Ichthyofile - The University of Kansas

Ichthyofile 
Number 2
Published by Steven Grant
30 October 2007
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The Library
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All Parts by Steven Grant1
PART 1
Fishes of the genus Botia Gray, 1831, in the Indian region
(Teleostei: Botiidae)
Key words: Botiidae, loaches, Botia, India, Nepal, Pakistan, Bangladesh, Bhutan,
Myanmar, Thailand, Tibet, Yunnan.
Introduction
In recent years there have been numerous works on fishes of the family Botiidae (I use
Botiidae rather than Cobitidae as per Šlechtová et al 2006); but most of them discuss
nomenclature of the names of genus rank and above, or are descriptions or synonymies of
the genera and species that inhabit waters of China, Laos, Thailand, Cambodia, and
Indonesia. In the last few years, only Kottelat (2004) and Ng (2007) have discussed
species of the genus Botiain any detail, describing new species from Myanmar. Nalbant
(2002) discusses Botia, but this is more about the interrelationships with other Botiidae
(phylogeny).
1
C/o 4 Milton Street, Castleford, England, WF10 1LW.
I will mainly discuss here the Botias of the ‘Indian region’ (see comments under
‘Distribution’).
The only recent work that has tried to resolve the identities and synonymies of ‘Indian
region’ Botia is that of Menon (1993). However the results have still left some doubt in
the minds of ichthyologists and aquarists alike. The synonymies proposed for the species
have little or no discussion about the reasons for them; and the photographs are of poor
quality. Having said this, Menon does provide some useful information that would not be
available otherwise. Before Menon’s review, the literature on Botia in this region has
been patchy, and much has relied on views of earlier authors, or misidentifications.
After discussions with a fellow aquarist (Andy Rushworth) we decided further
clarification was needed as we were becoming frustrated with the use of various names
for similar looking fishes that appear in the aquarium hobby, and the lack of clarity for
identification and discussion in scientific works. So a decision was made to try and see if
anymore light could be shed on the problem.
Building on the work of Menon, I will bring together here the myriad of papers on these
fishes, and combine this with information from preserved and live specimens, to provide
some (hopefully) useful information. However, I have found that although the status and
identity of some species has hopefully been resolved, I have also found this study has
produced more questions that cannot be fully resolved here. Rather than simplify and
reduce the number of names in usage, it has actually had the opposite effect as it is my
opinion that there are numerous species or subspecies within the genus that have
historically been lumped under one name. Although I accept that this work has perhaps
complicated the picture, I think this is more satisfactory than just lumping names together
when it appears that there are differences that should be recognised. The recent work of
Ng & Edds (2005) and Ng (2006) on Pseudecheneis catfish has shown that what was
once thought to be a few species spanning Nepal, India, Bangladesh, Myanmar Laos and
China are actually numerous species. Based on my interim findings I see a similar picture
with Botia.
As mentioned previously it is important to note that this is by no means an attempt to
thoroughly and finally review the genus as that is beyond my knowledge, time and
capabilities. The scarcity of specimens in collections, and the age and quality of them
makes it almost impossible to say with any absolute confidence that the results of this
study reflect a true picture of the genus. To do this, collections would have to be made
across the whole region, and fishes of differing ages and gender compared with that of
other localities; and what would be useful would be to capture the specimens and keep
them alive, observing their pattern changes as they grew. Menon (1993) did the former to
a degree, and is the only attempt so far to show this; but as stated previously he failed to
show or discuss reasons for his synonymies. He was better placed than most, having easy
access to important specimens in Indian collections, to show the variances in pattern and
colour across different distributions and the changes that happen as certain species groups
grow, or become sexually mature (also including morphological changes associated with
this). See ‘Identification by pattern’ for a further discussion of this point. Mitochondrial
DNA analysis may also be useful in any future studies, particularly for the B. almorhae
complex.
As well as access to important museum specimens, and information on them, the
information from those involved in the export, retail, and keeping of these fishes has been
invaluable. I consider that that information is a prime example when aquarists and
aquarium exporter/retailers can provide a valid and useful role in the study of fishes.
Materials and methods
Descriptions of colour and pattern and some meristics and morphometrics are by a
combination of live (or unpreserved) specimens (some with accurate locality data), and
any preserved specimens mentioned. Unfortunately I have not been able to physically
examine the preserved specimens myself, I have had to rely on images, x-rays, and
thankfully the staff that I have listed in the acknowledgements have kindly provided me
with any other information needed. Counts for caudal fin rays do not include the
procurrent or outermost principal rays. Counts for fin rays will class a ray as separate,
even if it shares the same pterygiophore as the preceding one. For dorsal rays, the last two
rays (sometimes counted as 1 ray or ½ ray by some workers) are split down the middle,
or are unbranched, and they usually do not have a normal or any pterygiophore.
However, I have counted them as 2 rays as when people are counting rays on live
specimens they will usually just see them as 2 rays, rather than 1 or ½ ray. Counts for
principal rays may not be accurate as some counts have been taken from references, or
un-x-rayed material, whereas some are from x-rays which usually shows some rays under
the flesh. I have used the term ‘sp.’ if I consider that this represents an undescribed
species/subspecies that I can also provide some specific information on their
zoogeography and how it differs from other species. I have used the term ‘form’ if the
differences are more subtle or their zoogeography is not as demarcated, but in either case
the names or labels attached to them are not intended to form available species or
subspecies names. For the B. histrionica complex I have used the term ‘form’ because of
ongoing work by others more qualified than I. Material used is in: BMNH (Natural
History Museum, London); USNM (Academy of Natural Sciences, Philadelphia); CAS
(California Academy of Sciences, California); MNHN (Museum National d'Histoire
Naturelle, Paris), AMS (Australian Museum), MCZ (Museum Comparative Zoology,
Harvard), KU (Kansas University), OSUS (Oklahoma State University, Stillwater),
NHSB (Bombay Natural History Society). Some specimens only have circa
measurements if they are taken from images with scale bars as the institution has not
been able to provide the information. Unfortunately I have not had access to specimens in
Indian or Pakistani museum collections, some of them being types. However, based on
the limited use of morphological and meristic data in fishes of the genus (and the
information provided by Menon and the original descriptions) this has not been unduly
problematic when discussing validity of some of the species.
