 20. 146. 147 

limpok 16. 18. 146. 147 

macrocephalus 13. 14. 146-149 

microncma 18. 19. 146 

micropus 145-147 

minimus 146 

minor 14. 17. 20. 149. 150 

schilbeides 16. 18. 146. 150 

sp. undct. 17. 150 
kuckenthali. Glyptostemum 135 
kuekenthali. Hemirhamphodon 155 
kuhh. Rohita (Rohita) 48 
kuhhi. Cobilis 95 
kuhlii. Oslcochilus 48 
kuhlii. Polyacanthus 171 
kukcnthali. Hemirhomphodon 155 
kusancnsis. Barbus 60 

laak. Ghptothorax 135 
Labeo 

bicolor 39 

chrysophekadion 45 

erythrurus 39 

ralcilcr41 

frcnatus 39 

hispidus 41 

munensis 39 

oblongus 32 

pleurotacnia 52 

rohitoides 45 

(Diplochcilos) rohitoides 52 
Labeobarhus tambra 8 1 

tambroides 81 
Labio 37 
Labiobarhus 5. 37 

Icplocheilus 5. 37 



ROBERTS-WESTERN BORNEO FISHES 



201 



lipocheilus 5, 37 

lineatus 38 
Labrus tnchoplenis 177 
lachnostoma. Acantopsis 96, 99. 100 
laclogeneus, Nemacheilus 19, 97, 107 
lacustre, Formosania 85 
laevis, Barbichthys 8. 16-19, 30 
lacvis var, sumatranus, Volz 30 
lacvis, Barbus 29, 30 
Laides 1 28, 1 3 1 

hexanema 14, 16, 131 
Lais 1 3 1 

hexanema 8, 131 
lais, Cryplopterus 147 
lais, Kryptopterichthys 147 
lais, Kryploptenis 18, 20, 146, 147 
lais, Silurus 147 
lala, Chanda 161 
lamani. Bagrus 120 
lampta, Cypnnus 40 
laosensis, Macrochinchthys 45 
laleristriga, Barbodes 65 
latcnslnga, Barbus 65 
lalenstriga, Puntius 13. 17. 65 
lalenstriala, Rasbora 73 
latcnslriata, Systomus (Barbodes) 65 
Laubuca 3 1 

dadiburjori 31 

dadyburjon 31 

(Euslira) maassi 31 
laubuca, Cypnnus 31 
lawak, Barbus 40, 41 
lawak, Kalimantania 18, 40, 41 
lawak. Puntioplites 29 
lawak, Puntius 40. 41. 60 
lawak. Systomus 40 
lazcra. Clanas 1 2 
leerii. Tnchogaster 16-20, 177 
leerii, Tnchopodus 177 
leerii. Tnchopus 177 
leerii. Wallago 3, 8. 16. 151. 152 
leiacanthus. Clanas 13. 18. 127. 128 
leiacanthus, Leiocassis 116 
leiacanthus, Leiocassis (Pseudomystus) 1 16 
leiacanthus. Ompok 151 
Leiocassis 9, 12, 111, 115-119, 124, 137, 138 

armatus 17, 18, 116, 117 

brcviceps 

chaseni 1 1 7 

doriae 1 1 7 

cf, fuscus 1 1 6 

hirsutus 1 1 5 

hosii 1 17 

inomatus 1 16 

leiacanthus 1 16 

mahakamensis 116. 126 

micropogon 17. 116, 117 

moeschii 120 

myersi 19, 116-118 

poecilopterus 1 1 7 

(Pseudomystus) bicolor 116 

(Pseudomystus) breviceps 1 16 

(Pseudomystus) fuscus 116 

(Pseudomystus) inomatus 1 16 

(Pseudomystus) leiacanthus 116 

(Pseudomystus) mahakamensis 1 16 

(Pseudomystus) moeschii 116 

(Pseudomystus) robustus 1 16 

(Pseudomystus) siamensis 116 

(Pseudomystus) stenomus 116 

(Pseudomystus) vaillanti 116 

regani 1 17 

robustus 1 16 



sp. indet. 119, 120 

stenomus 116, 120 

vaillanti 119. 120 
leiurus. Tetraodon 16. 186. 188 
Lepidocephalichthys21. 94. 103. 104 

guntea 96 

hasselti 8. 96, 103, 104 

hasseltii 103 

lorcntzi 96, 103. 104 

pallens 106 

pristes 16, 18, 19, 96, 105 

sandakanensis 96 

sandakanus 103 
Lepidocephalus I, 21, 94, 106 

macrochir 96, 106 

pallens 106 

spectrum 1, 19, 94, 96, 106 
lepidogaster. Gastromyzon 85. 87 
Lepidoglanis 85 

monticola 85 
leptaspis. Arius 6 
Lcptobarbus 3. 41 

hoevenii6, 8, 16. 17, 19, 41 

hosii 41 

melanopterus 18, 19. 41 

melanotaenia 41 
Leptocephalichthys 95 
leptochcila. Dangila 37, 38 
leptacheilus, Labiobarbus 37, 38 
Icptonema, Phalacronotus 145 
leptonema, Silurus 145 
leptosoma, Rasbora 66, 68, 72, 77 
Icucorhynchus, Achiroides 18, 19. 183 
leucorhynchus, Akysis 140 
Leuciscus 55, 57 

argyrotaenia 66, 67 

bankanensis 70 

Ccphalotaenia 72 

einthovenu 73 

helfrichii 77 

kalochroma 74 

macrochirus 44, 45 

oxygasler 55, 56 

thynnoides 80 

tnnema 42, 44 

uranoscopus 44, 45 

zamhczcnsis 30 
leveretti, Beaufortia 85 
liacanthus, Clanas 127 
Liachirus mclanospilus 183 
Ilea, Baganus22. 133, 134 
limpok. Cryplopterus 147 
limpok. Kryptopterus 16. 18. 146. 147 
limpok. Silurus 147 
lingua. Cynoglossus 186 
lincata, Dangila 16, 18, 38 
lineatus, Barbus 35, 65 
lineatus, Cyclocheilichthys 35 
lineatus, Labiobarbus 38 
lineatus. Puntius 18. 64. 65 
lineatus. Tetraodon 188 
Liniparhomaloptera 

dispans 85 
Liocassis 1 16 

baramensis 1 1 7 

doriae 1 1 7 

hosii 1 17 

merabensis 1 17 
lipocheilos. Chondrostoma 42 
lipocheilus, Labiobarbus 37 
lipocheilus, Lobocheilus (Gobionichthys) 42 
Lissorhynchus 40 
lissorhvnchus, Platvcara 40 



lithostoma, Pangasius 16, 131-133 
lobocheiloides. Schizmatorhynchus 79 
Lohocheilos 41 

cobilis 32 

hetcrorhynchus 79 

hispidus 41 

pleurotaenia 45 
Lobocheilus 41, 42, 79 

bo 41, 42 

falcifer 41 

(Gobionichthys) javanicus 42 

(Gobionichthys) lipocheilus 42 

heterorhynchos 79 

hispidus 13, 42 

pleurotaenia 52 

rohitoides 52 

sp. 16. 19 
Lobotes hexazona 164. 165 
longibarbis. Pangasius 131 
longipectoralis. Nemacheilus 107 
cf. longipectoralis. Nemacheilus 19. 97, 108 
longipectoralis, Noemacheilus 108 
lorentzi, Acaniophthalmus 95. 104 
lorentzi, Lepidocephalichthys 96. 103. 104 
Lucioccphalus 1 78 

pulcher 14. 16, 178 
Luciosoma 42, 43. 44 

Pellegrini 43. 44 

setigerum 16, 17, 43, 44 

spilopleura 16-19,42^4 

tnnema 8. 16-19,43, 44 

(Tnnematichthys) trinema 44 

webcni 43 
lucius, Channa 13. 14, 16, 18. 20. 170 
lucius. Ophicephalus 8. 170 
Lycothnssa 26 

crocodilus 16. 18. 19, 25, 26 



maassi. Chela 13, 18, 19 

maassi. Chela (Allochela) 31 

maassi. Chela (Malayochela) 31 

maassi, Eustira 31 

maassi, Laubuca (Eustira) 31 

maassi, Vaillantella 17, 19, 97, 109 

MacClellandi. Hymenophysa 101 

maclellandi, Hymenophysa 101 

macracantha, Botia 8, 16-19, 96. 102 

macracanthus, Bagnchthys 1 1 1-1 13 

macracanthus, Barbus 35 

macracanthus, Cobitus 102 

macracanthus, Cyclocheilichthys 35 

macracanthus, Hymenophysa 102 

macrocephalus, Cryplopterus 148 

macrocephalus, Kryptoptenchthys 145. 147 

macrocephalus. Kryptopterus 13, 14. 146-149 

macrochir, Belodontichthys 144 

macrochir. Chela 8 

macrochir, Cobitis 106 

macrochir, Macrochinchthys 17, 19 

macrochir, Lepidocephalus 96, 106 

Macrochinchthys 2, 8. 44. 45. 144 

laosensis 45 

macrochir 17. 19 

macrochirus 45. 55 

sn\dcri 45 

uranoscopus 45 
macrochirus. Leuciscus 44. 45 
macrochirus, Macrochinchthys 45, 55 
Macrognathus 2, 178, 180 

