Please note that all images of preserved specimens are copyright of the institution to which the specimen belongs; for photographs of live specimens the copyright is that of the photographer.

This is an updated and amended version of an article which appeared with the same title in Cat Chat, The Journal of the Catfish Study Group (UK), Volume 5 Issue Number 2, pages 5-17, June 2004.


his article has been written for aquarists and, to a degree, ichthyologists, to contribute towards resolving the identity and validity of the striped bagrid catfishes of the genus Mystus Scopoli, 1777. I say contribute towards as to attempt to finally resolve generic placements and give defining characters for all species is something only an ichthyologist could do as it is a huge task. Jayaram & Sanyal (2003) have attempted this but still finally elected to lump them all under Mystus pending further study (although they accepted in their Addenda that many of the species they covered should be included in Hemibagrus Bleeker, 1862).

Having disregard to the species that belong in Hemibagrus, Mystus is currently a genus containing at least two, possibly more, groups of species that will probably be split into separate genera at some point. Roberts (1994) split the genus into two groups based on several features. The ones that can be used for living fishes were the relative height and length of the adipose fin, length of the maxillary barbels, the shape of the caudal fin, and whether or not the fontanel (a gap/space in the structure of the top of the skull) is split by an epiphyseal bar (basically making it look like one, or two elongated grooves on the top of the head). The split can be a complete split or the bar can appear as an indent that does not fully split the groove.

Roberts restricted Mystus to the species with long and high adipose fins, long maxillary barbells, deeply forked caudal fins (with upper lobe being longer) and with the split fontanel. He did not propose any generic name(s) for the remaining species. Jayaram & Sanyal (2003) felt that more of the species could be included in the sensu stricto group to the ones included by Roberts, and felt that some of his characteristics were unreliable.

There is a generic name that already exists which could be resurrected in the future for some of the remaining species: Aspidobagrus Bleeker, 1862. Recent authors have mentioned Hypselobagrus Bleeker, 1862, but if one follows Roberts (1992) work on the synonymy of its type species, this genus would be a synonym of Mystus. Looking at the morphology of what is currently considered to be the type species of Mystus, I personally doubt how it can be included as belonging to the same morphological group as the species Roberts (1992) lists in sensu stricto. Basically its still a bit of a mess and I think that for the time being they will have to be referred to as Mystus, but I wouldn’t bet against Hypselobagrus and/or Aspidobagrus being resurrected.

With all the species I have tried to get images of the preserved type specimens and/or reproduced the original drawing if one was provided. The reason I have done this is that I have found that unless one refers to the type specimens or drawings of them, one cannot be 100% sure that you are talking about the species in question. I appreciate that for some people images or drawings of dead, sometimes now colourless, fish may not be the best way to assist in identification but misidentifications have been made previously and I feel this is the safest way. However, even with type material to refer to, some of the multi-striped species are still difficult to tell apart when looking at live fish, and sometimes one has to use the morphology of the fontanel (which is not easy in live fish) and supra-occipital process (a bony extension of the skull which extends towards the ‘basal bones’ of the dorsal fin). Sometimes the length, placement and height of the adipose fin in combination with maxillary barbell length and body pattern are a good defining character.

Roberts (1992) separates the ‘shoulder spot’ into two types. In some species it is a spot or a dark area, whereas in some species it is a semi-ocellus as it has a dark or black spot with a pale or bright imperfect ring around it:

Humeral semi-ocellus / spot

Placed immediately above the humeral process, which is a bone on the body immediately after the shoulder girdle in which the pectoral fin spine is inserted. The humeral process projects backwards and sometimes is angled upwards.

Tympanic semi-ocellus / spot

The spot overlays the tympanum, which is a membranous covering of the swim bladder. The tympanic semi-ocellus / spot is usually posteriorly further along the body in comparison with the humeral semi-ocellus / spot, and is usually bigger.

But for some species the only definite differences are that of measurements and proportions (morphometrics) and also counts of gill rakers, vertebrae etc (meristics). Obviously I have not mentioned these differences as they are useless for most users of this article.

I hope this article will go some short way in assisting anyone interested in the identification of this ‘group’ of catfish.

Mystus albolineatus Roberts, 1994 

See holotype CAS 79030 which originated from Prachinburi Market, Bangpakong basin, Thailand, but the species is also present in the lower Mekong basin, Cambodia. Largest type specimen is 13.5 cm SL. See also a paratype (ANSP 16453).