Distribution / zoogeography
By the term ‘Indian region’ I mean Nepal, Bhutan, Pakistan, India, and Bangladesh. I will
discuss connected areas: Myanmar (but not in too much detail as I am aware of work
ongoing on Botias of Myanmar), the region of Yunnan in China, and Thailand, as they
too have species of Botia within their borders. Based on what is known about the Tibetan
origin of some of the major rivers or tributaries which Botias do occur in (Negi, 1994),
one would think that Botias may occur in southern (Himalayan) Tibet (the Yarlung
Tsangpo [Brahmaputra system], and the Sengge Chu [Indus system]) However, Wu &
Wu (1992) and Zhang et al (1995) did not report any Botia from Tibet [Xizang], and
Edds (1993 & 2007) did not report any loaches in the upper waters of Nepalese rivers.
The waters there may be too cold and this may act as a natural barrier (see Petr 1999, and
Petr & Swar 2002).
Rather than dwell on countries and their man drawn borders, I should expand more on the
distribution in waters of the region; as I can say with certainty that fish do not require
passports to pass through different countries! The findings of Menon (pgs. 12-13 although not explicit on the point), and comments from Kottelat (2004) led me to
consider whether each major river system (and/or perhaps major river) has its own
species of Botia, and if this is the case, if this could contribute towards formulating the
identity and validity of the species in question.
I would have preferred to be able to include graphs with distribution markers for each
species or form, but I have not been able to access any that I could reproduce. The major
river systems or rivers that Botias are currently known to occur in are the following (only
certain and relevant tributaries or distributaries, and states/countries will be mentioned
and by no means does it infer that they are restricted to the ones mentioned here):
Indus River System: The Indus River rises in the Tibetan plateau and enters the
Himalayas in Ladakh. Thereafter it flows through the Ladakh region of Jammu and
Kashmir (India) before entering the plains of Pakistan. It has several tributaries that drain
the western Himalayas, the important ones being Jhelum River (originating in Jammu and
Kashmir), Chenab, Ravi, and Beas (all originating in Himachal Pradesh, India). The
Sutlej (or Satluj) River rises in southern Tibet, cutting through the main Himalaya into
the plains of Punjab (India) (Negi, 1994). Kabul River is a tributary. See Rafique (2000)
for more details.
Ganges River System: The Bhagirathi and Alaknanda rivers join to form the Ganges
River (all in the Uttarakhand Himalayas of India). The river then flows south east through
Himalayan valleys into the north India plains. The Gangetic Plains form a major river
basin and delta. Its tributaries include the Kosi, Sone, Karnali / Ghaghra and Gomti
rivers, and a major river, the Yamuna. The Yamuna joins the Ganges in Uttar Pradesh
(India). The River Hooghly branches off from the Ganges and drains into the Bay of
Bengal after passing by Calcutta [Kolkata]. The main stream of the river enters
Bangladesh (where it is known as the Padma river), where it meets with the Jamuna River
of the Brahmaputra system. They combine with the Meghna River and drain into the sea
in the Bay of Bengal.
Brahmaputra River System: The Yarlung Tsangpo River rises in the Tibetan Himalayas.
The river enters India in Arunachal Pradesh, after its confluence with two other rivers it
becomes the Brahmaputra River. It stretches through Assam (India). In Bangladesh it
splits into two branches. The larger branch is called the Jamuna which joins with the
Ganges (Padma). The other branch (Old Brahmaputra River) meets the Meghna (which a
tributary of are the Surma and Barak rivers). Both branches meet and then flow into the
Bay of Bengal. The Teesta (or Tista) is a tributary.
Salween River basin: Rises in Tibet, and flows through Yunnan (as the Nujiang River). It
runs south through Myanmar (as the Thanlwin), and then enters the sea. The Suriya River
is a tributary. The Ataran (or Kasat) River drains towards the Salween, where it enters the
Andaman Sea. They appear to be connected via the Gyaing River.
Irrawaddy [Ayeyarwaddy] River basin: Rises in the north east of Myanmar near the
border with China, and runs south west towards the ocean. In Yunnan it is known as the
Daying Jiang River; the Longchuan Jiang River in Yunnan (called the Shweli River in
Myanmar), also drains into the Irrawaddy. The Chindwin River of Myanmar flows into it,
and the Chindwin has connected rivers that enter into Manipur (India). Although the
Sittang River is not connected, it is theorised that the lower course of the Irrawaddy
occupied the valley, pre-Pleistocene era. I class the Pegu (or Bago) River as part of the
Sittang system. The Myitmaka River of the Irrawaddy drainage meets the Pegu River at
Yangon.
The results of this study (albeit far from being the full picture) have led me to believe that
generally one should pay close attention to the river system or drainage that Botia
specimens are found in, as it appears that species may be restricted to certain ones rather
than the very wide almost all encompassing ranges that have been given for them in
various publications over the years. As Kottelat (2004) states (when discussing
phylogeny of genera), zoogeography has to be derived from phylogeny, so I am not
trying to propose any phylogenetic tree using zoogeography, merely commenting that
zoogeography can be useful tools when considering identity or validity of species.
Because some of the river systems now merge, or have merged in the past, some species
will be present in more than one system or river. However, generally for the currently
described species the zoogeography when looking at major systems or rivers appears (on
a simplistic level) like this:
Indus system
Upper Ganges system
Mid course of the Ganges and the Himalayan drainage to it
Lower Ganges
Upper Brahmaputra
Lower Brahmaputra
Surma
Krishna
Sittang
Ataran
Tenasserim
B. birdi
B. almorhae
B. lohachata, B. dario
B. dario, B. rostrata
B. dario
B. dario, B. rostrata
B. dario, B. rostrata
B. striata
B. histrionica2
B. kubotai
B. udomritthiruji
The occurrence of B. striata in the Krishna, which is some distance from any of the other
systems/rivers above, and without any current hydrographic connection is discussed in
Menon (1993:24), where he also discusses ‘waves’ of species distribution.
Identification by pattern and/or colour
Aquarists know that the patterns of some Botia can change dramatically as they grow,
and can vary even in fish of the same size. Because of similarities in morphological and
meristic information between species, pattern can give us an important comparator to
help try and differentiate species. Using patterns alone has usually been frowned on by
some as methods of separating species; however Menon (pg. 13) stated that he had relied
2
But see notes under B. histrionica regarding which river system actually houses the true B. histrionica
mostly on colouration to distinguish the Botiidae species. Also Kottelat (2004) felt (with
some reservation) that for at least two types of species in this genus pattern was
“conspicuously diagnostic”. His comparison with another species was predominantly
based on the differences in pattern in ontogeny (as well as distribution). I have struggled
to come up with other factors that are as useful.