aculeatus4, 14. 16. 18-20. 178. 180 

aral 180 

erythrotaenia 18 



202 



CALIFORNIA ACADEMY OF SCIENCES 



perakensis 1 79 

siamcnsis 180 
macrolcpidola. Capoeta 40 
macrolcpidota. Hampala LI, 17, 18. 40 
macrolcpis, Bogoda 162 
macrolcpis, Chanda 16. 17. 19. 162 
macrolcpis. Channa 18 
macrolepis. Parambassis 159. 162, 163 
macroncma, ."Xkysis 136 
cL macronema, Akysis 136 
macroncma. Bagrus 120 
macronema. Hypsclohagrus 122 
macroncma. Callichrous 8 
macronema. Silurodes 150 
macronema. Silurus 1 50 
macronemus. Bagrus 120, 122 
Macrones 120. 124 

bo 124 

bongan 122, 124 

cavasius 1 22 

howong 124 

kajan 124 

micracanthus 120 

nignceps 8, 122 

planiceps 122 

wolffii 126 

u^ckii 126 

(Hcmibagrus) lilamenlus 122 
Macromchthys 120 
macrophthalmus. PoKncmus 16. 166 
Macropodus pugnax 173 
macropterus, Bagnchlhys 19. I 12-1 15 
macroptcrus. Pseudohagnchthvs 1 14 
Macropleronotus. magur 127 
macroptcrus. Bagnchlys 16 
macroplcrus. Bagroidcs 111. 114 
macrostoma. Bella 171. 172 
maculata. Pararhynchobdella 180 
maculata. Rasbora 69 
maculata. Rhynchobdella 180 
maculatus. Barbus 60. 61 
maculatus, Mastacembelus 1 1. 13, 14, 16-20, 178- 

181 
maculatus var chrvsogaster 181 
maculatus var. dictyogaster 181 
maculatus, Systomus (Barbodes) 61 
niaculiceps. Nemacheilus 17, 19. 97. 108 
iTiagnons. Calamiana 168 
magur. Clarias 1 27 
magur. Macropleronotus 127 
inahakamensls, Leiocassis 116, 126 
mahakamensis, Leiocassis (Pseudomystus) 1 16 
major, Akysis 1 34 

major. Glyplolhorax 16, 19, 134, 135 
majus, Glyptoslemum I 34 
malabarica. Calopra 165 
malayanus, Osphromenus 176 
malayanus, Sphaerichlhys 176 
Malayochela 3 1 
niarchei, Micracanthus 171 
margmata. Prislolepis 165 
marginalus. Barbus 45 
marginatus. Myslacolcucus 17. 45 
marginatus. Ophiocephalus 46 
mariac, Acanlhophlhalmus 98 
marmorala. Eleolns 166 
marmorata. Oxyeleolns 16. 19, 166. 167 
rnarmoratus. Nandus 164 
martcnsi. Doryichthys 159 
mancnsn. Doryichthys 14. 16, 17. 158. 159 
marlcnsii. Syngnathus 8. 159 
marulioides. Channa 8. 17. 170 
marulioides. Ophiccphalus 170 



marulioides, Ophiocephalus 8 
Mastacacemblus. catenalus 181 
Mastacembelus II. 22. 178-182 

argus 181 

armatus 178. 179. 181. 182 

billitonensis 181 

canciloides 153 

catenalus 181 

dayi 179, 182 

eatenatus 181 

crythrotacnia II. 16. 178. 179. 181 

favus 178, 179, 182 

gucntheri II, 179, 181 

keilhi 178. 179. 181 

maculatus II, 13, 14, 16-20. 178-181 

maculatus var, chrysogaster 181 

maculatus var. dictyogaster 181 

notophlhalmus 16. 17. 178. 179. isi. 182 

perakensis 181 

undulalus 178, 179, 181. 182 

umcolor 17, 178-180, 182 

vaillanli 179. 181 
mastacembelus, Ophidium 180 
Mastocembelus 180 
Malsya 45 

tnegalepis, Gaslromy/on 187 
megalolepis. Chela 31, 57 
mcgalomycter. Aperioptus 103 
megalomyctcr, Ellopostoma 16, 18. 94. 96. 103, 

104 
megalops, Cyclocheilichthys 37 
mcgalura, Setipinna 26 
Megarasbora 44 
mcinkeni, Rasbora 69 
mekongensis. Cyclocheilichthys 35 
mcladcrma, Clarias 4, 127 
melanochir. Anus 16, 18, 19. 21. I 10. 1 I 1 
mclanochir, Cephalocassis 1 10 
melanochir. Coilia 26 
mclanochir. Engraulis 26 
melanochir. Setipinna 18. 20. 24. 26 
melanoderma, Clarias 4, 20, 127 
melanogaster. Acrochordonichthys 137, 138 
cl melanogaster. Acrochodonichthys 14, 136. 138 
mclanopleura. Ostcochilus 17. 19, 46. 47 
melanopleura. Rohila 45. 50 
niclanopleurus. Ostcochilus 50. 51 
mclanoptcra. Channa 170 
melanopterus. Bagroides 1 1 5 
melanopterus. Balanliocheilos 8. 17. 19. 29 
melanopterus. Barbus 29 
melanopterus. Leplobarbus 18. 19.41 
melanopterus. Ophiccphalus 170 
melanopterus. Ostcochilus 50-51 
mclanorhyncha. Euryplcura 183 
melanorhyneha. Synaptura 183 
mclanorhynchus. Achiroidcs 16. 18-20. 183 
mclanorhynchus. Plagusia 183 
mclanosoma. Clarias 127 
nielanosoma. Eleotris 166 
mclanosoma. Ophiocephalus 170 
melanospilus, Liachirus 183 
nielanotaenia, Leplobarbus 41 
mclapterus, Bagroidcs 8, 16, 19. I 15 
mclasoma, Channa 1 70 
melasoma, Clanas 127 
mclasoma, Ophiccphalus 1 70 
merabensis, Liocassis I 1 7 
Micracanthus 171 

marchei 1 7 1 
micracanthus, Bagrus 120 
micracanthus, Hyselobagrus 120 
micracanthus, Macrones 120 



micracanthus. Mystus 8, 14. 16. 18-20. 120, 121 

micranodus, Bagnchlhys 19, 112-115 

Microcanthus 171 

microcephalus, Ostcochilus 16-19, 46, 47, 52 

microcephalus, Rohita 52 

microlepis, .Ambassis 160, 162 

microlepis, Ambassis (Bogoda) 162 

microlepis, Barbus 78 

microlepis, Barynolus 8. 36 

microlepis, Capoeta 33, 36 

microlepis, Cyclocheilichthys 18-20, 33, 35, 36 

microlepis, Cynoglossus 183, 184, 185 

microlepis, Dangila 38 

microlepis, Datnioides 16, 17, 164 

microlepis, Neobarvnotus 36 

microlepis, Parambassis 162 

microlepis, Rohtee 78 

microlepis, Rohteichthys 18, 19. 78. 79 

microlepis. Systomus 78 

microlepis. Toxotes 8. 16, 18. 19, 165 

Microncma 145 

phalacronotus 149 

lypus 145, 149 
micronema, Callichrous 150 
micronema. Cryptoplerus 149, 150 
microncma, Kryptoplerus 18, 19, 146 
micronema, Pangasius 8, 131-133 
micronemus, Silurus 145, 149 
micropeltes, Channa 8, 13, 169, 170 
micropelles. Ophiccphalus 170 
micropeltes. Ophiocephalus 8 
Microphis 

caudatus 159 

ignoralus 1 59 
micropogon. Bagrus 1 1 7 
micropogon. Cryplopterus 145 
micropogon. Leiocassis 17. 116. 117 
micropogon. Phalacronolus 145 
micropogon. Silurus 145 
microps. Sundasalanx 18. 27 
micropus, Cryptoplerus 147 
micropus. Kryptoplerus 145-147 
microstoma, Parhomaloptera 84 
micruroptcrus, Phalacronotus 149 
minimus. Kryptoplerus 146 
mino. Batrachocephalus 1 10 
minor, Kryptoplerus 14, 17, 20, 149. 150 
minor. Sundasalanx 2 
Misgumus barbaluloidcs 8. 103 
modeslus. Chonerhinos 8. 19. 184. 186, 187 
modeslus, Chonerhinus 187 
modeslus, Tetraodon (Arothron) 187 
modeslus, Xenopterus 8 
Modigliana 106 

papillosa 106, 107 
mocschii, Leiocassis 120 
Monocirrhus 164 
Monoplerus 183 

albus 16, 183 

javanensis 183 
monlanus, Ophiocephalus 
monlicola, Gastromy/on 87 
monlicola, Lepidoglanis 85 
moolenburghae, Pseudculropius 18, 19, 129 
moolenburghi. Hemisilurus 16, 145 
morala. Cypnnus (Morulius) 45 
Morulius 45 

chrysophekadion 16, 45 
Mugilogobius 167, 168 

sp undet. 13, 16, 168 
niullcn. Silurus 1 5 I 
mullitilis. Polislonemus 166 
nnillililis. Polynemus8. 16. 18. 19. 166 



