Journal of Fish Biology (1998) 53, 645–657

Article No. jb980731

A revision of the akysid catfish genus Breitensteinia

Steindachner with descriptions of two new species
H. H. N*‡  D. J. S†
*Department of Biological Sciences, National University of Singapore, Kent Ridge,
Singapore 119260 and †Department of Zoology, The Natural History Museum,
Cromwell Road, London SW7 5BD, U.K.

(Received 6 February 1998, Accepted 8 May 1998)

The highly specialized akysid catfish genus Breitensteinia is revised; two new species are
described. Breitensteinia cessator is described from Sumatra and western Borneo. It can be
distinguished by its large, wide-set eyes and evenly scattered brown spots on the dorsal and
pectoral fins. Breitensteinia hypselurus is described from western Borneo. It can be distin-
guished by its relatively tall neural spines on caudal vertebrae, smooth posterior edge of dorsal
spine, short caudal peduncle, and fewer vertebrae. Breitensteinia insignis is sexually dimorphic.
 1998 The Fisheries Society of the British Isles

Key words: Siluriformes; Breitensteinia; Sumatra; Borneo; relationships.

INTRODUCTION
The highly specialized and distinctive catfishes of the akysid genus Breitensteinia
Steindachner inhabit the middle and upper reaches of rivers in Borneo and
Sumatra. The genus has been considered monotypic since its description by
Steindachner (1881), based on Breitensteinia insignis Steindachner, 1881 from the
Barito River in southern Borneo. However, very little is known of these strange
catfish because of their extremely poor representation in museum collections.
Roberts (1989) suggested that material identified as B. insignis from Sumatra
and western Borneo might not be conspecific with the holotype of B. insignis.
Until now it has been impossible to follow up Roberts’ suggestion because of the
paucity of material other than the holotype of B. insignis, which is in poor
condition.
During an on-going survey of the Barito River drainage by the second author
and collaborators from the Museum Zoologicum Bogoriense, fresh material of
B. insignis was obtained which has enabled us to redescribe B. insignis and
compare material from the Barito River with material from Sumatra and western
Borneo. The comparison has made it clear that the material from Sumatra and
western Borneo belongs to two new species.

MATERIALS AND METHODS

Measurements were made point-to-point with dial calipers and recorded to 0·1 mm.
Ng & Lim (1995) is followed except: head length (LH) was measured from the tip of the
‡Author to whom correspondence should be addressed: Tel.: 065 874 2969; fax: 065 779 2486; email:
scip7116@leonis.nus.edu.sg
645
0022–1112/98/090645+13 $30.00/0  1998 The Fisheries Society of the British Isles
646 . .   . . 

snout to the posteriormost extremity of the fleshy opercular flap; preanal, prepelvic and
prepectoral lengths were measured from the tip of the snout to the anterior base of the
dorsal, anal, pelvic and pectoral fins, respectively; and interpelvic distance was measured
as the distance between the bases of the last (posteriormost) ray of the left and right pelvic
fins. Measurements are reported as proportions of standard length (LS) except for
subunits of the head which are presented as proportions of LH.
Fin ray counts were obtained under transmitted light using a binocular dissecting
microscope and follow Hubbs & Lagler (1947) with a small modification: unbranched fin
rays are indicated with lower-case Roman numerals. Gill raker counts follow Roberts
(1992). Vertebral counts follow Siebert & Richardson (1997) and were taken from
radiographs. This protocol is more elaborate than usual but recording more vertebral
characters provides insight into the evolution of Breitensteinia, as was also demonstrated
in the cyprinid genus Rasbora by Siebert (1997). The number in parentheses following a
particular fin ray, vertebral, branchiostegal ray, or gill raker count is the number of
examined specimens with that count.
Drawings of specimens were made with a Nikon SMZ-10 fitted with a camera lucida.
Specimens examined for the present study are in the following collections: American
Museum of Natural History, New York (AMNH); California Academy of Sciences, San
Francisco (CAS); Collection of Maurice Kottelat, Cornol (CMK); Museum Zoologicum
Bogoriense, Cibinong (MZB); The Natural History Museum, London (BMNH);
Nationaal Natuurhistorisch Museum, Leiden (RMNH); National Museum of Natural
History, Smithsonian Institution, Washington, D.C. (USNM); Instituut voor
Systematiek en Populatiebiologie, Universiteit van Amsterdam (ZMA); and Zoological
Reference Collection, National University of Singapore (ZRC).
The phylogenetic classification of de Pinna (1996) is followed since it is based on
sound principles and a broad anatomical survey of relevant South-east Asian catfishes.
In this classification Breitensteinia is recognized as a closer relative of Parakysis than of
Akysis.