Having witnessed the different patterns of many specimens (live and some preserved)
from different river systems or major rivers, I agree that in this genus pattern is an
important factor to use as a diagnostic tool. As aquarists we are well placed to see in live
specimens the colours and more importantly the patterns that these fish exhibit at various
growth stages, and aquarists and exporters like the groups that frequent Internet forums
such as Loaches Online and Petfrd, have played an important role in obtaining visual and
anecdotal information on the varying patterns of specimens, be it due to ontogeny,
variations, or aberrant patterns. Bogdan Janiczak, and Mark Duffill have taken
photographs of varying specimens as they grow, and these have contributed to this article.
I have found that although patterns do have variations (and in the B. almorhae complex
this becomes more problematic), and there are aberrant patterns, in some species the
number and positioning of bars is almost always constant. The problem is when the bars
start to anastomose3 due to ontogeny, but I also feel that in some species they can
‘disintegrate’. By this I mean when the solid dark bars end up with pale (colour depends
on species/form) spots within them.
As well as or instead of anastomosis and ‘disintegration’, the dark bars can start to reduce
or break off to form smaller dark particles and end up being unconnected dark spots,
blotches, bars or reticulations (depending on the species); and/or in at least two species,
some or all of the bars split to form two separate bars. I will call this fragmentation.
Based on what I have observed, I agree with Kottelat that patterns can be very useful
and/or diagnostic when trying to identify and separate Botia species. However it is not
failsafe as some species have a very variable pattern. For the similar B. almorhae
complex, colour differences (albeit subtle) can also play a role.
Morphology & morphometrics
Menon (pg. 13) used the following morphological and morphometric characters to help
differentiate the species: body depth, length of the snout, length of ventral fins in relation
to anal opening, and depth of caudal peduncle in relation to its length. I have found that
the majority of these generally unhelpful as they can all fluctuate depending on size and
gender. Hora (1922) used the length of snout in relation to the distance between the eye
and the opercle (snout length compared to remaining part of measurement for head
length), and then the position of the eye as to which portion of the head it was placed.
Again, I find these factors unreliable for the same reasons. He then drilled down further
using the position of the dorsal fin, eye size, and pattern. I have found that lateral head
length in SL is so close and overlapping (all specimens I used 3.1 – 4.6) in all Botias to
not be of any use. Out of any of the proportionate morphology measurements,
proportionate body depth can be useful, and proportionate snout length more useful,
3
To connect, or branch together. This word appears to have first been used by Hora (1932), but has also
been used by Menon, and Kottelat.
when used in conjunction with other factors (colour, pattern, distribution). However,
depending on the number of specimens used, and the proportion of gender in the
specimens, results can be misleading so they should not be given too much weight on
their own. When using proportionate body depth in SL it has to be borne in mind that
smaller fish will be not as deep bodied, even when taking into account their SL, and
female specimens will have a deeper body in SL. Snout length as a proportion of SL also
seems to vary, and appears to be attributed to gender (males with proportionately longer
snouts).
Hora also stated that in two similar species the swimbladder was of different relative size,
and that the anterior chamber differed in the proportion of it that was enclosed in bone.
He (1935) again commented on this and provided some drawings. Menon did not
mention this character for Botia, but on page 14 mentioned it for Cobitids and stated it
was not of any use. Of the Botia specimens that I have seen x-rays of, they almost all
have a very similar if not the same structure to the bony portion of the swimbladder. The
only one specimen with a notable difference is that of BMNH 1860.3.19.910, which has
the anterior portion encased in a narrow tubular shape. Unfortunately this specimen has
no identifiable locality data but is possibly B. lohachata, although I have been unable to
compare it with x-rays of topotypical B. lohachata.
Incidentally, I have compared the morphology of the swimbladder of numerous Botia
specimens from BMNH , USNM and AMS, with that of some Syncrossus, and some
Yasuhikotakia (USNM 107853) and found that, they are all more encased in bone
(anterior chamber) than in Syncrossus and Yasuhikotakia. This is something that Taki
(1972) discussed in some detail.
Meristics
Menon (pg. 13) mentions the number of branched dorsal fin rays as a character to aid
segregation of species but this is probably because he included Syncrossus berdmorei in
Botia. I have noticed that some populations, forms or species have a certain consistent
range of dorsal fin ray counts when compared to other species or forms or populations of
them. However, the differences are only small and the counts will sometimes overlap,
therefore as a character I consider that on its own it is unreliable.
Meristics of other fins are similar and vary within the same species, population, or form.
The only possible difference is that of a higher ventral ray count, but this is only one or
two extra rays.
Vertebral counts are usually within 3-4 of counts for other species, with the range being
the same or very similar. The average count can be different albeit usually by only a low
number.
To summarise I have found that the major river system and/or particular ranges, or
tributaries/distributaries of systems can be a good starting point to differentiating species
because those found in them tend to have different (albeit only subtle) base colours, and
also patterns in terms of the number and placement of them, and the way in which the
bars anastomose, disintegrate, and fragment, to specimens found in other systems etc.
Whether this is enough to justify considering them as different species or subspecies
needs further work using methods such as mitochondrial DNA analysis, as the factors
discussed here may not be enough.
Results
Botia Gray, 1831 – type species Botia almorhae Gray, 1831
Synonyms:
Hymenphysa MClelland, 1839 (and its unjustified emendation Hymenophysa Bleeker,
1858 (see Part 2).
Diacantha Swainson,1839
See Kottelat (2004: 12) for the characteristics of the genus.
Menon (1993) erected two complexes for the Botia species: the B. almorhae complex,
and the B. striata complex (his B. berdmorei complex is not relevant here as that species
belongs in Syncrossus Blyth, 1860 – see Grant (2001), and Kottelat (2004)). He did not
define the complexes, but based on his comments on pages 12, 13 and 24, he appears to
have grouped them together based on a combination of pattern and distribution. He
considered that B. almorhae, B. birdi, B. histrionica and B. dario belonged to the B.
almorhae complex, and B. striata to its namesake complex.
I too consider that species complexes do exist within Botia, but differ slightly on their
members. Without further study and specimens the description of the complex is only
loosely based, and is like Menon, mainly based on body and head shape, size, and
colour/pattern. Future DNA studies will be able to define them specifically and may of
course produce different results. I know that this is not scientific but to the trained eye it
is easy to see which species are closer to one than to perhaps others.
Botia almorhae complex
Members: B. almorhae; B. birdi; B. lohachata; sp. Teesta; sp. Kosi; (possibly B. blythi).
Diagnosis: Pattern consisting of black, grey, or brown Y and i bars (usually 3 Y’s and
approximately 4-5 i’s) in young, which usually disintegrate and/or anastomose to varying
degrees in ontogeny. Base colour whitish, yellow, golden, or very light green. Size up to
approx. 16cm SL.