ROBERTS-WESTERN BORNEO FISHES 



203 



multifills. Trichidion 166 

mulliradiala, Glaniopsis 84 

munensis. Labeo 3Q 

muraemformis, Acantophthalmus 96 

myersi, Acantophthalmus 96 

myersi. Leiocassis 19, 1 16-1 18 

mycrsi. Rasbora 13. 16-18. 66. 69. 74. 75 

cf. myersi. Rasbora 74 

myersi. Pseudogastromyzon 2. 82 

Mystacoleucus 45 

margmatus 17, 45. 46 
mysla.x. Ophicephalus 170 
Mystus4, 8, 12, 21, 22. 116. 120. 122-125 

baramcnsis 122-124 

bimaculatus 121 

bo 124 

hocourti 124 

bongan 1 24 

cavasius 122-124 

cf. gulio 4 

cf. planiceps 123 

chilala 24 

gulio 4. 126 

howong 1 24 

johorensis 122 

kajan 124 

micracanthus 8. 14, 16, 18-20, 120. 121 

nemurus 13. 16-19, 120-126 

nigriceps 14, 16-19. 120, 122-124 

olyroides 16. 124-126 

pahangensis 122 

planiceps 123 

wolffii 126 

wycki 126 

wyckii 17. 19. 126 

nama. Chanda 160. 161 
nandoides. Catopra 165 
Nandus 164 

borneensis 164 

marmoratus 164 

nandus 164 

nehulosus 14. 16. 18-20. 164 
nandus. Coius 164 
nandus. Nandus 164 
nanensis. Banlius 31 
Nangra 140 

nantus. Chonerhmus 187 
naritus. Tetraodon 186. 188 
nasula. Platycara 40 
nasulus, Pangasius 16, 130, 133 
Navicula 59 

nebulosa, Homaloptera 13, 14, 17, 18, 84, 85. 88 
nebulosus. Bcdula 164 
nebulosus. Nandus 14. 16. 18-20. 164 
nebulosus. Wallago 152 
nefastus. Chonerhinos 18. 184. 186. 187 
ncilli, Osteochilus 48 
Nemacheilos 106 
Ncmacheilus 3, 6. 21. 94. 100. 106. 108. 109 

beavani 107 

cf. longipectoralis 19. 97. 108 

cupiplerus 109 

Iasciatus96, 106. 107. 108 

kapuascnsis 17. 94. 97. 107 

laclogeneus 19. 97. 107 

longipectoralis 107 

macuhceps 17. 19, 97, 108 

saravacensis 14, 97, 107. 108 

selangoncus 14. 16-20. 97. 107. 109 
Ncmalocheilus 106 
nemurus, Bagrus 120, 121 
nemurus, Macrones 8 



nemurus, Mystus 13, 16-19, 120-124, 126 
Neobarynotus 33 

microlepis 36 
Neochela 3 1 
Neogastromyzon 83, 91. 93 

nieuwenhuisi 83. 84 

nicuwenhuisii 93 

pauciradiatus 83. 93 
Neohomaloptera 83. 93. 94 

johorensis 13. 84. 85. 94 
Ncopangasius 131 

nicuvvenhuisii 1 31 
nieuhofi. Clarias 128 
nieuwenhuisi, Bagarius 133 
nieuwenhuisi, Gastromyzon 93 
nieuwenhuisi, Glyptolhora.x 134, 135 
nieuwenhuisi. Neogastromyzon 83. 84 
nieuwenhuisi. Pangasius 131 
nieuwcnhuisii, Neogastromyzon 93 
nieuwenhuisii, Neopangasius 131 
nieuwenhuisi!. Pangasius 132 
nigra. Homaloptera 91 
nigriceps. Bagrus 120. 122 
nigriceps, Hypselobagrus 122, 124 
nigriceps, Macrones 122 
nigriceps, Mystus 14, 16-19. 120. 122-124 
nigriloba. Crossocheilus oblongus var. 32 
nigrovindis. Tetraodon 186. 188 
mm. Puntius 60 
nitidus. Barbichthys 30 
nitidus. Barbichthys laevis 30 
nobilis, Ctenops 176 
nobilis. Osphromenus 177 
Noemacheilus 5. 106. 107 

fasciatus 107 

longipectoralis 108 
notatus. Osphromenus 176. 177 
notophthalmus. Mastacembelus 16. 17. 178. 179, 

181, 182 
Notopterus 24 

borneensis 17-19. 24 
notopterus, Gymnotes 24 
Nukta 79 

nukta. Schismatorhynchos 79 
nunus. Brachygobius 167. 168 

oblonga. Cobitis 97 

ohlongus. Acantophthalmus 17. 19. 96-98 

oblongus. Crossocheilus 13. 16-18. 20. 32. 33 

oblongus. Crossochilus 32 

oblongus var. mgnloba. Crossocheilus 32 

oblongus. Labeo 32 

obscurus. Acrochordomchthys 1 38 

obtusirostns. Barbus 45 

ocellata. Cyrene 6. 38 

ocellata. Dangila 17-19. 38 

ocellata. Homaloptera 88 

oclocirrhus. Cobitis 95 

octozona. Eirmotus 16. 20. 38 

oetik. .Anus 1 10 

ogilviei. Homaloptera 14. 85. 89 

olfax. Osphromenus 177 

oligactis. Gobiopsis 168 

oligolepis. Anabas 8 

oligolepis, Osteochilus 53 

oligolepis. Rohita 53 

Olyra 126 

olyroides, Mystus 16, 124-126 

Ompok4, 150-152 

bimaculatus 4 

eugeneiatus 18, 19. 150. 151 

hypophthalmus 8. 17-19, 150, 151 

leiacanthus 151 



sabanus 18, 151, 152 

siluroides 150 

weberi 10. 151 
Ophicephalus 169. 170 

bankanensis 169 

bivittatus 170 

lucius8. 170 

marulioides 8. 170 

melanopterus 1 70 

melasoma 1 70 

micropeltes 170 

mystax 170 

pleurophthalmus 170 

punclatus 169 

rhodotaenia 1 70 

serpentinus 170 

spintalis 170 

stevensii 170 

striatus 8. 170 

urophthalmus 170 
Ophidium 

aculcatum 178. 180 

mastacembelus 180 
Ophiocephalus 169 

baramensis 170 

bistrialus 170 

diplogramme 170 

gachua 170 

melanosoma 170 

polylepis 170 

studen 170 

vagus 8 
ophiolepis, Homaloptera 17. 19, 84, 85, 89, 90 
Opsandium 30 
Opsanus 30 
Oreichthys 38 
oresigencs, Barbus 60 
ornalus. Pelteobagrus 111. 126, 127 
ornatus, Pseudobagrus 126 
orientahs, Channa 10. 18. 169. 170 
orthogomata. Homaloptera 18, 84, 85, 89 
Oryzias 1 

Oryzias javamcus 1 
Oshimia 171 

osphromenoides, Sphaenchthys 13. 175. 176 
osphromcnoides selatanensis. Sphaerichthys 1 76 
Osphromenus 177 

deissneri 1 74 

gourami 177 

gouramy 176 

malayanus 176 

nobilis 177 

notatus 176, 177 

olfax 177 

satyrus 177 

siamensis 177 

striatus 171 
Osphronemus 3. 177 

goramy 3, 16-18, 177 
Osteochilus 1-3. 10. 21. 45. 46. 49, 50. 52. 142 

borneensis 16. 18. 46. 47 

brachynotopterus 52 

brcMcauda 48 

duostigma 48 

enneaporos 13. 17 

enneaporus 46—48 

hasselti 13. 14. 46^9 

hasseltii 16 

hasselti tweediei 48 

intermedins 16. 19. 46-48, 50 

jentinkii 52 

kahajanensis 16. 18, 46, 47, 49, 50 

kappenii 17-19,45-47.49, 51 



204 



CALIFORNIA ACADEMY' OF SCIENCES 



kuhlii 48 

melanoplcura 17. 1*^. 46. 47 

nuianoplcurus 50. 51 

mclanoplerus 50-5 1 

microcephalus 16-19. 46. 47, 52 

neilli 48 

ohgolcpis 53 

plcurolaenia 46. 47 

saravvakcnsis 46 

scapulans 47 

schlegcli 16. 19,46,47. 5.3 

spilurus 13. 14, 16. 18-20.46,47. 53 

tnporos 16. 17. 19. 54 

inporus 46-49. 54 

Miialoidcs 47 

Mltatus 8. 47. 52 

vvaaniicrsi 18. 19 

waandcrsu 13. 16, 46, 47, 54, 55 
Osteoglossiim 23 

formosum 23 
Oxybarbus 33 

heteronema 36 
Oxyeleolns 166 

marmorala 16. 19. 166, 167 

urophlhalmoides 20. 167 

urophlhalmus 167 
Oxygaster 55. 56. 76. 77 

anomalura 16. 18. 55. 56 

hypophthalmus 14. 16. 18. 19. 56 

oxygaster 55 

oxygaslroidcs 17-19. 56. 57 
oxygaslcr. Chela 56 
oxygaster. Cypnnus 56 
oxygaster, Leuciscus 55. 56 
oxygaslcr, Oxygaster 55 
oxygastroidcs. Chela 57 
oxygastroides, Leuciscus 57 
oxygastroides, Oxygaster 17-19. 56. 57 