BREITENSTEINIA STEINDACHNER, 1881

Type species: Breitensteinia insignis Steindachner, 1881, by monotypy.

Gender: Feminine.

Diagnosis
A genus of akysid catfish lacking an adipose fin but having a long low adipose
ridge, with a very long and slender caudal peduncle, with gill opening not
extending above the base of the pectoral spine, with 11–12 principal caudal-fin
rays, and with 42–45 vertebrae.

Discussion
Two features of the diagnosis are especially noteworthy. Species of
Breitensteinia conform to the caudal-fin principal ray pattern of other akysids,
which is unusual among catfishes (Roberts, 1989); i.e. they have a low principal
ray count and more rays in the upper lobe than in the lower.
The elongate form of Breitensteinia is reflected in their high number of
vertebrae [they have eight to 12 more vertebrae than other akysid species
(Roberts, 1989)], which is also a clear indication that the genus is monophyletic.
All Breitensteinia species have certain other features in common. Among these
are: head depressed and broad, covered with small tubercles; anterior nostril
tubular; occipital process narrow, its tip tapering and reaching predorsal plate;
dorsal origin nearer tip of snout than caudal flexure; body with tubercles
arranged in five to six longitudinal rows on each side; dorsal profile rising evenly
    647

but not steeply from the tip of snout to the origin of dorsal fin, then sloping
gently ventrally from there to caudal peduncle; ventral profile horizontal to
origin of anal fin, then rising to caudal peduncle; dorsal origin nearer tip of snout
than caudal flexure; caudal-fin truncate.
Little is known about the biology of Breitensteinia, but they are very lethargic
in aquaria (Ferraris, 1991; pers. obs.). However, that prawns are included in
their diet and that they can be taken with hook and line suggests they might not
be so lethargic in the wild. Ferraris also observed periodic moulting of the skin
in specimens held in aquaria. We document sexual dimorphism in B. insignis
below. The three known species of the genus are so similar we are confident all
members of the genus will be revealed as sexually dimorphic when further
material becomes available.

ARTIFICIAL KEY TO THE SPECIES OF BREITENSTEINIA

1(a) Posterior edge of dorsal spine smooth (see Fig. 4); neural spines on caudal
vertebrae relatively tall (see Fig. 5); length of caudal peduncle 26·0% LS; snout
length 35·5% LH; interorbital distance 30·9% LH; 42 vertebrae (western Borneo)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. hypselurus sp. nov.

1(b) Posterior edge of dorsal spine serrated (see Fig. 4); neural spines on caudal
vertebrae relatively short (see Fig. 5); length of caudal peduncle 30·7–36·8%
LS; snout length 27·2–34·1% LH; interorbital distance 19·2–26·9% LH; 44–45
vertebrae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2(a) Eye diameter 3·9–5·3% LH; interorbital distance 25·4–26·9% LH; brown
spots on dorsal and pectoral fins evenly scattered (Sumatra & western Borneo)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. cessator sp. nov.

2(b) Eye diameter 1·9–3·2% LH; interorbital distance 19·2–25·1% LH; brown
spots on dorsal and pectoral fins concentrated in a dark band near the edges
(southern Borneo). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. insignis

BREITENSTEINIA CESSATOR SP. NOV.

Figures 1–2 and 4–5.

Breitensteinia insignis (nec Steindachner)—Vaillant, 1901: 76–81, figs 16–18;

Weber & de Beaufort, 1913: 374–375, fig. 151 (in part); Kottelat et al., 1993: 75,
pl. 37 (in part).
Breitensteinia cf. insignis—Roberts, 1989: 140–141, fig. 108 (in part).

Holotype
MZB 3693, 186·1 mm LS; Borneo: Kapuas mainstream 58 km NE of Sintang
and 1 km downstream from Sebruang (025·5 N 11152·5 E): T. Roberts et al.,
16 August 1976.

Paratypes
CAS 49415, 2 ex., 88·9–181·2 mm LS and USNM 230304, 1 ex., 190 mm LS;
data as for holotype. CMK 6035, 1 ex., 165 mm LS; Sumatra: Palembang,
648 . .   . . 