This species has a white or pale stripe running along the lateral line. The stripe is bordered above and below by dark patterning. Some specimens have a dark spot at the dorsal fin base; some a small dark triangular spot at the middle of the body near the base of the caudal fin (midpeduncular spot); some with a humeral semi-ocellus.

Can be visually distinguished from other species by combination of a very long adipose fin which originates more or less directly after the dorsal fin, very long maxillary barbells which extend at least to the posterior point of the body, the white lateral streak, and the caudal fin lobes curling inwards.

Mystus armatus (Day, 1865)

Day described this species as new in 1865a from rivers and backwaters of Cochin, India, but did not include any details regarding its colour or pattern, but did thankfully say (amongst other morphological details) that the base of the adipose fin equalled that of the anal fin, the supra-occipital reached the basal bones of the dorsal fin, and the fontanel almost reached the origin of the supra-occipital process.

In 1865b he used the same morphological information but went on to say that the colour and pattern was “Bright leaden silvery, lightest along the sides and with a purplish gloss over the cheeks. A black spot just anterior to the root of the dorsal spine. Fins finely dotted with minute black points.” He then stated that the adipose fin commenced a short distance anterior to that of the anal fin.

Unfortunately no figure was published of the whole fish and as yet the type specimens have not been correctly located although BMNH 1865.7.17.21 may constitute one of the syntypes (one of a number of type specimens originally used by Day), on the other hand it may be the specimen used by Day in 1865b.

There is no mention of stripes, but the reason I have included this species is that in Day (1877, Plate CI fig. 3.) a drawing is shown of a fish (which in my opinion is probably AMS B.7573, which Ferraris et al. 2000 have correctly stated is not a type specimen of M. armatus) with a mid-lateral stripe terminating in a blotch on the body near the base of the caudal fin, and the adipose fin being much longer than the anal fin base. In my opinion this is a misidentification by Day and this has led to Jayaram (1954) and subsequently (Ng 2002) and Jayaram & Sanyal (2003) incorrectly considering M. armatus as having a stripe along the body. The colour and pattern information from Day (1865b) means that M. armatus looks similar to Mystus cavasius (Hamilton, 1822), but with a smaller adipose fin than Hamilton’s species.

In my opinion the specimen figured in Day (1877) represents Mystus dibrugarensis (Chaudhuri, 1913), a species revalidated by me in 1999 (see under species heading for more details).

Mystus atrifasciatus Fowler, 1937 

See holotype ANSP 67907 (which originates from Pitsanulok, Thailand) and paratype ANSP 67908. Largest type specimen is 8.62 cm SL but is believed to grow to at least 11 cm SL.

Colouration as per original description: “Back and upper surface of head brown. Dark to blackish grey median lateral band, wide as vertical eye diameter and including lateral line, bounded above by whitish parallel longitudinal narrower band its whole extent, and below by whitish colour of under surfaces of body. Pale brownish streak, narrowing behind, back from pectoral axil until over front of anal. Iris grey. Lips pale or whitish. All barbels pale, with brown margins and nasal and maxillary pairs darker. Fins all more or less dull brownish.”

Mystus bleekeri (Day, 1877) 

See paralectotype AMS B.7999 which supposedly originates from Seharunpore/ Seeharanpore/ Sethrampoor, West Bengal State which now appears to be a suburb of Calcutta, India. And also specimen CAS 93966.

Colour from original description: “Brownish grey, with two longitudinal bands, one above the other below the lateral line, some specimens have a dark shoulder spot and a dark band along the middle of the anal fin. The fins are mostly darkest at their edges.” The “dark shoulder spot” is a tympanic spot.

Said to attain 12.5 to 13.5 cm SL.

Similar to Mystus atrifasciatus but M. bleekeri has shorter maxillary barbells (in M. atrifasciatus they extend to or past the caudal peduncle), the posterior edge of the adipose fin is angled off (versus rounded in M. atrifasciatus), and has the fontanel indented (versus not). Also similar to M. vittatus (Bloch, 1794); see under that species for information.

Mystus bocourti (Bleeker, 1864)

Junior synonym: Prajadhipokia rex Fowler, 1934

M. bocourti was originally described from the Mé-Nam River at Bangkok, Thailand; it is also present in Laos and Cambodia. It reaches up to 24 cm (SL). See holotype (MNHN 1553).