pabda. Silurus (Callichrous) 150 
pachyderma. Acrochordonichthys 137. 138 
Pachystomus 30 

schagra 30 
padangcnsis. Capoeta 45 
pahangensis. Acanthophthalmus 106 
pahangcnsis. Mystus 122 
palavanensis. Barbus 61 
palavcncnsis. Barbus binotatus var 6 1 
palenibangensis, Kryptoptenchthys 145 
palemhangensLS, Silurus 145-147 
palenibangensis. Tetraodon 8. 16. 20. 186, 188 
pallens, Lepidocephalichthys 106 
pallcns, Lepidocephalus 106 
paludicola, Parosphromenus 174 
Panchax pictum 1 7 1 
panchax, Haplocheilus 31 
Pangasius 3, 13, 131-133 

de/waani 130 

hexanema 1 3 1 

hocksi 10, 132, 133 

humeralis 16. 131-133 

juaro 133 

hthosioma 16. 131-133 

longibarbis 131 

micronema 8. 131-133 

nasutus 16. 130. 133 

nieuwenhuisi 131 

nicuwenhuisii 1 32 

pangasius 12 

polyuranodon 16. 130-133 

ponderosus 1 30 

rios 132 

sp, undet, 131 



pangasius, Pangasius 12 

pangasius, Pimelodus 131 

pangia, Acaniophthalmus 96, 98 

pangia, Cobitis 95 

Pangio 95 

papillosa, Modigliana 106, 107 

Papyrocranus 24 

Parachela 

breitenstcini 8 
Paracrossochilus 39. 40, 57 

acerus 17, 57 

bicomis 57 

vittatus 13, 17, 57 

paracrossocheilus. Crossocheilus 

paradiseus. Polynemus 166 

Paradoxodacna 1. 160. 161 

piratica I. 8, 16, 18, 19. 159-161 
Parakysis21, 136, 141. 142 

anomalopteryx 18, 19. 142. 143 

verrucosa 14 1-143 
Parambassis 1. 161. 163. 164 

apogonoides 1, 11, 159, 161. 162 

confinis 164 

gigas 164 

macrolcpis 159. 162. 163 

microlepis 162 

wolffi 164 

vvolITu 159. 162-164 
Paranandus 165 
Paraphiocephalus 169 
Pararhynchobdella 1 80 

maculata 180 
Parapsilorhynchus 40 
Parasilurus 143. 144 
Parhomaloptcra 

microstoma 84 
Parluciosoma 21. 44. 66 
Parophiocephalus 171 

unimaculatus 171 
Parosphromenus 174 

deissneri 13. 14. 20. 174. 175 

filamentosus 1 74 

paludicola 174 

parvulus 13. 174. 175 
parvulus. Parosphromenus 13. 174. 175 
pauciperfbrata. Rasbora 13. 14. 16-18. 66. 69. 74, 

77 
pauciradiatus, Neogastromy/on 83. 93 
Pcctcnocypris 1. 2. 57. 59 

balacna 1.2. 16. 18. 58, 59 

korlhausae 57. 58 
Pellegrini. Luciosoma 43, 44 
Pclleobagrus 111. 126 

ornatus 111. 126. 127 
pentapterus. Clanas 128 
penta/ona, Barbus 65 
penta/ona johorensis, Punlius 65 
penta/ona penta/ona, Puntius 65 
pentazona, Puntius 13, 16, 63-65 
perakensis, Cobitophis 96, 97 
perakensis, Mastaccmbelus 179. 181 
Perca scandens 1 7 1 
Peridinium 58. 59 

phaiosoma. Hemirhamphodon 13. 16-18. 155. 158 
phaiosoma. Hemirhamphus 154. 155 
phaiosoma. Silunchthys 14. 19, 151, 153, 154 
phaisoma, Silurus 151 
Phalacronotus 145 

leploncma 145 

micropogon 145 

micruropterus 149 
phalacronotus. Micronema 149 
phalacronotus. Silurus 149 



phasa. Clupea 26 
philippina. Rasbora 69 
Philypnoidcs 169 

surakartensis 169 
picta. Betta 171 
pictuni. Panchax 171 
Pimelodus 

anus 1 10 

baganus 1 33 

borneensis 110. Ill 

cyanochloros 1 35 

lulvidraco 126 

pangasius 1 31 

piratica 9 

platypogon 135 

plalypogonoides 136 

\ariegatus 1 38 
pinguis. Tetraodon 188 
Pinnulana 59 

piratica. Paradoxodacna 1.8. 16. 18. 19. 159-161 
pisonis, Gobius 166 
Plagusia 

melanorhynchus 183. 184 

waandersi 1 86 
planiceps. Macrones 122 
planiceps. Mystus 123 
Platacanthus 103 

agrensis 103 
Platycara 40 

nasuta 40 
platyccphalus. Acrochordonichthys 137 
platycephalus. Callieleotns 166 
platypogon. Glyptothorax 17. 18. 135 
platypogon. Pimelodus 135 
platypogonoides. Pimelodus 1 36 
platypogonoides. Glyptostemon 134. 136 
plalypogonoides. Glyptostemum 8. 17. 134-136. 

140 
Platystacus chaca 143 
pleurophthalma, Channa 18 
pleurophthalmus. Channa 17 
pleurophthalnius, Ophicephalus 170 
pleurostigma. Gobius 169 
picurotaenia. Diplocheilichthys 51 
pleurotaenia. Labco 52 
pleurotaenia. Lobochilos 45. 52 
pleurotaenia. Osteochilus 46. 47 
podonemus, Barbus 43 
Poecilia 106 

poecilopterus, Leiocassis 1 17 
pogonognathus. Hemirhamphodon 11-14. 16. 18- 

20, 154-158 
Polistonemus 166 

niultililis 166 
polota. Coius 165 
polola. Datnioides 164. 165 
Polyacanthus 

cinthovenii 1 7 1 

hasselti 171 

hell'nchii 171 

kuhh 171 
Polycentropsis 164 
Polycentrus 164 
polylepis, Ophiocephalus 170 
poKlepis, Thynnichthys 16-19, 80, 81 
Polynemus 166 

borneensis 166 

hilleri 166 

hornadayi 166 

macrophthalmus 16, 166 

multilihsS. 16. 18. 19. 166 

paradiseus 166 

quaterdecimfths 166 



ROBERTS-WESTERN BORNEO FISHES 



205 



polyporos. Chnsophekadion 45 

polyporos, Rohita 45 

polyspilos, Barbus 61 

polyuranodon. Pangasius 16. 130-133 

pondcrosus, Pangasius 130 

polamophllus. Clupea (Clupeoides) 25 

praecox, Sundasalanx 27 

pristes. Lepidocephalichthys 16, 18. 19,96, 105 

Prislolepis 164 

fasciata 7, 14, 16, 17, 19, 165 

fasciatus 8, 165 

grootii 165 

marginata 165 
proctozysron, Barbus 60 
proctozysron, Punlius 59 
proctozysron, Puntiopliles 59 
Prosphromenus 175 
Protomyzon 

aphelocheilus 84 

bomeensis 84 

gnswoldi 83, 85 

whitehcadi 85 
Pseudambassis 161 
Pseudeutropius 128. 129 

brachypopterus 12, 17-19. 129 

moolenburghae 18. 19. 129 
pseudobaganus. Akysis 18. 136, 138-140 
Pseudobagnchthys 111, 123 

macropterus 1 14 
Pscudobagrxis omatus 1 26 
Pseudobelta 171 

pugnax 173 
Pseudogastromyzon 2 

fasciatus 85 

myersi 2, 82 
Pscudogobiopsis 167. 168 

jurongensis 168 

sp. 13 
Pseudogobius 167, 169 

sp. undet. 20. 169 
Pseudogyrinocheilus 81 
Pseudolais 131 

telranema 131. 132 
Pseudomystus 1 16 
Pseudopangasius 131 
pseudopterus. Clupeoides 24 
Pseudosilurus 1 50 
Pseudotylosurus angusticeps 194 
Pscphurus gladius 8 
Psilorhynchus tentaculatus 40 
pugnax. Betla 8. 10, 13, 16, 18-20. 171-173 
pugnax. Macropodus 173 
pugnax. Pseudobetta 173 
pulcher. Diploptenis 178 
pulcher. Luciocephalus 14. 16, 178 
pulcher. Puntius 60 
punctatus. Ophicephalus 169 
punctulatus. Gastromyzon 87 
puntio. Cypnnus 60 
Puntioplites 3. 6. 27. 30. 40. 41. 59 