F. 1. Breitensteinia cessator. CAS 49415, paratype, 189 mm LS, Borneo: Kapuas Basin

Tulangbawan River; donated by Vivaria, 29 May–4 June 1988. RMNH 7847, 1

ex., 168·2 mm LS; Borneo: Sintang; J. Büttikofer, 1894. ZMA 104.653, 1 ex.,
75·3 mm LS; Sumatra: Jambi; P. H. Moolenburgh, 1909.
Roberts (1989) was not certain that specimens of Breitensteinia from Sumatra
and western Borneo were conspecific with B. insignis (from southern Borneo),
but could not conclude further due to the state of preservation of the holotype
(the only specimen then known) of B. insignis. Subsequent collection of fresh
material of B. insignis has enabled us to validate Roberts’ suspicion.

Diagnosis
Breitensteinia cessator is differentiated from its congeners by the following
unique set of characters: eye diameter 3·9–5·3% LH; interorbital distance
25·4–26·9% LH; snout length 27·2–31·8; length of caudal peduncle 30·7–36·8;
44–45 vertebrae; and brown spots on the dorsal and pectoral fins evenly
scattered. A photograph of the new species is presented in Fig. 1.

Proportional measurements
In % LS: head length 16·5–18·7, head width 12·0–18·2, predorsal distance
27·2–34·3, preanal length 54·7–62·3, prepelvic length 36·5–40·3, prepectoral
length 16·5–18·1, body depth at anus 6·3–7·0, length of caudal peduncle
30·7–36·8, depth of caudal peduncle 1·2–2·0, pectoral-fin length 14·5–15·3, length
of dorsal-fin base 4·2–6·0, pelvic-fin length 8·7–10·7, length of anal-fin base
9·0–10·8, caudal-fin length 8·1–10·5. In % LH: snout length 27·2–31·8,
interorbital distance 25·4–26·9, eye diameter 3·9–5·3, nasal barbel length 5·8–7·5,
maxillary barbel length 41·6–48·7, inner mandibular barbel length 12·8–21·1,
outer mandibular barbel length 33·8–50·4.

Counts
Fin ray counts: dorsal I,4,i (4); pectoral I,6,i (4); pelvic i,4,i (4); anal ii,6 (2),
ii,6,i (1) or ii,7 (1); caudal 6/5 (4). Branchiostegal rays 5 (4). Gill rakers 1+3 (2).
Vertebrae counts are reported in Table I.

Fin spine characters

Dorsal spine with 7 (2), 8 (1) or 9 (1) large serrae posteriorly. Pectoral spine
stout, with 8 (1), 14 (1), 15 (1), or 16 (2) large serrae posteriorly.

Colour pattern
Dorsal surface of head brown, with dark-brown spots scattered randomly
throughout. Dorsolateral and lateral surfaces of body dark brown, with about
11–13 light-brown spots on dorsolateral surface extending from predorsal area to
    649

F. 2. Map showing distribution of Breitensteinia species: B. cessator (); B. hypselurus (); B. insignis
().

base of caudal fin. Adipose ridge light brown. Small patch of light brown on
predorsal area present occasionally. Belly, chest and ventral surface of the head
cream or light brown, with few scattered brown spots. Dorsal and pectoral fins
cream with numerous brown spots scattered throughout. Pelvic and anal
fins cream with one or two dark-brown bands, one present occasionally at the
base of the fins and another in the distal half of fins. Caudal-fin cream, with
dark-brown band in the distal half and with a narrow midlateral prolongation.
Barbels and pectoral spines cream or light brown, with dark-brown spots present
sometimes on dorsal surfaces.

Comparisons
Breitensteinia cessator resembles B. insignis closely, but is distinguished by its
larger and more widely set eyes (eye diameter 3·9–5·3% LH v. 1·9–3·2; interorbital
distance 25·4–26·9% LH v. 19·2–25·1). The brown spots on the dorsal and
pectoral fins in B. cessator are also more evenly scattered (v. concentrated in a
dark band near the edges).

Distribution and ecology notes

Presently known from the Batang Hari and Tulangbawan drainages in
Sumatra and the Kapuas River drainage in western Borneo (Fig. 2). Two of the
specimens examined by Roberts (1989) had small prawns in their stomachs.

Etymology
From the Latin cessator (idle fellow), in allusion to the sluggish nature of the
fish. Used as a noun in apposition.
650 . .   . . 