It is sometimes still listed in the genus Heterobagrus Bleeker, 1864, although this is currently considered to be a synonym of Mystus.

This species is usually uniform silver, bronze or platinum in colour but I have seen at least one specimen that has two light bands above and below the dark lateral line (Burgess, 1989, Plate 4).

As per Roberts (1994) it also sometimes has a humeral semi-ocellus and / or a dark spot on the body near the base of the dorsal fin spine.

It is easily differentiated from the other species by the extraordinary long dorsal fin.

Mystus canarensis Grant, 1999

This name was described by myself as a replacement name for Hara malabarica Day, 1865(b) as that species belongs in Mystus and contrary to Jayaram (1954) and other authors is not a junior synonym of Mystus malabaricus (Jerdon, 1849). There are no known preserved type specimens of M. malabaricus (Jerdon, 1849) therefore the original description of the colour and pattern are important: “blueish leaden above, silvery beneath; fins yellowish.” No semi-ocellus, or stripe(s) on the body are mentioned.

The type specimen of M. canarensis (AMS B.7624) measures 11.1 cm SL and originates from Canara, Karnataka State, India. It was also the specimen figured by Day (1877, Plate CI fig. 2) as “Macrones Malabaricus”, showing a fish with a tympanic semi-ocellus, a dark stripe along the lateral line, and spots on the fins making the upper portion of the dorsal fin and lower third of the anal fin appear blackish. It is similar at first glance to Mystus dibrugarensis and Mystus rufescens (Vinciguerra, 1890) - see under species headings for differences.

Mystus carcio (Hamilton, 1822)

There are no known preserved type specimens of this species which was described from the ponds of northern Bengal.

In Hamilton (1822) there is no reference to any drawing but as per Hora & Law (1941), Plate 23, Figs. 60 of Hamilton (1822) are erroneously captioned as Pimelodus batasio (which is now Batasio batasio), and should have been captioned as Pimelodus carcio. It is quite obvious when one reads the description of batasio and carcio that Hamilton made a mistake with designating which fish figs 60 represented. Thus we can quite clearly see what M. carcio looks like. The specimen in my image matches exactly the written description and figures in Hamilton (1822).

This species is a dwarf one, only reaching approx. 3-4cm SL.

It can be differentiated from all others by its small adult size, very small adipose fin, a dark horizontal mark across the tympanum, greenish yellow colouration, large laterally placed eyes, and relatively long and wide fontanel which is distinctly split by a thick epiphyseal bar. Sometimes confused as representing M. vittatus.

? Mystus colvillii (Günther, 1874)

See image of a type specimen and see discussion under Mystus pelusius (Solander, 1794).

Mystus dibrugarensis (Chaudhuri, 1913)

See image of the holotype, which came from Dibrugarh, Assam, India and measures 6.8 cm total length. Colour and pattern described as: “Head grey, dorsal side dark brown, body brownish. The membranous covering of the air bladder behind the gill openings is black, and a black line from above this membrane extends through the middle of the side to the middle of the root of the caudal fin, ending in a black circular blotch. The barbells are black, except the inner mandibular, which, with the fins, is dull white.”

This species was originally described as Macrones montanus var. dibrugarensis. Contrary to Jayaram (1954) and subsequent authors it is not a junior synonym of Mystus montanus (Jerdon, 1849). There are no known type specimens for M. montanus so again therefore the original description is important: “greenish above and on the fins; yellow on the cheeks and beneath.” There is no mention of any stripe(s) on the body, or semi-ocellus. The fish pictured in Day (1877, Plate CI, fig. 4) captioned as “Macrones Montanus”, is not the true M. montanus of Jerdon but appears to represent Mystus pulcher (Chaudhuri, 1911). Day’s incorrect identification has (as in armatus and malabaricus) caused later ichthyologists to misidentify the true M. montanus.

M. dibrugarensis differs from M. canarensis by having the supraoccipital process raised, long, and touching the proximal radials (versus not raised, very short, and not touching); body not elongated; caudal fin lobes being equal (versus upper lobe being longer than lower lobe). Also see notes on M. pulcher and M. rufescens.

Mystus gulio (Hamilton, 1822)

This is the type species of Aspidobagrus. See image of an eyeless specimen.

Originally described from “Higher parts of Gangetic estuaries”, this species lives in fresh and brackish waters. Sometimes when young it can exhibit pale stripes along the body.

It is easily differentiated from the other species by the combination of its greyish silver colour and small adipose fin.