Botia dario complex
Members: B. dario; B. striata
Diagnosis: Pattern consisting of usually 7-9 blue, green, grey or black body bars that
usually contain thinner lighter bars within them, the bars can anastomose. Base colour
usually yellowish to golden. Size up to approx. 12cm SL.
I have chosen B. dario as the namebearer of the complex (rather than B. striata as in
Menon) as I have a hunch that B. dario will probably be the basal member of the
complex.
Botia rostrata complex
Members: B. rostrata; sp. ‘Upper Brahmaputra’; sp. Ladder; (possibly B. blythi)
Diagnosis: Pattern consisting of 8-10 black to dark brown body bars that exhibit paler and
numerous spots in juveniles, and in adults the bars can anastomose, sometimes
completely obscuring the vertical bars on the anterior of the fish. Base colour whitish, or
yellowish to gold, sometimes with greenish or bluish tinges. Size up to approx. 12cm SL.
Botia histrionica complex
Members: B. histrionica (all forms); B. kubotai; B. udomritthiruji.
Diagnosis: Pattern consisting of 5 black bars (1st across nape and opercle) in juveniles
that have either no pale disintegration or (usually) 2-3 pale spots in small specimens. In
adults some of the 5 bars can split into two, or can anastomose, disintegrate, and/or
fragment to varying degrees. Base colour golden, yellow, light brown or whitish
(sometimes with a hint of green). Size up to approx. 13 cm SL.
Botia dario (Hamilton, 1822)
Junior synonyms:
Cobitis geto Hamilton, 1822
Canthophrys (Diacantha) zebra Swainson, 1839
Botia macrolineata Teugels, De Vos & Snoeks, 1986
Preserved specimens used: BMNH 1855.12.26.691, Ganges River, India, presented by
McClelland, 1 spec., c. 77mm SL; BMNH 1867.2.14.74-75, Cachar, India, presented by
Colonel Playfair, 2 spec. c. 61mm SL – 67mm SL; BMNH 1870.11.30.47, North East
Bengal, presented by T. Jerdon, 9 spec., c. 45 mm SL – 81 mm SL; MNHN 1984-0344,
(paratypes of Botia macrolineata), “+/- 100 km from Bombay, India”, presented by M.
Vanderweyer, 2 spec., 40.1 – 42.9 mm SL; CAS 94008, Ganges River, Patna, Bihar
State, India, presented by Tyson Roberts, 3 spec. 60.9 mm SL – 63.1 mm SL; ANSP
85779, “Bombay”, presented by Bombay Natural History Society, 1 spec., 67.0 mm SL;
AMS B.7534, “Sone River, Bengal”, presented by F. Day, 1 spec., 57mm SL.
Distribution: The type locality described by Hamilton was “northern rivers of Bengal”.
This is not specific, but historic Bengal relates to the region which is now current day
West Bengal and small parts of current day Bihar, Orissa, and Tripura (all states of
India), and Bangladesh. Hora (1935) states that Hamilton’s original notes and what he
knew of his journeys show that the specimen was from “Dumdumma”, which I believe
may equate to Dum Dum in West Bengal, which is near Kolkata, and if correct would
mean that the specimen probably came from the Hooghly River. It is definitely found in
the states of Assam (Sen, 1985; Banarescu & Nalbant, 1968; Menon, 1993; and
specimens above), Uttar Pradesh (Hora, 1922; Srivastava, 2004), Bihar (Srivastava, 2004;
and specimens listed above), Arunachal Pradesh (Nath & Dey, 2000), and West Bengal
(Shaw & Shebbeare, 1937, - Singhimari stream, and Chel River, Western Duars –
Northern Bengal; Menon, 1993). Petr (1999) lists it as occurring in Gaylegphug River,
southern Bhutan. Sen (1995) reports on specimens caught in Meghalaya (Khasi Hills,
Jaintia Hills, and Garo Hills, which are connected to either the Brahmaputra River, or the
Barak River, which connects to the Meghna River). Barak basin, Manipur (Vishwanath,
2002). Ataur Rahman (2005) reports it as being abundant in streams of Mymensingh,
Sylhet, Dinajpur, Rangpur and Chittagong Hill-Tracts, all in Bangladesh. See under
‘Specimens’ for instances in the Ganges. It is therefore predominantly found in the
middle to lower Brahmaputra River System (notably including the Meghna River and
Barak River), and Ganges River System. Sen (1985 & 1995) lists it as occurring in
Punjab but this is doubtful. It is not found in the waters of the Bombay [Mumbai] area
(see under discussion of Botia macrolineata).
Meristics: Dii-iii 8-12; Pi 11-15; Vi 6-7; Ai-iii 5-6; C 19.
Colour and pattern: Base colour yellow (dusky or light), yellowish gold, or greenish gold.
Snout dusky. A wide, dark bar across the top of the head, extends through and below the
eye; sometimes the bar is not separate from the dusky area on the snout and therefore
most of the head is dark. After the eye bar there is a bar that usually starts just above the
lateral line, and the ventral half of the fish, covering edge of the operculum and the area
on the body posterior to the insertion of the pectoral fin. The body then almost always has
seven bars, which connect over the ridge of the back. The first bar starts around the nape
area and almost always is angled diagonally backwards as it travels towards the ventral
part of the body. The next bar along has a similar angle. The next bar anterior edge is
almost always placed prior to the insertion of the dorsal fin spine. This bar also usually
extends at a backward angle as it travels down towards the vent. The next bars anterior
edge starts around the fourth or fifth branched dorsal ray insertion, its posterior edge
starting around the insertion of the last dorsal fin rays; it is more vertical than the
preceding bars. The next bar is also more vertical than the angled ones; its posterior edge
is near the insertion of the anal fin. The next bar is almost vertical and its edges usually
lie within the confines of the base of anal fin. The final bar is across the caudal peduncle,
and sometimes extends onto the base of the caudal fin rays. Sometimes the bars have a
thin line of the base colour within them (usually increasing in ontogeny). Sometimes the
bars end in the ventral half, or join the next bar ventrally and/or dorsally, which can give
the appearance of ‘u’, ‘n’, ‘v’, ‘Y’, ‘H’ or ‘o’ shapes (see Sane & Chhapgar, 1985, for 24
diagrams of the variations they found in specimens from Silchar, Assam (probably from
Barak River?)). The bars can be slate grey, dark green, or almost black (darker the
smaller the specimen). Across the base of the dorsal fin, the colouration of the two body
bars usually extends on to the base of the rays and spine. Across the middle to slightly
upper half of the dorsal fin, the spine and rays usually have a dark stripe. The first few
rays of the pectoral fin are sometimes dusky; sometimes basal portions of the rays are
dark, and sometimes there is a dark band on the rays along the middle of the fin. The
ventral fins sometimes have a dark blotch at the base of some of the rays, which can
sometimes cover the whole of the fin, apart from the distal portion. The anal fin
sometimes has a diagonal band across the lower half. The caudal fin can have two (or
three in specimens over 5cm SL) vertical bars, but they are sometimes angled diagonally.