F. 3. Breitensteinia hypselurus. RMNH 16048, holotype, 121·9 mm LS, Borneo: Kapuas Basin.

BREITENSTEINIA HYPSELURUS SP. NOV.

Figures 2–3 and 4–5.

Breitensteinia insignis (nec Steindachner)—Weber & de Beaufort, 1913: 374–

375 (in part); Kottelat et al., 1993: 75 (in part).
Breitensteinia cf. insignis—Roberts, 1989: 140–141, fig. 108 (in part).

Holotype
RMNH 16048, 1 ex., 121·9 mm LS; Borneo: Sanggau; Westenenk, 1894.

Diagnosis
Breitensteinia hypselurus (Fig. 3) is differentiated from its congeners by its
relatively tall neural spines in the caudal vertebrae (Fig. 5), shorter caudal
peduncle (length of caudal peduncle 26·0% LS v. 30·7–36·8), longer snout (snout
length 35·5% LH v. 27·2–34·1), more wide-set eyes (interorbital distance 30·9%
LH v. 19·2–26·9) fewer vertebrae (42 v. 44–45), and smooth (v. serrated) dorsal
spine (Fig. 4).

Proportional measurements
In % LS: head length 21·5, head width 18·2, predorsal distance 34·3, preanal
length 62·3, prepelvic length 46·5, prepectoral length 22·1, body depth at anus
8·1, length of caudal peduncle 26·0, depth of caudal peduncle 3·2, pectoral-
fin length 18·4, length of dorsal-fin base 5·7, pelvic-fin length 10·2, length of
anal-fin base 11·6, caudal-fin length 13·0. In % LH: snout length 35·5,
interorbital distance 30·9, eye diameter 4·6, nasal barbel length 4·2, maxillary
barbel length 72·1, inner mandibular barbel length 18·3, outer mandibular barbel
length 38·2.

Counts
Fin ray counts: dorsal I,4,i (1); pectoral I,6,i (1); pelvic i,4,i (1); anal ii,7,i (1);
caudal 6/5 (1). Branchiostegal rays 6 (1). Gill rakers 0+3 (1). Vertebrae counts
are reported in Table I.

Fin spine characters

Dorsal spine without serrae on posterior margin. Pectoral spine stout, with 11
(1) large serrae posteriorly.
    651

F. 4. Schematic illustration of the dorsal spines of: (a) B. hypselurus, RMNH 16048, holo-
type, 121·9 mm LS; (b) B. cessator, CAS 49415, paratype, 189 mm LS (dorsal spine of B. insignis
similar).

F. 5. Schematic illustration of the caudal vertebrae of: (a) B. hypselurus, RMNH 16048, holotype,
121·9 mm LS; (b) B. cessator, CAS 49415, paratype, 189 mm LS (caudal vertebrae of B. insignis
similar).
652 . .   . . 

T I. Vertebral counts of Breitensteinia

B. hypselurus B. cessator B. insignis

Holotype n=5 n=5

Total vertebrae 42 44–45 44–45

Prehaemal vertebrae 15 14–17 15–16
Haemal vertebrae 27 28–30 28–29
Peduncular vertebrae 14 15–16 15–16
Anal fin position 21 21–22 20–22

Colour pattern
Dorsal surface of head brown, with very few dark-brown spots scattered
randomly throughout. Dorsolateral and lateral surfaces of body dark-brown,
with about eight light-brown spots on dorsolateral surface extending from
predorsal area to base of caudal-fin. Adipose ridge light brown. Belly, chest and
ventral surface of the head cream or light brown. Dorsal, pectoral, pelvic and
anal-fins cream with a dark-brown band in the distal half of fins. Caudal-fins
cream, with dark-brown band in the distal half and with a narrow midlateral
prolongation. Barbels and pectoral spines cream or light brown, with dark-
brown spots.

Comparisons
Breitensteinia hypselurus is distinguished easily from its congeners by its
relatively tall neural spines on the peduncular vertebrae (Fig. 5), shorter caudal
peduncle (26·0 v. 30·7–36·8% LS), longer snout (snout length 35·5% LH v.
27·2–34·1), more widely set eyes (interorbital distance 30·9% LH v. 19·2–26·9),
fewer vertebrae (42 v. 44–45), and smooth (v. serrated) dorsal spine (Fig. 4).

Distribution
Known only from the Kapuas River basin in western Borneo (Fig. 2).

Etymology
From the Greek hypselos (high) and oura (tail), in allusion to the relatively tall
neural spines of the caudal vertebrae. Used as a noun in apposition.