Can reach approx. 40cm SL.

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Mystus horai Jayaram, 1954

See image of a type specimen.

It has been included here as it was originally described as a sub species of Mystus vittatus (Bloch, 1794), although according to the original description there are no stripes on the body of M. horai, just a faint black ‘shoulder’ mark. Preserved colouration said to be brownish yellow above, dull grey underneath. Type specimens originate from the Indus River, Kalabagh, Pakistan; the largest specimen being 8.4 cm SL.

Mystus keletius (Valenciennes, 1840)

Originally described from Java, and Pondicherry. Ng (2002) has examined the type specimens and concluded that the one from Java represents Mystus nigriceps (Valenciennes, 1840); therefore restricting the type locality for keletius to Pondicherry, India.

See image of the lectotype (MNHN A.9011) from Pondicherry, India; the specimen measures approx. 9cm SL.

Valenciennes states that it was very similar to Mystus tengara (Hamilton, 1822) in its colouration and pattern, but differed due to the supraoccipital and humeral processes being more granulated, the supraoccipital process being longer, the dorsal fin appearing rounder and the maxillary barbells being shorter.

Baensch & Evers (2002) show a picture and state that it has a yellow/silvery colouring, with a “dark shoulder mark”, two silvery to golden strips bordering the lateral line, and black marks near the dorsal and caudal fin. Jayaram & Sanyal (2003) state it is "Brownish turning dull white beneath. A dark shoulder spot and a light band along lateral line present. Dorsal and caudal fin tips tinged black, anterior portions of anal fin black".

Ng (2002) feels it is probably a synonym of Mystus armatus, or Mystus vittatus (Bloch, 1794). But as one can see from the image of the lectotype, this species is much more slender and elongated than M. vittatus and the shape of the fontanel differs. Also, as mentioned earlier the pattern of M. armatus is being misunderstood anyway, so I feel that M. keletius is a valid species.

? Mystus misrai Anuradha, 1986

See discussion under Mystus pelusius (Solander, 1794).

Mystus multiradiatus Roberts, 1992

See holotype (CAS 76119) which originates from Prachinburi market, Thailand. Largest type specimen is 12.8 cm SL.

The small gap between supraoccipital process and basal bones of dorsal fin spine help visually differentiate this species from the similar M. atrifasciatus. It is also distinguished from the similar M. bleekeri by the much less conspicuous and non-indented fontanel, and the lack of dark tympanic spot.

Mystus mysticetus Roberts, 1992

See holotype (CAS 76121) which originates from Nakorn Phanom market, Thailand. Largest type specimen is 12.9 cm SL.

This species has a humeral semi-ocellus, and the tympanum is darkly pigmented but is not a semi-ocellus. Tips of anal and caudal fin often black.

The laterally placed eyes (visible when viewed from above or below), and the combination of small adipose fin, humeral semi-ocellus, and dark tympanum differentiate this from all other species.

Mystus oculatus (Valenciennes, 1840)

Type locality is the coast of Malabar, India.

See holotype (MNHN 1195). According to Day (1877) there is no stripe(s) present on the body. The colour/pattern is said to be “silvery, lightest beneath, a dark spot at the commencement of the base of the dorsal fin, which is also black tipped, a darkish band likewise along the middle of the fin”. The reason I have included it here is that Jayaram & Sanyal (2003) state it has a dark band along the lateral line. They may be taking this erroneously from the figure in Day (1877), which in my opinion is just trying to illustrate the lateral line itself, not any band of colour or pattern along it.

The difference (or lack of it) between this species and M. armatus warrants further investigation.

Mystus pelusius (Solander, 1794)

Junior synonyms: Bagrus halepensis Valenciennes, 1840
                            Macrones aleppensis Günther, 1864
                            ? Macrones colvillii Günther, 1874
                            ? Mystus misrai Anuradha, 1986

M. pelusius is currently considered to be the type species of Mystus but the earliest designation has not yet been fully resolved. Solander described the species in 1794, from Kowick (Coic?) River, Aleppo, Syria based on earlier invalid names proposed by Russell in 1756 and Gronow in 1763.