In some small (circa. under 3cm SL) specimens the first bar is in fact a blotch or spot, and
the second bar is sometimes missing or very faint.
Discussion:
Cobitis geto Hamilton, 1822 - Prior to Menon, 1993 (who classed C. geto as a junior
synonym of B. dario) there had been much debate about the validity of C. geto. Günther
(1868) appears to have been the first to consider C. geto as the young of B. dario and
chose B. dario as the valid name, therefore establishing B. dario as the senior synonym.
He did not give any justification for this. Day (1872) questionably listed it as a junior
synonym of B. dario. Various other authors have listed it as valid, but without any or
much explanation. In 1922 Hora classed C. geto as valid, based on some poorly preserved
specimens from Gorakhpur (Uttar Pradesh, India). He said the “colouration is, however
very distinct and seems to be characteristic of the species”. However, in 1932, Hora said
that on further examination of the material he was of the opinion that C. geto is in reality
a young form of B. dario; the slight differences in the colouration of the two forms being
due to the age of the specimens.
I have considered whether C. geto is in fact Botia sp. ‘Upper Brahmaputra’. However, I
do not consider that C. geto is synonymous with that species or with the similar B.
rostrata (see under Botia sp. ‘Upper Brahmaputra’ for a discussion).
Hamilton described C. geto and C. dario in the same publication. I have already
discussed where Hamilton’s specimen of B. dario came from, and C. geto was described
from the ponds in the north-eastern parts of Bengal. Hora (1935) states that the C. geto
specimen was from Goalpara in Assam, which would mean the specimen probably was
caught in the Brahmaputra. Rao (personal communication) has fished in the vicinity of
Goalpara and did not catch any Botia, but as you can see under distribution, B. dario is
found in the Brahmaputra. The specimens used are not preserved in any institution. When
one looks at the drawings that he published of both species, a person unfamiliar with live
Botia species may be excused for classing them as different. Hamilton obviously saw
these fish (or the drawings of them) and considered them different because of the
apparent differences one may see in the pattern, but also because of alleged
morphological and meristic differences.
The drawing of B. dario is immediately recognisable as the fish we accept as B. dario
today. The pattern is either of a semi-adult, but probably of an adult specimen as it has
the paler lines distinctly formed within the dark bars, and the caudal fin has the 3
vertical/diagonal bars that have usually formed in specimens of 5cm SL+ (and Hamilton
states that it grows to 2 or 3 inches in length). The drawing and text description of the
colour and pattern shows that there was no pattern in any of the fins except the caudal.
This can easily be explained by the fact that the fish was stressed (or dead!) at the time
the notes and drawings were made, and that in some 5cm+ specimens the markings on
those fins can be pale.
The colour and pattern of geto was described as “yellow colour, with transverse black
bars running obliquely backward on the sides. One of the bars passes through the eye,
which is black, with a gold-coloured ring round the pupil. The fins are yellow, with two
black spots on the tail.” No size is given for the species, and the drawing is small.
Although this does not definitely mean the specimen used was small, I consider that it
was, based on how Hamilton’s drawings represented the fishes he described, and based
on him stating it had no lateral line. If one looks at small B. dario specimens, some do
exhibit a yellow base colour, darker bars than adults, and some have only two marks on
the caudal fin (or with two more very pale indistinct marks at the extremities of the
lobes). Sane & Chaapgar (1985) show one such small specimen, whose pattern is almost
exactly like that of the drawing of C. geto. What is very noticeable is that in the drawing
of C. geto, the dark area around the snout is shown, the band through the eye, and then
seven bars on the body, the first few being more oblique than the others, and the
gradually becoming vertical, and the last one being a wider band and covering the caudal
peduncle. The pattern on the caudal, the colour, the number of body bars, and the
placement of them represent a small B. dario.
In the text of the descriptions he also lists some other differences between the two species
he described:
•
•
•
He stated that B. dario had 6 pairs of barbels, whereas C. geto had 8. All Botia
species have 8 barbels and Syncrossus 6, so we can rule out C. geto being a small
Syncrossus (Yasuhikotakia have 6 also but it is doubtful that any occur in this
area). We know that B. dario does have 8 barbels, so this must be a mistake in the
text and the drawing.
He stated that B. dario had a lateral line, and C. geto did not. As far as I can
ascertain, all Botia species have a lateral line. If this is the case, I can only assume
that this is a mistake by Hamilton, possibly caused by the small size of the
specimen.
He gave the following fin meristics: D11 P13 V8 A7 C18 in B. dario, D12 P13
V9 A7 C no count in C. geto. All these counts (some include branched and
unbranched rays) are within the range for B. dario, with the exception of the
ventral fin count. 9 ventral rays is not the norm for Botia, except one found in the
Khasi Hills of Meghalaya (see further on); although sometimes the last branched
ventral ray is split to the base and is sometimes counted as one ray, or two. Some
Syncrossus species have 9 and the drawing of C. geto does seem to show a long
pointed snout, but as stated earlier Hamilton clearly states that 8 barbels are
present in C. geto, thus ruling out a young Syncrossus. I therefore consider that
again, this count is a mistake, possibly due to the specimen being small.
Based on all the above information, I consider that C. geto was a young B. dario, and as
Günther chose B. dario to have priority, C. geto is its junior synonym. However, this
species has a wide distribution and there are meristic differences in fin ray count ranges.
Having said this, the species appears to have a similar pattern and appearance across its
range, but perhaps with some differences in the base colour and darkness of the body
bars. DNA analysis may later show that certain populations are distinct from each other
and the possibility of different subspecies being in existence shouldn’t be ruled out. If
this happened, C. geto is an available name if the Assamese population turns out to be
different to the population in the lower Ganges’ distributaries.
Botia macrolineata Teugels, De Vos & Snoeks, 1986 – This species was described from
“+/- 100 km from Bombay, India”. It was based on 3 specimens. They were imported by
a Belgian tropical fish company, and that is where the locality information will have
originated. They had been kept alive in an aquarium for just over 2 years, before being
preserved. This species first came to my attention, when Menon (1993) published a
drawing as an addendum, and stated that it differed from B. dario by it having 11
branched dorsal rays, whereas B. dario was only known to have up to 10. The drawing
used does look like a B. dario, but also seems to be slightly different. Upon obtaining the
original description I was surprised that the drawing in Menon was not the drawing used
in the description. The drawing used in the description is that of a B. dario. Having seen
the paratypes of the species, these two represent a B. dario in all aspects.