Discussion
Roberts (1989) treated the only known specimen of B. hypselurus (RMNH
16048) as conspecific with B. insignis because at that time it was the only known
male specimen of Breitensteinia. However, he recognized that the differences
between it and the other material he had to hand were of such a nature that they
might not be attributable to sexual dimorphism but to differences between
species. Fresh material of B. insignis has revealed the nature of sexual dimor-
phism in Breitensteinia (see discussion of B. insignis) and shows that sexual
dimorphism in Breitensteinia takes a different form than the differences between
B. hypselurus and other Breitensteinia species. Roberts’ (1989) suspicion that
two species might be involved was correct.
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F. 6. Breitensteinia insignis. ZRC 41964, 137·6 mm LS, Borneo: Barito Basin.

Breitensteinia hypselurus has a shorter caudal peduncle than B. cessator and

B. insignis (see below) and fewer peduncular vertebrae (Table I), which is to be
expected if the position of the anal-fin along the vertebral column is constant as
the total number of vertebrae varies among species. In an evolutionary context
either the number of peduncular vertebrae in B. hypselurus is reduced or that of
B. cessator and B. insignis is increased. We argue below that the latter has
occurred.

BREITENSTEINIA INSIGNIS STEINDACHNER, 1881

Figures 2 and 6–7.

Breitensteinia insignis—Steindachner, 1881a: 213–215, 1881b: Fig. 2; Vaillant,

1901: 76–81, Figs 16–18 (in part); Weber & de Beaufort, 1913: 374–375 (in part);
Kottelat et al., 1993: 75 (in part); de Pinna, 1996, 7.

Material
NMW 55042, holotype, 129·8 mm LS: Borneo: Barito River, Muara Teweh;
H. Breitenstein, 1880. AMNH 58378, 2 ex., 110·2–110·7 mm LS; locality and
collector unknown. BMNH 1997.17.7:1, 1 ex., 130·0 mm LS; Borneo:
Kalimantan Tengah, Sungai Barito, market at Puruk Cahu and Sungai Barito at
Muara Laung; D. Siebert, O. Crimmen & A. H. Tjakrawidjaja, 20–22 February
1991. BMNH 1997.17.7:3–5, 3 ex., 117·1–147·6 mm LS; ZRC 41964, 1 ex.,
137·6 mm LS; Borneo: Kalimantan Tengah, Sungai Barito at Muara Laung;
D. Siebert, O. Crimmen & A. H. Tjakrawidjaja, 20–22 February 1991.

Diagnosis
Breitensteinia insignis is differentiated from its congeners by the following
unique set of characters: eye diameter 1·9–3·2% LH; interorbital distance
654 . .   . . 

F. 7. Schematic illustration of the ventral view of the pelvic fins and external genitalia of B. insignis: a.
male (BMNH 1997.7.17:3, 117·1 mm LS) and b. female (ZRC 41964, 137·6 mm LS).

19·2–25·1% LH; snout length 28·6–34·1% LH; length of caudal peduncle 33·7–
36·5% LS; 44–45 vertebrae; and brown spots on the dorsal- and pectoral-fins
concentrated in a dark band in the distal half.

Proportional measurements
In % LS: head length 16·8–18·2, head width 11·1–13·2, predorsal distance
26·7–28·5, preanal length 55·1–56·6, prepelvic length 36·7–38·2, prepectoral
length 13·9–18·9, body depth at anus 5·5–7·1, length of caudal peduncle
33·7–36·5, depth of caudal peduncle 1·3–1·6, pectoral-fin length 13·1–16·0, length
of dorsal-fin base 3·3–5·5, pelvic-fin length 8·1–9·7, length of anal-fin base
9·1–10·1, caudal-fin length 7·2–9·4. In % LH: snout length 28·6–34·1, interorbital
distance 19·2–25·1, eye diameter 1·9–3·2, nasal barbel length 4·0–6·5, maxillary
barbel length 40·6–49·8, inner mandibular barbel length 13·1–18·0, outer man-
dibular barbel length 33·2–43·9.

Counts
Fin ray counts: dorsal I,4 (1) or I,4,i (5); pectoral I,5,i (1) or I,6,i (5); pelvic i,4,i
(2) or i,5 (4); anal i,6,ii (1) or ii,7 (5); caudal 6/5 (5) or 6/6 (1). Branchiostegal
rays 5 (5). Gill rakers 0+4 (1) or 1+4 (1). Vertebrae counts are reported in
Table I.