Roberts (1994) tentatively considered colvillii (from the Tigris River at Baghdad, Iraq) and misrai (from Lake Antioche, Syria and Tigris River, Baghdad) to be junior synonyms of pelusius, although he did note that in the future colvillii may prove to be distinct. Thanks to Anthony Troncale of the American Museum of Natural History I have now managed to see the original drawing of pelusius from Solander (1794), and the text of the description, which is very basic in terms of colour and pattern information, just saying it was predominantly dark silver. The colour pattern of the alleged junior synonym misrai (see image of a type specimen) is (in alcohol): pale yellowish brown with head slightly lighter. No spots or stripes present.

It’s hard to say without comparing type specimens* but the drawing of pelusius looks more similar to colvillii than it does to misrai, although the profile of the dorsal fin is different in colvillii than the drawing of pelusius.

M. colvillii is said to be olivaceous in colour, with three narrow, white, parallel, longitudinal stripes, one along, one above, and one below the lateral line. The type specimens are “9 inches long”.

* according to Eschmeyer et al there are no known type specimens of pelusius. Roberts (1994) lists BMNH 1955.6.25.1 as a syntype of pelusius whereas Eschmeyer et al list it as a Günther specimen for the name Macrones aleppensis Günther, 1864 (a synonym of pelusius). In view of the date it was catalogued (1955) it is unlikely to be a syntype of pelusius, unless the specimen had been overlooked earlier or donated to the Natural History Museum at a later date.

Mystus pulcher (Chaudhuri, 1911)

Described from four small specimens (6.7cm total length) from near the Yunnan border, upper Myanmar. See image of a type specimen.

Described as “Dorsal and upper part of the body dark brown, with lighter or paler whitish brown stripes: one median, from the tip of the snout to the base of the dorsal spine, and two lateral longitudinal on each side, one above and the other below the middle line, which is distinguished by being dotted black for the openings of the lateral organs.” “…nasal and maxillary barbells blackish brown, adipose fin dark brown, dorsal, anal and caudal fins are brownish with black spots on the membranes between the rays.”

It has an intensely black tympanic spot/semi-ocellus, and also a spot near the caudal peduncle, followed by a thin white band. Its supraoccipital process meets the basal bones of the dorsal fin.

The difference in length of the supraoccipital process differentiates it from M. canarensis. The light body stripes differentiate it from M. dibrugarensis. Differs from M. rufescens by the fact that its fontanel is not split by an epiphyseal bar, and it has a shorter adipose fin.

Mystus rhegma Fowler, 1935

See drawing and image of holotype ANSP 61748 which originates from Bangkok, Thailand, and measures 4.96cm SL. Roberts (1994) lists specimens up to 10.6cm SL.

As per original description: “Very light or pale brown, lower or under surfaces more or less whitish. Upper surface of head and back sprinkled with dark grey dots. Band of dark dots along lateral line and broader one along lower side of trunk and tail parallel. Iris greyish, also maxillary barbell, other barbells whitish. Outer edge of adipose fin dusted with dark grey dots. Caudal dark grey. Other fins pale or whitish.”

This is a small and graceful looking species, which reminds me of species of the South American pimelodid genus Pimelodella Eigenmann & Eigenmann, 1888.

The lack of any semi-ocellus, the thin dark band across the lateral line, and large gap between the small suproccipital process and basal bone of the dorsal fin spine differentiate this species from all the others.

Mystus rufescens (Vinciguerra, 1890)

The type specimen measures 7.4cm (total length?), and originated from Meetan, Tenasserim Provinces, Myanmar. See image of the type, and another preserved non type specimen.

Vinciguerra described the colour and pattern as body tawny reddish, with one spot near the humeral region and one on the caudal peduncle. If one looks at the figure of the holotype there appears to be a thin black stripe along the lateral line. Roberts (1994) states that in live specimens that he identified as rufescens, there were two pale longitudinal stripes on the body, one above the lateral line, one below. He also states that the spot referred to as being near the humeral region by Vinciguerra, is actually a tympanic spot.

Differs from the similar M. canarensis, M. dibrugarensis and M. pulcher by having its fontanel split by an epiphyseal bar.

Mystus vittatus (Bloch, 1794)

Junior synonym: ? Pimelodus tengara Hamilton, 1822.

Bloch described the colour and pattern as head, ‘back’ fins, and caudal chestnut brown. Other fins steel coloured. Stripes light blue, with yellow interspaces. The plate of the holotype shows the general shape of the body and fins but the accuracy of its pattern is doubtful. See exclusive images of the holotype of M. vittatus: ZMB 2939 that originates from Tranquebar, Tamil Nadu, India.