The difference given by Teugels, De Vos & Snoeks is that B. dario has a smaller eye (2.1
– 2.3 times in snout length v 3.0 times in B. dario). In the specimens available to me I
have found great variation in eye in snout length measurement. Even specimens of a
similar size have different proportions. The range of 2.1-2.3 (based on 3 specimens) for
B. macrolineata falls within the range I have found in 16 specimens (2.2 – 3.1).
The difference of 11 branched dorsal rays in B. macrolineata highlighted by Menon, is
given as 11-12 by Teugels, De Vos & Snoeks. Two of lot BMNH 1870.11.30.47 have 11,
and one has 12 branched rays (the last ray is not always branched). The AMS specimen
has 11. Sen (1995) reports on some of his specimens having 11, and I have seen the
images of live B. dario specimens which also have 11.
Teugels et al also give a caudal fin ray count of 26-28, but this count includes the
procurrent and outermost principal rays.
Another issue regarding B. macrolineata is the type locality. If the type specimens had
indeed been collected from the watercourses in Bombay (Mumbai), then this would have
been a marked deviation from where B. dario occurs. The rivers around Bombay are not
currently connected with those of the Ganges, Brahmaputra, or Meghna rivers (although
Rao & Yazdani (1977) discuss previous connections with the drainages of the Eastern
Himalayas). B. dario are indeed exported from Bombay, as a friend of mine has live
specimens that have been traced back as exported from Bombay, and these specimens
exactly match B. dario. ANSP 85779 is listed as being from Bombay, and was donated
by the Bombay Natural History Society. However, the fish exported from Bombay are
not caught there; they are caught from the range known for B. dario (likely to specifically
be Assam), transported to Bombay, and then shipped out (Chhapgar, personal
communication). I am told that a similar problem happened with Chela dadiburjori
(Menon, 1952) when Axelrod et al (1962) published a new record from the “vicinity of
Bombay” based on aquarium specimens that were exported from there. C. dadiburjori is
not from Bombay (Sane & Chhapgar, 1983).
Based on the identical colour and pattern, body and head shape, flawed locality data, the
eye in snout length measurement and dorsal fin and caudal fin ray counts falling in the
range for B. dario, I have conclude that B. macrolineata is a junior synonym of B. dario.
Whilst I was waiting for Part 2 of this work to be finished, Ng (2007) also decided that B.
macrolineata was a junior synonym of B. dario, based on comparisons with information
in Teugels et al (1986) and Ng’s B. dario specimens.
Botia striata Narayan Rao, 1920
Junior synonym:
Botia striata var. kolhapurensis Kalawar & Kelkar, 1956
Botia strigata incorrect subsequent spelling (see Axelrod et al, 1972?)
Botia ‘weinbergi’ unpublished and unavailable name (see Axelrod et al, 1972?)
Botia dayi (nec Hora) Rao & Yazdani, 1977.
Preserved specimens used: BMNH. 1919.11.19.8-12, (holotype and some paratypes of B.
striata), Thumga [Tunga] River, Shimoga, Mysore State, India; NHSB uncatalogued,
Panchganga River, Kolhapur, India, 1 spec., 36mm SL.
Distribution: Krishna River System - Tunga River, Karnataka; Koyna River
distributaries, and Kolhapur (Panchganga river subsystem), Satara District, Maharashtra,
India.
Merisitics: Dii-iii 9-10; Pi-ii 9-12; Vi 6-7; Ai-iii 5-7; C 19.
Colour and pattern: Base colour light yellow to gold, some specimens being more golden
than others, and having a metallic effect. Parts of all or all of the upper lip and/or the
lower lip can be orange or red. The head usually contains 3 bands (+1 along from the
snout, along the ridge of the head), and the body 9 bars. These bands and bars are of a
darker colouration, which can vary from almost black, purplish, through dark grey, to
greenish grey. These bars and bands usually contains lines of the (lighter) base colour,
smaller fish tend to have fewer and wider lines, but as the fish grow the lighter lines can
be thinner and there are more of them, and they can become fine reticulations. This gives
the impression of the dark bands and bars being thinner, but in actual fact they are still
usually there, they are just not as distinctly separate from the base colour as in smaller
specimens. Chhapgar & Sane (1985) show 17 examples of the variations that can occur.
Some specimens have the bars and bands connected with each other, horizontally, or
diagonally, some even to the point where one cannot see any ascertainable vertical or
diagonal bands or bars. In some ways, the variations are similar to those shown by Sane
& Chhapgar (1985) for B. dario, as some have ‘H’ or ‘Y’ patterns. Because of these
variations, the following description pertains to the average specimen.
Along the ridge of the head, from the tip of the snout to the area aligned with the middle
of the eye, is a band of the darker colouration. There is usually a band that starts on the
top of the head (just posterior to the aforementioned band) and spreads diagonally
downwards to the upper lip. The band usually ventrally contains a line of the base colour
within it. There is usually another band that originates posterior to the aforementioned
one, travels downwards diagonally, passing through the eye. It usually ventrally contains
a line of the base colour within it. There is then a bar which about halfway down the head
starts to run diagonally anteriorly, and passes over the opercle. The next bar starts to run
diagonally posteriorly, about halfway down, and ends around the base of the pectoral fin
(mainly over the base of the rays). There are two more similar bars, which are less
diagonal than the preceding bands and bars. The next bar usually has its middle at the
insertion of the leading ray of the dorsal fin; this bar also being slight diagonal, running
posteriorly. The next bars are not diagonal; the first one is within the base of the dorsal
fin rays; the next three on the posterior half of the body, and the last being on the caudal
peduncle and base of the caudal fin rays. The base of the dorsal fin usually has some
markings where the fourth and fifth dark body bar meet the fin, and there can be two or
three crescentic stripes along the middle to distal half of the rays. The pectoral fin usually
has three thick dark bands; one at the base, one in the middle, and another one in the
distal half. The ventral and anal fins usually have two dark bands, one usually near the
base and the other in the distal half. The caudal fin usually has three to five dark
crescentic bars (the smaller the specimen, the fewer bars there are), and the distal margin
of the fin is usually dark.
Discussion:
Botia striata kolhapurensis – described on the basis of 6750 specimens from Kolhapur.