bulu 16-19. 40. 60 

falcatus 60 

proctozysron 59 

lawak 29 

waandersi 16. 18, 41, 60 
Puntius 3. 21, 38, 58,60-63, 65 

alius 62 

anchisporus 14, 16, 18, 19, 60 

binotatus 13. 14. 17. 60. 61 

binotatus banksi 61 

bramoides 14. 16. 18. 61 

bulu 60 

collmgwoodi 17. 62 



elongatus 61 

endecanalis 18. 19. 38. 62-64 

cugrammis 16 

eugrammus 16. 17, 64. 65 

everetti 64 

fasciatus 65 

foerschi 63, 64 

gonionotus 62 

hemicterus 61 

hexazona 65 

lalcristnga 13, 17, 65 

lawak 40, 41, 60 

lineatus 18, 64. 65 

nini 60 

pentazona 13. 16. 63-65 

penlazona johorensis 65 

pentazona pentazona 65 

pulcher 60 

proctozysron 59 

rhomboocellatus 13. 16, 18, 63-65 

schwancfeldi 66 

schwanenfeldii 60. 62, 66 

sealei 6 1 

sibukensis 61 

streeten 8 1 

sumatranus 8 

letrazona 60 

(BarboidesI bramoides 62 

(Barbodes) fasciatus 64 

(Barbodes) schwanenfeldi 66 

(Barbodes) tetrazona 65 

(Puntius) waandersi 60 
pusillus, Dermogenys 154, 156 
cf pusillus, Dermogenys 154 
pustulosis, Gynnocheilus 17, 81, 82 

quadnfasciatus. Chaetodon 164, 165 
quadrifasciatus, Datnioides 164, 165 
quaterdecimfilis. Polynemus 166 
quinquemaculatus. Barbus 61 

Raiamas 30 
Raja uamak 22 

Rasbora 2. 21. 22. 44. 57. 59. 66. 67. 69. 72. 73. 
75-77 

agilis 13, 14, 16, 17. 19. 20. 66-68 

argyrotaenia 8. 16. 19, 20, 66-68, 72, 74 

axelrodi 2. 1 1 , 1 3. 2 1 . 44. 66-68, 70 

bankanensis 11, 14, 16-20. 66. 68, 70, 71, 74 

beauforti 10, 66, 71 

borapctcnsis 68 

borneensis 16, 18-20, 68, 71. 72 

bnttani 14. 18, 19, 66, 68. 72 

caudimaculata 13. 68. 72, 74, 76 

cavern 68 

cephalotaenia 13, 14, 66, 68. 72 

cf ennealepis 17, 68 

cf myersi 74 

daniconius 66, 68 

dorsimaculata 72 

dorsiocellata 16. 18-20. 68. 73 

dusonensis 13. 16. 20. 66, 68. 73, 74 

eintho\eni 13, 16 

einthovenii 68, 73 

elanga 68 

elegans 17, 68. 73. 74 

ennealepis 17. 66, 68, 73, 74 

heteromorpha 68 

hubbsi 68 

jacobsoni 68 

kalochroma 17, 68. 74 

latenstnata 73 

leptosoma 66, 68, 72, 77 



maculata 69 

meinkeni 69 

myersi 13, 16-18, 66, 69, 74, 75 

pauciperforata 13. 14. 16-18, 66, 69, 74, 77 

phihppina 69 

rasbora 69 

reticulata 69 

rutteni 69, 75 

sarawakensis 13, 16, 17, 69. 75, 76 

semilineata 69 

sp. undet. 16-18, 20. 77 

steineri 69 

stigmatura 76 

subtilis 18. 59.66. 69. 75. 76 

sumatrana 16. 18. 76. 77 

taeniata 66. 67 

tawarensis 69 

tornieri 72 

irilineata 14. 16, 18, 19. 66, 69. 75-77 

tubbi 69 

urophthalma 69 

vateriflons 69 

volzi 17. 66. 69. 74. 77 

volzi-fasciata 77 

volzi var. fasciata 77 
rasbora. Cypnnus 66 
rasbora, Rasbora 69 
Rasborichthys 77 

hclfnchii 18, 19, 78 
regani, Leiocassis 1 17 
remotus, Chonerhinos 184 
rcpasson, Barbus 37 

repasson, Cyclocheilichthys 17-19, 33. 35. 37 
rerio. Brachydanio 70 
reticulata. Rasbora 69 
revcrsa. Botia 17. 96, 101-103 
rhodotaenia. Ophicephalus 170 
rhomboocellatus. Puntius 13. 16. 18. 63-65 
Rhynchobdella 178 

maculata 180 
ridens. Gastromyzon 17. 84. 85. 87. 88 
rios. Pangasius 132 
ripleyi. Homalopterula 88 
robustus. Ambassis 162 
robustus, Leiocassis 116 
robustus, Leiocassis (Pseudomystus) 116 
Rohita 

borneensis 47 

brachynotoptcrus 52 

chrysophekadion 45 

cyanomelas 45 

enneaporos 47 

hasselti 48 

kahajanensis 49 

kappenii 49 

kilogcneion 45 

kuhli 48 

melanopleura 45. 50 

microcephalus 52 

oligolepis 53 

polyporos 45 

rostellatus 48 

schlegeli 52 

iriporos 54 

wandersii 54 

viltata 52 
rohitoides. Labeo 45 
rohitoides, Labeo (Diplocheilos) 52 
rohitoides. Lobochilus 52 
Rohtee 

microlepis 78 

(Osteobrama) vigorsi 78 

vigorsi 78 



206 



CALIFORNIA ACADEMY OF SCIENCES 



Rohlcichthys 78 

microlcpis 18, 11. 78. 7Q 
rosea. Dangila ."(8 
rostellatus, Rohita 48 
rubra, Barbus schwanefeldi \ ar, 66 
rubnpinna. Filirasbora 41 
rubnpinnis, Cydochcilichthys 35 
rugosus, .Acrochordonichlhss 137 
rupicola, Chopraia 88 
rultcni. Rasbora 69, 75 
R\nihobdclla 178 



sabanus. Ompok 18. 151. 152 
sadanundio. Ciobius 16'* 
Sagittabanlius 30 
salmolucius. Banlius 30 
sandakancnsis, Lepidocephalichlhys 96 
sandakanus, Acanlophthalmus 95 
sandakanus, Lcpidoccphalichthys 103 
Sandclia 171 

sanguineus, Silunchlhys 16. 151. 153. 154 
sanhocnsis. Sinogastromyzon 85 
saravacensis. Nemacheilus 14. 97. 107. 108 
sarawakensis. Rasbora 13. 16. 17, 69, 75. 76 
sarawakcnsis, Osteochilus 46 
satyrus. Osphromenus 177 
scandens. Anabas 171 
scandens, Perca 1 7 1 
scapularis, Osleochilus 47 
Scatophagus argus 6 
Scenedcsmus 59 
schacra. Cypnnus (Banlius) 30 
schagra, Pachystomus 30 
schilbeides. Cryploplerus 16. 150 
schilbeides. Hemisilurus 150 
schilbeides. Kryptopterus 18. 146. 150 
Scliisnialorhynchos 79 
helcrorhynchos 17. 79 
heterorhynchus 79 
lobocheiloides 79 
nukia 79 
schlcgch, Osleochilus 16. 19. 46. 47. 53 
schlegcli. Rohila 52 
schlegeli. Toniisloma 6 
Schwanefeldi. Barbus 66 

schwanefeldi. Punlius 66 

Schwanefeldi. Punlius (Barbodcs) 66 

Schwanefeldi. Syslomus (Barbodes) 66 

Schwanenlcldii. Barbus 65 

schwanenfeldii. Punlius 60. 62. 66 

Sciaena jaculator 165 

sclcronema, Ceraioglanis 16. 144 

scleronema. Hemisilurus 144. 145 

Scleropages 23 

formosus6. 16. 19. 22-24 
leichardti 23 

sealei. Barbodes 61 

sealei. Puntius 61 

selangoncus. Nemacheilus 14. 16-20.97. 107. 109 

sclalanensis. Sphaerichthys 175. 176 

sclalancnsis. Sphaerichthys osphromenoides I 76 

scmicinclus, Acanlhophthalmus 18, 98 

semicinctus, Acantophthalmus 16. 19. 20. 96. 98 

Scmilabeo 81 

semilacvis. Cynoglossus 184 

seniilineata. Rasbora 69 

sepal. Trichopus 177 

scphen. Hypolophus 22 

Serrasalmus 1 1 

serpentinus. Ophiccphalus 170 

setigcrum. Luciosoma 16. 17, 43. 44 

seugerus. Barhus 42. 43 



Selipinna 26 
breviceps 25 
mcgalura 26 

melanochir 18. 20. 25. 26 
Shacra 30 

shacra, Cypnnus (Banlius) 30 
shelfordi. Acanlhophthalmus 13, 14. 16-18.98 
shelfordi, Acantophthalmus 16, 18. 96, 98 
Siaja 33 

siaja, Capoeta 33, 35 
siaja. Cyclocheilichthys (Siaja) 35 
siamensis, Epal/eorhynchus 32 
siamensis. Leiocassis 1 16 
siamensis, Leiocassis (Pseudomystus) 1 16 
siamensis, Macrognalhus 180 
siamensis, Osphromenus 177 
sibukensis, Punlius 61 
sicboldu, Bagrus 122 
signata, Belontia 1 7 1 
signifer, Himantura 2, II, 16, 20, 23 
Silonia 128 