Since its description in 1822 from “Ponds of India”, M. tengara has been discussed as being very similar to or perhaps a junior synonym of M. vittatus. Jayaram & Sanyal (2003) have classed them both as valid but they did not have access to the holotype of M. vittatus and as such they may have been using non type specimens and information from years of wrongly identified specimens which has just added to the confusion. They may have used misidentified specimens of M. carcio as representing M. vittatus, and therefore incorrectly distinguishing M. vittatus as distinct from M. tengara.

If one looks at the original drawings of M. tengara (see images) and compares them with the images of the holotype of M. vittatus, you will see that the structure of the fontanel and supraoccipital process are very similar, the only difference being that in the drawing of tengara the front portion of the fontanel is thin, the second is wider, whereas this is the opposite in the holotype of vittatus. This could be a mistake in the drawing. Everything else from the description of tengara appears to fit vittatus. If one looks at the images of ANSP 85780 from Bombay, India (which definitely represent vittatus in view of the exact same fontanel and supraoccipital morphology when compared to the holotype) you will see that the pattern, and shape of the body, especially relating to the high back and adipose fin matches that of the drawing of tengara. The adipose fin on the holotype of vittatus has shrunk in size due to its age.

The only possible evidence I have seen so far for classing M. tengara as possibly valid is that of specimen CAS-SU 34858 from Calcutta, India (see images). If you look at the lateral image of this specimen it appears visually very similar to that of ANSP 85780. But if you compare the fontanels of each specimen you will see that the fontanel of the CAS specimen is long and evenly portioned, unlike that of the holotype of vittatus and of the ANSP specimen. I feel that for the validity of tengara to be properly determined, future ichthyologists need to bear in mind that validity and identity of carcio when comparing lots of specimens from different localities. Until then, I feel that tengara should be classed as a questionable synonym of vittatus.

M. vittatus will reach at least 8 to 9cm SL.

M. vittatus differs from most species (except carcio) by the higher number of dark body bands, but can also be differentiated from the most similar species by:

M. atrifasciatus differs due to atrifasciatus having less dark lines on the body and having a thinner, non indented fontanel.

M. bleekeri differs due to bleekeri’s more elongated body, adipose fin and fontanel.

M. carcio differs due to carcio’s distinctly separated fontanel, much smaller adipose fin, and laterally placed eyes.

M. multiradiatus differs due to multiradiatus having a humeral semi-ocellus and having the fontanel not as visible.

M. mysticetus differs due to mysticetus’s large laterally placed eyes.

Leiocassis argentivittatus (Regan, 1905)

See image of a type specimen.

I have mentioned this fish here as in some aquarium publications it is placed in Mystus, due to Jayaram (1978). Jayaram & Sanyal (2003) place it in Mystus, however, Dai (1999) places it in Leiocassis Bleeker, 1858.

Gromov (1970) feels this species and the next one probably warrant their own new genus.

Described as being brownish with a silvery dark line across the middle and one along the ridge of the back, and a similar colour blotch on the upper portion of the dorsal fin, and two lines on the caudal fin.

Leiocassis mica (Gromov, 1970) new combination

See image of a type specimen.

Described from numerous small (approx. 3.5 cm) specimens from Lake Ommi, middle Amur basin, Russia. It was originally described as a Mystus species but its placement in Mystus is doubtful due to the shape and placement of the mouth, and the overall pattern and especially that of the caudal fin, which refers it towards Pelteobagrus Bleeker, 1864 but it has too few anal fin rays for that genus. If one follows Dai’s (1999) generic key it belongs in Leiocassis, and not Pelteobagrus, or Pseudobagrus Bleeker, 1859. It may well end up in a new genus along with argentivittatus and Leiocassis virgatus (Oshima, 1926).

Acknowledgements

Thanks to Jon Fong of the California Academy of Sciences, California, USA; Mark Sabaj of the Academy of Natural Sciences, Philadelphia, USA; Mark McGrouther & Sally Reader of the Australian Museum, Sydney, Australia; Rémi Ksas of the Muséum National d’Histoire Naturelle, Paris, France; Dr Peter Bartsch of the Institut fuer Systematische Zoologie, Museum fuer Naturkunde der Humboldt-Universitaet zu Berlin for the provision of the images of preserved specimens. Anthony Troncale of the American Museum of Natural History, New York, USA for the drawing of pelusius.

References

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Catalog of Fishes – online version. http://www.calacademy.org/research/ichthyology

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