The type specimens are no longer traceable in the Kolhapur, Satara and Sangli districts
collection, or the Taraporevala Marine Biological Station, Bombay (Daniel & Sane,
personal communication). I am told that thousands of specimens are exported from
Kolhapur and Sangli after the rainy season, to Europe and the USA. Kalawar & Kelkar
gave the following differences:
•
•
•
•
•
Maximum size of B. s. kolhapurensis 57mm Vs 95mm for B. striata. They state
that they never catch them at bigger sizes. This can be easily explained by the
localities where they are being caught, as it appears from information I have
obtained from collectors of this species and others, that only certain sized
specimens are sometimes found in certain locations.
Pectoral fin total ray count of 11-12 in B. s. kolhapurensis, and 13-14 in B. striata.
As can be seen in B. dario, a few more rays in fins is not enough to differentiate
species or even subspecies.
Depth of body in SL 3.5 to 3.75 in B. s. kolhapurensis Vs 3.33 in B. striata. This
can be explained by the differences in size of the specimens, how well fed or
preserved they were, or their gender.
Diameter of eye in head length is 4 to 4.25 Vs 5. As can be seen in B. dario,
differences in eye in head or snout length is not uncommon.
“Bands” on the body fading into the ground colour as they reach the ventral
surface, not completely surrounding the body Vs the opposite. Chhapgar & Sane
(1979) state that they examined hundreds of specimens of B. s. kolhapurensis,
they found two colour variations. In some specimens (mostly those that were
lighter coloured) the bands fail to meet the mid ventral line, but in other (darker)
specimens, posterior to the anal fin they do meet mid-ventrally, although on the
anterior portion of the body they stop short on the sides. They also state that in
juvenile specimens there are a few broad black bars on the body, and as they
grow, each broad bar splits up into (usually) three narrow stripes, until they reach
the final body colouration of the adult with numerous narrow stripes. They show a
drawing of a juvenile, and an adult. The adult is identical with Naryan Rao’s B.
striata, and the juvenile also looks similar to a young B. striata. Nothing in the
description of the pattern is different to that of a B. striata, other than the bars
stopping short in the ventral half of the fish (in some specimens). I have seen
aquarium specimens where depending on mood and surroundings, the fish
appears to have the full body bars, or the same specimen can have the ventral half
of the bars missing, and also some of the colour and patterning in-between the
bars of the upper half of the body can ‘disappear’, in what appears to be a fright
pattern. It matched exactly the figure of a juvenile in Chhapgar & Sane. I saw this
happen in a specimen that had been in a plastic bag and a white polystyrene box
for a few hours. After a short while in its new aquarium it developed the full body
bars. Also, Chhapgar & Sane state that they examined the specimen in Rao &
Yazdani (1977) and concluded that it was a B. s. kolhapurensis. The specimen
shows the bars fully reaching the ventral side of the body. Thanks to Sane, Daniel
& Giri of the Bombay Natural History Society I have an image of a preserved
topotypical specimen of B. s. kolhapurensis. The specimen is 36mm SL and was
caught at Kolhapur, in the Panchganga River, a tributary of the Krishna River.
This specimen is identical with B. striata, and shows the bands encircling the
body.
Based on all the above, B. s. kolhapurensis is a junior synonym of B. striata, and I do not
consider that subspecies level is warranted (even with the slightly divergent distribution).
Botia rostrata Günther, 1868
Senior synonym?:
Junior synonym?:
Botia nebulosa Blyth, 1860 – nomen dubium
Botia dayi Hora, 1932
? Botia geto (nec Hamilton) Day, 1878 & 1889
Botia dayi Shaw & Shebbeare, 1937
Botia dayi Ataur Rahman, 1989 & 2005
? Botia rostrata Sen, 1995
? Botia rostrata Ng, 2007
Preserved specimens used: BMNH 1855.12.26.694, syntype of B. rostrata, “Ganges”,
Bengal, presented by McClelland, 1 spec., 58mm SL; BMNH 1860.3.19.114, syntype of
B. rostrata, “Assam?”, presented by Griffith, 1 spec., 110mm SL. See discussion below
on localities.
BMNH 1872.4.17.104, “North East Bengal” (listed in the BMNH records as from
Bangladesh), presented by Jerdon, 1 spec., 107mm SL. Identity uncertain.
Distribution: Based on information I have from collectors, from references, and the
locality data for the junior synonym dayi:
Ganges system: Mahananda River (see discussion below), at Siliguri, at base of
Darjeeling Himalayas, West Bengal (Hora, 1932; Shaw & Shebbeare, 1937; and possibly
due to nebulosa), then possibly far eastern Bihar (both India), then possibly lower Ganges
and Padma; Bangladesh (Ataur Rahman, 2005 – who does not give specific areas or
localities).
? Lower Brahmaputra system: Sen (1995) had specimens of B. rostrata from
Rongrenggiri, Garo Hills, Meghalaya (probably in the Simsang [Someswari] River). He
also listed it as present in the Khasi Hills, but had no readily identifiable specimens to
back this up (see discussion below under B. blythi). Having said this, specimens that may
be conspecific with the true B. rostrata have been found in the Khasi Hills: Wahumiam
River, a tributary of Shella River, Shella, Khasi Hills, Meghalaya, (drains towards the
Surma and Meghna) (Rao, personal communication), and a specimen similar to B.
rostrata appears in Ng (2007) from the Piyain Gang River in Bangladesh, and this may
be conspecific with the true B. rostrata, or may be the same species or form found in the
Shella system. Therefore B. rostrata (or a similar species or form) is probably found in
the mid to lower courses of the Brahmaputra drainage, around the Old Brahmaputra,
Surma, and Meghna rivers / systems.
Merisitics: Dii-iv 8-10; Pi-ii 11-12; Vi 6-7; Aii-iii 5-7; C 19.