Silurichthys 142, 151, 153 
hasselti 13, 17, 151 
hasseltii 151, 153 
phaiosoma 14, 19. 151, 153, 154 
sanguineus 16, 151, 153, 154 
siluroides, Ompok 150 
Silurodes 150 
eugeneiatus I 50 
hypophthalmus 150 
macronema 1 50 
Silurus 143, 144 
anguillaris 127 
apogon 145 
attu 151 
batrachus 127 
bicirrhis 145, 147 
bimaculatus 150 
(Callichrous) pabda 150 
cahanus 126 
cryptopterus 145, 147 
docmak 1 20 
hypophthalmus 150 
lais 147 
Icptonema 145 
limpok 147 
macronema 150 
niicronemus 145, 149 
micropogon 145 
mullcri 151 

palemhangensis 145-147 
phaiosoma I 5 1 
phalacronotus 149 
Mttatus 120 
silus, Chonerhinos 184 
simulans, Tetrodon 188 
sinensis, Tachysurus 1 10 
singanngan, Bagrus 122 
Sinogastromyzon 

sanhocnsis 85 
Sinohomaloptera 
holTmanni 85 
\aoiancnsis 85 
smaragdinus, Thryssocypns 2, 18, 79. 90 
snyderi. Macrochinchthys 45 
sophore. Cypnnus 60 
Sosia 137 

chamaeleon 137 
sp. indcl.. Anus 126 
sp. undct.. Achiroides 16. 183 
sp undct . Calamiana 10. II. 16. 18. 19. 167, 168 
sp undct. Crossochilus 19 
sp undct.. Cyclocheilichthys 18 



sp. undet.. Hemirhamphodon 16. 155. 158 
sp. undct.. Hyperoaslromy/on 84 
sp. undet.. Kryptopterus 17. 150 
sp. undet . Leiocassis 1 19. 120 
sp undet,, Lobocheilus 16, 19 
sp undct., Mugilogobius 13, 16, 168 
sp, undet , Pangasius 131 
sp, undct., Pseudogobius 20, 169 
sp. undet,, Pseudogobiopsis 13 
sp undet,, Rasbora 16-18, 20, 77 
sp, undct,, Stigmatogobius 13, 169 
sp. undet,, Sundasalanx 27 
spectrum, Lepidocephalus 1,19, 94, 96, 106 
Spcrata 120 

Sphaerichthys 10. 175. 176 
acrostoma 175-177 
osphromenoides 13, 175, 176 
osphromenoides selatancnsis 1 76 
selatenensis 175, 176 
vaillanti 9, 16, 18, 19, 175-177 
spilopleura, Luciosoma 16-19. 42-44 
spilurus, Dangila 53 

spilurus, Osteochilus 13, 14, 16, 18-20.46.47.5: 
spinosus. Doryichthys 159 
spiritalis. Ophicephalis 170 
splcndens. Bella 1 7 1 
Slaurastrum 59 
stcinen. Rasbora 69 
stenomus. Bagrus 1 16 
sienomus. Leiocassis 1 16, 120 
stephcnsoni, Homaloplcra 17, 18, 84, 90 
cf stephcnsoni, Homaloplcra 85, 90, 91 
slevensu, Ophiccphalus 170 
stngatus, Barbus 62 
Stigmatogobius 167, 169 
brocki 13, 169 
sp undel. 13. 169 
siigmalura, Rasbora 76 
stirhngi. Anus 1 10 
cf stirlingi. Anus 1 10 
slormi. Hcmianus 
slormii, Cephalocassis 1 10 
slormii. Anus 16, 21. 110. Ill 
slrecten, Punlius 81 
stnala, Channa 8, 13, 19, 20, 170 
slnatus, Glyptoslemon 1 34 
stnalus, Ophiccphalus 8, 170 
slnatus, Osphromenus 1 7 I 
stngatus, Barbus 62 
studeri, Ophiocephalus 170 
sublihs, Rasbora 18, 59, 66, 69, 75. 76 
sumatrana. Rasbora 16. 18. 76. 77 
sumalranus. Barbichthys laevis var. 30 
sumatranus. Barbus 60 
sumalranus. Barbus laevis var, 30 
sumalranus. Barbus (Puntius) 60 
sumalranus. Puntius 8 
sundaicus. Bagrus 1 10 
Sundasalanx 2. 9. 27 
microps IS, 27 
minor 2 
praecov 27 
superhus. .Acantophthalmus 16, 20, 96, 98 
surakartensis. Phihpnoides 169 
Symphysodon 13 
Synaplura 
achira 183 
melanorhyncha 183 
Syncrossus 101 

bcrdmorei 101 
Syngnathus 
boaja 158, 159 
deokhaloides 158, 159 



ROBERTS -WESTERN BORNEO FISHES 



207 



hcterosonia 1 59 
jullieni 159 
martensii 8. 1 59 
/onalus 159 
Systomus 
albuloides 27, 28 
apogon 35 
apogonoides 35 
bclinka 60 
bulu 60 
fasciatus 64 
janthochir 36 
lawak 40 
microlepis 78 
(Barbodes) bolinka 60 
(Barbodes) fasiatus 64 
(Barbodes) goniosoma 61 
(Barbodes) lateristriga 65 
(Barbodes) maculatus 61 
(Barbodes) schwanefeldi 66 
(Puntius) bulu 60 
waandersi 60 



Tachysurus ! 10 

sinensis 1 10 
taenia, Cobilis 95 
taeniata, Betta 13, 14, 172-174 
taeniata, Dangila 38 
taeniata, Garra 40 
taeniata. Rasbora 66, 67 
tambra, Barbus 80 
tambra, Labeobarbus 81 
tambra. Tor 18 
tamhroides, Labeobarbus 8 1 
tambroides. Tor 17, 81. 82 
tapeinopterus, Encheloclanas 128 
tapeinopterus, Heterobranchus 128 
tapiensis, Cyclocheilichthys 35 
tawarensis, Rasbora 69 
teijsmanni, Clanas 128 
Telara 26 
telara, Clupea 26 

tcmminckii, Helostoma 6, 16, 18, 19, 177, 178 
temminkii, Helostoma 178 
tentaculatus, Psilorhynchus 40 
testudineus, Anabas 171 
testudineus, Anthias 171 
Tetraedron 58, 59 
tetranema, Pseudolais 131, 132 
Tetraodon 185-186 

cambodgiensis 188 

fluvlatilis 6 

leiurus 16, 186, 188 

lincalus 188 

(Arothron) modestus 187 

naritus 186 

nigrovindis 186. 188 

palcmbangensis 8. 16. 20. 186, 188 

pinguis 188 
tetra/ona, Barbus 64. 65 
tetra/ona. Punlius 60 
tctra/ona, Puntius (Barbodes) 65 
Tetrodon 11, 188 

bergii 188 

hilgendorfii 188 

simulans 188 
teysmanni, Clanas 127, 128 
Theraponidae 
Thryssocypns 2, 79 

smaragdmus 2, 18, 27, 79, 80 
Thynmchthys 77, 80 

polylepis 16-19, 80, 81 



thynnoides 8, 18, 80 

vaiUanti 80 
thynnoides. Leuciscus 80 
thynnoides. Thynmchthys 8. 18, 80 
liong, Glyptothorax 134, 135 
Tomistoma schlegeli 6 
Tor 3, 80 

ct'. tambra 80 

tambra 18 

tambroides 17, 81. 82 
tornieri. Rasbora 79 
Toxotes 165 

jaculator 165 

microlepis 8. 16. 18. 19. 165 
Trachycorystes 12 
Trachycorystes galeatus 1 2 
Tnchidion multifilis 166 
Trichogaster 177 

leerii 16-20. 177 

trichopterus 16. 177 
Tnchopodus 

Iceni 177 
Trichopsis vittata 171 
trichopterus. Lahrus 177 
trichopterus. Tnchogaster 16. 177 
Trichopus 

leerii 177 

sepal 1 77 
tnfasciata. Betta 171 

trilineata. Rasbora 14. 16. 18. 19. 66. 69, 75-77 
trinema, Leuciscus 42, 44 
irincma, Luciosoma 8, 16-19, 43, 44 
Trinematichthys 42 
tnporos, Osteochilus 16, 17, 19, 54 
triporos, Rohita 54 
triporus. Osteochilus 46^9. 54 
truncatus. Amblyrhynchichthys 6. 16. 18-20. 29 
truncatus. Barbus 28 
tubbi. Rasbora 69 
twecdici. Homaloptera 14. 16. 18. 19. 84. 85. 91, 