Fin spine characters

Dorsal spine with 5 (2), 6 (1), 7 (1) or 9 (2) large serrae posteriorly. Pectoral
spine stout, with 12 (1), 13 (3) or 14 (2) large serrae posteriorly.
    655

Colour pattern
Dorsal surface of head brown, with very few dark-brown spots scattered
randomly throughout. Dorsolateral and lateral surfaces of body dark brown,
with about 11–14 light brown spots on dorsolateral surface extending from
predorsal area to base of caudal-fin. Adipose ridge light brown. Small patch of
light brown on predorsal area present occasionally. Belly, chest and ventral
surface of the head cream or light brown, with few scattered brown spots.
Dorsal-, pectoral-, pelvic- and anal-fins cream with a dark-brown band in the
distal half of fins. Caudal-fins cream, with dark-brown band in the distal half
and with a narrow midlateral prolongation. Barbels and pectoral spines cream
or light brown, with dark brown spots present sometimes on dorsal surfaces.

Comparisons
Breitensteinia insignis has a smaller eye than B. cessator and spots on the
dorsal- and pectoral-fins concentrated near the edges rather than scattered over
the whole area of the fins (see above); it has a longer caudal peduncle than B.
hypselurus as well as a serrated rather than smooth dorsal spine (see above).

Distribution and habitat notes

Our specimens of B. insignis represent the first record of fresh material
collected since its original description; at present the only verified locality records
for B. insignis come from the Barito River drainage of southern Borneo (Fig. 2).
Its occurrence in other river basins of southern Borneo would not be surprising
since among the fish fauna of southern Borneo there is more regional endemism
than river basin endemism.
The recent Barito River material was taken at night with hook and line, by
children who were fishing along the mainstream. Day-time fishing in the same
area yielded another akysid, Acrochordonichthys sp.

Discussion
Sexual dimorphism in Breitensteinia is documented here for the first time.
Males have the anus situated immediately in front of a genital papilla, which is
located immediately posterior to the pelvic-fin base (Fig. 7). The genital opening
is situated at the tip of the papilla, covered by a fleshy flap. In females, the anus
is situated more posteriorly and the genital opening is located at the tip of a short
genital appendage (Fig. 7). The pelvic fins of females are also more closely-set
(inter-pelvic distance 2·1–2·6% LS v. 2·6–2·8) and usually overlap posteriorly
(v. laterally spread). This form of dimorphism was first reported by Ng &
Kottelat (1996) in Akysis fuscus among akysids. However, in A. fuscus it is the
males and not the females that have more closely-set pelvic fins which overlap
posteriorly.

RELATIONSHIPS
De Pinna (1996) dealt extensively with the relationships of Breitensteinia
among other South-east Asian catfishes in his phylogenetic analysis of the
families Sisoridae, Akysidae, and Amblycipitidae. We have evidence to extend
those results to a hypothesis of relationships within Breitensteinia. The low
656 . .   . . 

neural spines in the peduncular region of B. insignis and B. cessator are clearly
derived among akysids and are evidence that B. insignis and B. cessator are each
other’s closest relative; the taller neural spines of B. hypselurus are like those
found in species of the sister group (Acrochordonichthys, de Pinna, 1996) to
Breitensteinia. The difference in number of vertebrae among species of
Breitensteinia can be interpreted as additional evidence in support of this
hypothesis. Breitensteinia stands out as derived among akysids with a great
increase in number of vertebrae. Breitensteinia insignis and B. cessator represent
the extreme of this trend, with the largest increase in number of vertebrae.
The trend is evident in number of peduncular vertebrae too. Breitensteinia
insignis and B. cessator have a longer caudal peduncle than B. hypselurus and
this is reflected in an increased number of peduncular vertebrae (Table I).
The phylogenetic relationships of the genus are [B. hypselurus (B. insignis,
B. cessator)].

The authors thank R. Arrindell (AMNH), D. Catania (CAS), M. Kottelat (CMK),

M. van Oijen (RMNH), L. Parenti (USNM), and I. Isbrücker (ZMA) for the loan
of material; E. Mikschi (NMW) for hospitality and help in Vienna; and Tan Heok
Hui and Kelvin Lim for help with the figures. This project has been supported
partially by research grant RP 960314 from the National University of Singapore to
P. K. L. Ng.

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