94 
Iweediei, Osteochilus hassclti 48 
twecdiei, Wallagonia 152 
Tylognathus 

bo 41 

heterorhynchus 79 

hispidus 42 
typus, Micronema 145, 149 

uarnak. Raja 22 

undulatus, Maslacembelus 178, 179, 181, 182 
umcolor, Maslacembelus 17. 178-180, 182 
unimaculata, Betta 172 
ummaculatus, Parophiocephalus 171 
uranoscopus, Leuciscus 44, 45 
uranoscopus, Macrochirichthys 45 
urophthalma. Rasbora 69 
urophthalmoides, Eleolns 167 
urophihalmoides, Oxyeleotns 20, 167 
urophthalmus. Elcotns 167 
urophthalmus. Ophicephalus 170 
urophthalmus. Oxyeleotns 167 

vagus. Ophiocephalus 8 
Vaillantella 94. 109 

euepiptera 16. 17-19, 97, 109 

cuepiplerus 120 

flavofasciata 109, 110 

maassi 17, 19, 97, 109 
vaillanti, Leiocassis 119, 120 
vaillanti, Maslacembelus 179, 181 
vaillanti, Sphacnchthys 9, 16, 18, 19. 175-177 
vaillanti, Th\nnichthys 80 



Vaimosa 

balteata 169 

brocki 169 

jurongensis 168 
valenciennesii, Barbus 35 
Vanncmania 83 
variabilis, Discognathus 40 
vanegatus, Akysis 136, 142 
vanegatus, Pimelodus 138 
vanus, Acrochordomchthys 138 
vaterifloris, Rasbora 69 
vcrmicularis, Acanthophthalmus 95 
verrucosa, Parakysis 141-143 
vigorsi, Osteobrama 78 
vigorsi, Rohtee 78 
vittata, Rohita 52 
vittatoides, Osteochilus 47 
vittatus, Crossocheilus 57 
vittatus, Osteochilus 8, 47, 52 
vittatus, Paracrossocheilus 13, 17, 57 
vittatus, Silurus 120 
vittata, Tnchopsis 171 
volzi, Rasbora 17, 66, 69, 74, 77 
vobi-fasciata, Rasbora 77 
volzi var. fasciata. Rasbora 77 

waandersi, Arelia 185 

waandersi. Barbus 60 

waandersi. Cynoglossus 18. 19. 184. 186 

waandersi, Osteochilus 18, 19, 46, 47 

waandersi, Plagusia 186 

waandersi, Puntioplites 16, 18, 41, 60 

waandersi, Puntius 60 

waandersi, Puntius (Puntius) 60 

waandersi. Systomus 60 

waandersii. Osteochilus 13. 16. 54. 55 

waandersii, Rohita 54 

Wallago 2, 151, 152 

dinema 144 

heterorhynchus 145 

leeni 3, 8, 16, 151, 152 

nebulosus 1 52 
Wallagonia 151 

twcediei 1 52 
wassinku, Homaloptera 88 
weben, Callichrous 151 
weberi, Ompok 10, 151 
whitcheadi, Protomyzon 85 
wcbcni, Luciosoma 43 
whitcheadi, Protomyzon 85 
Whitleyia 161 
Whitleyina 161 
wolffi, Parambassis 164 
wolffii, Acanthoperca 164 
woUTii, Ambassis 161, 162 
wolffii. Bagrus 126 
wolffii, Chanda 16, 19, 20, 164 
wolffii, Hypselobagrus 126 
wolffii, Macrones 126 
wolffii, Mystus 126 
wolffii, Parambassis 159, 162-164 
wycki, Mystus 126 
wyckii, Bagrus 126 
wyckii, Hemibagrus 126 
wyckii, Macrones 126 
wyckii. Mystus 17, 19, 126 

xanthomelas, Brachygobius 14, 16, 19, 167, 168 
Xcnentodon 152, 153, 155 

cancila 153, 155 

canciloides 14, 17-20, 153, 155 

sp. 153 



208 



CALIFORNIA ACADEMY OF SCIENCES 



Xcnomystus 24 
Xenopterus modeslus 8 

yaolancnsis, Sinohomaloptera 85 



yarrclli. Baganus 16, 18. 133, 134, 140 
yarrclli, Bagrus 133 

zambczcnsis, Leuciscus 30 



zollingcn, Homaloptera 17, 84, 85, 91, 92 
zonatus, Syngnalhus 1 S'' 



General Index 



aerial olfaction 177 
acl quahben 4 
Agcneiosidae 1 2 
Ahnclt, H, 8, 176 
airbreathing 

anabanloids 177 

catfishcs 127 
Akysidac 11. 136 
alcian-alizarin technique 2 
Algemeene Ronsl-en Lelter-Bode 5 
Ambassidac 159 
Amblycipitidae 12 
Amhoina 4 

American Museum of Natural History 
Amphiliidae 12 
Anahantidae 170 
Anabanloidea 20 
anchovies 25 
Andresen, A. J. 7 
angler cat fishes 143 
ants, as fish food 158 
archer fishes 165 
Ariidae 11, 110 
Atlas Ichthyologiquc des Indes 

Neerlandaises 7 
Aucheniptcridae 12 
axial gland, calfishes, 1 30 
Bacillanophyceac 59 
Badidae 165 
Bagndae 1 1 1 
Balon, E K, 9 
Baram River 9 
Bariliinac 44 
Barito River 6 

Batang Lupar 6 

Bcaufon, L. F, de 7 

Belonidae 1 52 

Belontiidac 171 

Bern Natural Histor> Museum 22 

Bianco. P G. 10 

Blceker. P. 6, 7 
Atlas 7 

autobiography 7 
bibliography 7 
type specimens 7 

blind fishes 

Lcpidotcphcihn 106 

blowgun fishes 165 

Boeseman, M, 4, 7 

Boic, F. 6 

Boie. H, 6 

bent acl 4 

bony tongues 23 

Botiinae 94 

Boulenger, G. A, 9 

breeding tubercles 
. Icaninpsis 99 
Rarhmai 100 



Ikinliiis 3 1 
Brachydanio 70 
Cri^ssocheilus 32 
Liiciosoma 44 
Rashom 69-70, 75 
Schizmalorhynchos 79 
Breitenstein, H. 8 
autobiography 8 
collections 8 
British Museum of Natural History 2, 9 
Brooke, J. 9 
bubble nesting 1 7 1 

Bulletin des Sciences Naturelles et de Geologic 5 
bumblebee gobies 167 
Buttikofcr. J, 9 

California Academy of Sciences 2 
carcinophagy 1 10 
carps 27 

catch-all genera 21 
catfishcs 110-152 
Chacidae 143 
Chandidae 169 
Channidae 169 
Chaper, M. 9, 29 
Clanidae 127 
climbing perches 171 
Clupeidae 24 
coal beds 6 
Cohitidae 94 
color change, ontogenetic 
Channa 169-170 
MasJaceinhchis 1 8 1 
PuiUius 64 
color pattern reversal 
Bona 130 
Mastacembelus 181 
Compagno. L, J, V, 2 
continuous reproduction 10, 1 I 
Cranoglanidae 12 
Cuvier, G. 5 

cutaneous papillae, cephalic 
Cyclochcilichlhys 38 
Eirmolus 38 
Cynoglossidae 185 
Cyprinidae 27 
Danioinac 70 
Dasyatidae 22 
Datnioididae 164 
Datnioidinae 164 
De Histona Piscium 4 
designation of type species 3, 21 
diadromy 1 1 
Doeson River 6 
Duncker, G 9 

Dutch Expeditions to Central Borneo 9 
egg eating 1 3 
egg size 

Calaintana 168 



Chonerhmos I 1 
Hemirhamphodon 1 57-158 
Pscudogobiopsis 1 69 

Rashnra 69 
Einthoven, J. 7 
elasmobranchs 22 
Eleotridae 166 
Engraulidae 25 
equatorial river systems 1 
Everett, A, 9 
evolutionary center 1 
Fallours. S. 4 
featherfins 24 
fecundity 69. 168 
Feldman, M. A, 7 
Ferrusac's Bulletin 5 
Field Museum of Natural History 2 
filter feeding 177 
fin-eating 187-188 
fire eel 1 8 1 
Fische Syriens 6 

fish communities, species composition 2 
Fishes of the Indo-Australian Archipelago 7 
Fly River 1 1 
Fowler. H. W. 9 
Freihofer. W. C. 20 
Gastromyzontinae 82-83 
generic typification 21 

Mycins 1 20 
genital papilla 

catfishcs 1 2 

Hemirhamphodon 1 54 

MaslaLemheltis 1 8 1 
giant gouramy 177 
glass perches 1 59 
gobies 167 
Gobiidae 167 
Goulding, M. 3 
gouramies 177-178 
Grabowski. F. 8 
Gunlher. A. 9 
Gynnocheilidae 81 
Hacker. R. 8 
halfbeaks 1 53 
Hanilsch. R. 9 
Hardenberg. J. D. F. 10 

collections 10 

species inquirendae 10 

type specimens 10 
Hasselt, J. C. van 5. 6 
Heckel. J. .1, 6 
Hclfrich. C. 7 
Helostomatidae 177 
Hemiramphidae 152 
herrings 22 
higher classification 

Ak\sidae 21. 136 

anabantoids 20 



ROBERTS-WESTERN BORNEO FISHES 



209 



catfishes 111, 126. 136. 141-142 

Channidae 20 

Datnioididae 21, 164 

Elloposloma 94, 103 

Luciocephalidae 20 

Liiciosoma 44 

Olyridae 126 

Parakysis 141-142 

Peclenocypris 59 

Peleteohagrus 1 1 1 

Rashora 67-70 
hillslream loaches 81 
Histoire Naturclle des Poissons 5 
Holthuis. L. 4, 7 
Homaloptendae 81 
Homaloptennae 82 
Hose. C. 9 

hydrographic history' 1 
Ice Age maxima 1 
Ictalundae 4 
ikan gahus 4 
ikan pachil 3 
ikan sumpit 165 
ikan tapah 3 
Ingcr. R. F. 9 

Institut Royal des Sciences Naturelles '. 
intraspecific vanation 

Acrochordomchlhys 1 37-138 

Anabas 171 

Breitenslenna 141 

Kryplopterus 1 46- 1 47 

Leiocassis 1 1 7 

Mastacemhelus 178-179 

Mystiis 120-124 

Puntnis 6 1 
introductions 

Oryzias 1 

Oiphwnemus 177 
kabos 4 

Kappen, E. J. F, van 7 
Kapuas River 1. 2 
Kasetsart Museum of Fisheries 2 
Kinabatangan River 9 
kissing gouramy 177 
Kuhl, H, 5 
Kukenthal. W. 9 
lakes distnct. Kapuas 6 
leaf fishes 164 
lectoype designation 
lepidophagy 161 
Liem. K. F. 20 
loaches. 94 
lobate testes 12 
Lobotidae 164 
Lorentz. H. A. 9 
Luciocephalidae 178 
Maatshappij ter Bervorderung 9 
Macklot, H. C. 6 
Mahakam River 9 
Malay Peninsula 1 
Martens. E. von 8 
Mastacemhelidae 178 
Mauntius 177 
Melawi River 6 
migrations 

Cosmochilus 3 

Wallagoi. 122 
minnows 27 
Moara Teweh 6, 8 
Mochokidae 12 
Molengraaff, G. 9 

mosquito larvae as fish food 1. 167 
Mount Kinabalu 6 



Mount Raja 6 
Muller. S. 6 

collections 6 

travels 6 
Museum National d'Histoire Naturelle 2, 9 
Museum of Comparative Zoology 2. 3 
Museum Zoologicum Bogorense 2, 3 
myrmecophagy 158 
Nandidac 164, 165 
National Inland Fishenes Institute 2 
Naturhistorisches Museum Wien 6, 8 
Natuurkundig Commissie 5. 6 
needlefishes 152 
neegen-oogen 4 
Nemacheilinae 94 
neotype designation 

Chonerhinos modestus 8 

Afysnis micracanihus 1 20 
Nieuhof .1. 4 
Nieuwcnhuis, A. W. 9 
nomina nuda 5 

non-seasonal reproduction 156 
noodlcfishcs 27 
Notoplendae 24 
Olyndac 12. 124 
oophagy 13 
oral brooding 10. 110. 159 

Ariidae 110 

Bclontiidae 171. 176 

Chandidae 11. 159. 162 

Luciocephalidae 11. 178 

Osteoglossidae 1 1 
Osphronemidae 177 
Ostanophysi I 
Osteoglossidae 23 
ostracophilous larvae 2 
oviducai fertilization 12 
ovipositor 

Masuicembelus 181 
ovovivipanty 10 

llcmirhamphodon 1 56-1 58 
paddlefish 8 
Pepke. H. J. 8 
Pangasiidae 130 
Parakysidae 141 

parental care (see also oral brooding) 12-13 
pectoral gland 1 30 
pectoral pore 1 30 
penne visch 4 

periodicity, reproductive 10-12 
Peters, W. C, 8 
Pleiffer, I. 6 
Phalacronoti 145 
Phallostcthidae 9 
phyloplankton 58-59 
pike gouramy 11, 178 
pipefishes 158 
planl-eating 3, 58-59 
Pleistocene sea levels 1, 186 
Plotosidae 12 
Polynemidae 166 
Popta, C. M. L. 9 

Preussische Ost-Asien Expedition 8 
pnapium 9 

priman. freshwater fishes 1 1 
Pristolepidae 165 
protoanabantoids 165 
protrusible jaws 

LucHKcphalus 20 
pulTerfishes 186 
Pyrophyta 59 
radiography 22, 148 
raritv 1 1 



Regan. C. T, 9 

Renard, L. 4 

reproduction 10 

reproductive guilds 10 

reproductive periodicity 10-12 

reproductive seasonality II, 156 

Rhodcinae 2 

nee fish 1 

Rijksmuseum van Natuurlijke Historic 2, 6, 7, 9 

Rofen R. R 7 

Salangidae 2 

Salmomformes 2 

scale eating 

Chonerhonos 187-188 

Paradoxodacna 1 6 1 
Schilbeidae 128 
Schwaner, C. 6 
Schwaner Mountains 6 
sea catfishes 1 10 
sea levels 1 

secondary freshwater fishes 1 1 
Senckenberg Museum 9 
sexual dichromatism 

Rashora axelrodi 67-70 
sexual dimorphism 

Barbiiaa 100 

Cobitidae 100, 105 

Cypnnidae 

Gobiidae 168 

Gymnochanda 159-160 

Hemiramphidae 157 

Mastaccmbelidae 181 
Sigal, D. 7 
Silundae 143 
Sisoridae 133 
sleepers 166 
Smith, H. M. 5 
Smithsonian Institution 2, 3 
snakeheads 169 
Soleidae 183 
soles 183 

South China Sea I 
spawning strategies 1 2 
species inquirendae 

Rashora beaufortt 1 1 

Sphaerichlhys osphromcnoidcs 176 
spermatophores 12 
spermalozeugma 12 
spiny eels 1 78 
Steindachner, F. 8 
Stevens, M, J, N, 7 
stingrays 22 
Sunda River 1 
Sundaland I 
Sundasalangidae 27 
Synbranchidae 182 
synchronous egg clutches 11-12 
Syngnalhidae I 58 
synonymies 21 
Temminck, C. J. 5 
testes morphology, catfishes 1 2 
Tetraodontidae 186 
Teweh River 8 
Thepass, A, H, 7 
Theraponidae 164 
thrcadfins 166 

Tokyo LIniversity of Agnculturc 10 
tongue soles 185 
Toxotidae 165 
transparency 

Gymnochanda 1 60 

Kryplopterus 147, 149 

Sundasalanx 27 



210 



CALIFORNIA ACADEMY' OF SCIENCES 



Ivpc-spccies designation 21 

unculi 2 

unculiferous epidermal plaques 
Akysidae 136 
Sisoridae 133. 136 

unculiferous larvae 2 

unculiferous menial disc 
(ianu 40 

unculiferous lips 
Cnlochciliclilhys 36 
Ostimiulus 4S-5\. 53-55 
Pseudogaslwmv-on 2 

unculiferous paired fins 



Cohitidae "^Q 

C>prinidac 30 

Homaloplendae 82 
unculiferous thoracic organ 

(ilvploihorax 132 
Universita degli Studi dell'Aquila 10 
LIniversiteit Alexander von Humboldt 8 
Vaillant, L <) 
Vaillantellinac <)4 
Valenciennes, A. 5 
Valcntijn. F. 4 
Valenlini, M, 4 
Valcntim's Naturkamer 4 



vijfoogen 4 

Vlaming, C. de 4 

vyj oogh 4 

Weber. M. 7 

uhiptailcd stingrays 22 

Willughby. F. 4 

wit visch 4 

Wolff. J. 7 

Yangtze paddlefish 8 

Zoological Museum of Amsterdam 3. 10 

Zoologischcs Museum Hamburg 3 



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