Colour and pattern (based on all localities discussed above): Base colour can be whitish,
but usually yellowish to golden, but can also have a greenish or even bluish tinge. Tip of
the snout, barbels, and lips can be golden to orange. The head and body have dark bands,
dark brownish to black. On the head there tends to be a dark band across the interorbital
region, which encircles the eyes, then moves horizontally towards the snout, and this
band can be disintegrated. A short bar is usually present under the eyes and one on the
opercle. There is usually a dark bar across the ridge of the snout. In younger specimens
the body bars are more distinct but tend to anastomose with each other more on the
anterior part. There tends to be a bar after the opercle, then two placed before the dorsal
fin insertion, two under the dorsal fin base, two after the dorsal fin, and then another one
or two near the caudal peduncle, the last bar usually being on the caudal peduncle. There
is usually a wider pale bar in-between the penultimate pair of dark bars and the last pair
or last single bar. The bars disintegrate and anastomose, and as the specimens get bigger
they anastomose and disintegrate to a greater degree. The dorsal fin usually has a dark
line across its base, then one across the upper half of the branched rays, then sometimes
another near or on the distal margins of the rays. The caudal has dark bands that vary
depending on the size of the specimen. The pectoral fins can have 3 to 4 thin dark bands
and the rays can be golden to orange. Ventrals 2 to 3 thin dark bars and same colour
range as pectoral fins, anal fin can have 2 thin dark bars and same colour range as ventral
fins and pectoral fins. The larger specimens tend to have the orange to golden colours on
the snout and lower fins. The pattern is similar to some B. kubotai but that species tends
to have fewer body bars (usually 5), and doesn’t appear to get the vivid colours of B.
rostrata. An aquarium specimen that came in with some members of the B. almorhae
complex (probably sp. Kosi) had the same pattern but the base colour was not as vividly
gold, but was whitish to yellow; this may represent the true B. rostrata, and the Shella
specimens shown here, and the ones in Sen (1995) and Ng (2007) may be a different
species or form.
Discussion:
Günther used the above type specimens to describe B. rostrata. He had what appeared to
be definite (albeit not very specific) locality data of Ganges, Bengal for one syntype, but
the other syntype has a questionable locality of Assam. In view of the differences
between Assam specimens (called in this publication Botia sp. ‘Upper Brahmaputra’) and
the lower Ganges specimens, I felt it was necessary to designate a lectotype. Which one
is chosen could determine which is the true B. rostrata, either the Ganges specimen
(possibly making B. dayi and B. nebulosa synonyms), or the “Assam?” specimen,
possibly meaning B. dayi / B. nebulosa a similar but separate species. However, at the
request of the BMNH and because it is not yet crucial for nomenclatural reasons, I am not
designating a lectotype. Due to a mix up with labelling in the 1800’s it is not now clear
that the smaller specimen was from the Ganges and the larger from “Assam?” (as had
been stated by Günther); they may have been mislabelled prior to Günther using them
(Maclaine, personal communication). In any case, there are pros and cons for choosing
either specimen due to one (could be either of them) having a clearer but still not defined
locality data, and possibly the other having been figured (larger specimen), but being in a
poorer state of preservation than the smaller one. It is not even definite (but likely) that
they are the same species as each other. They probably would not be if one was from
Assam and the other from the Ganges in “Bengal” based on what I have found. The
questionable locality of “Assam?” for one specimen could obviously be incorrect. Both
specimens appear to have the same number of fin rays, and the larger specimen only
having 1 more vertebra than the smaller (29 versus 28 total vertebrae). The larger
specimen does have a visibly longer snout but is almost twice the size (SL) of the smaller
one, and is probably an adult male (although proportionate snout length in SL is very
close – 5.9 times in the large specimen, 6.1 in the smaller specimen). The drawing in
Günther of the snout region of the larger specimen is somewhat misleading because the
specimen is slightly emaciated, thus pronouncing the structure of the bones underneath.
On the x-ray of the specimen there is a clear gap in-between certain bones behind the
snout region, and this gap is not visible on the smaller syntype. However, this appears to
just be because the larger syntype has its jaws protruded due to its mouth being opened.
Any ichthyologist working on Botia may take the view that it is more stable if the Upper
Brahmaputra ‘form’ is given the name B. rostrata, thus leaving the other ‘form(s)’ with
another scientific name (B. dayi, B. nebulosa or even perhaps B. blythi – see below). I
was tempted to do this rather than upset the relatively long held view of B. rostrata and
perhaps even the use of the name (due to the issue of B. nebulosa and even perhaps B.
blythi being possible senior synonyms, although article 23.9 of the Code may be usable).
However, based on the information to hand I consider that although the available figures
of proportionate snout length in SL shows the B. dayi specimens have a shorter snout (7.6
– 7.8 Vs 5.9 – 6.1 in B. rostrata syntypes) it is probable that the syntypes of B. rostrata
are conspecific with B. dayi because the probable locality of the B. rostrata syntypes
match closer to that of B. dayi than sp. Upper Brahmaputra, and the meristics are closer.
However, just because the B. dayi specimen locality drains into the Ganges where we
know one of the B. rostrata syntypes was supposed to have come from, it doesn’t mean
that they are definitely the same. The other issue is that the Shella specimens also have
different morphometric counts (possibly shorter snouts) and may be much more vividly
coloured than Ganges and Mahananda specimens. This conclusion has been based on the
above specimens, information from various other references, and information from non
preserved material with accurate locality data. Further study is needed with the use of
numerous freshly preserved specimens from the Mahananda, Teesta, Shella, Surma, Old
Brahmaputra, lower Ganges, Brahmaputra in West Bengal, Assam, and Arunachal
Pradesh.
Body Depth in SL
B. rostrata syntypes (2)
B. dayi syntypes (2 of)
Shella specimens (4, unpreserved)
Piyain Gang River (Ng, 2007) (1)
‘Ganges’ specimen (1, unpreserved)
4.6 – 6.1
4.1 – 4.7
4.0 – 5.3
3.6
4.3
sp ‘Upper Brahmaputra’ from Guwahati (5, unpreserved) 3.7 – 4.3
Snout Length in SL
5.9 – 6.1
7.6 – 7.8
6.6 – 8.9
8.2
5.9
6.9 – 8.8
Near to the end of this publication being finished, Ng (2007) considered B. rostrata to be
valid and not a synonym of B. almorhae, and thus did not agree with Menon or Kottelat.
Obviously the information here shows that I consider them to be distinct. Their pattern,
zoogeography and morphometrics show this. The only issue I would note is that the
specimens he used to compare the two may not be true B. rostrata or B. almorhae. The
specimen he used to represent B. rostrata may not be conspecific with the true one, and
some of the B. almorhae specimens (including the one pictured) are in my opinion sp.
‘Teesta’.
Botia nebulosa – Menon (1987) classed this species as a junior synonym of
Acanthocobitis botia (Hamilton, 1822), and to date this has been accepted (although only
tentatively by Kottelat, 1990). After reading the account of this species in Day (1878) I
accepted that this synonymy was possibly correct and so didn’t give it much further
thought. However, I was invited by Kamphol to look at Blyth’s original description of B.
nebulosa, and to look at the differences between it, and how Day described it in his
Fishes of India. Blyth clearly thought his B. nebulosa was a Botia, and not in the same
genus as other current members of Acanthocobitis, which he grouped under the genus
“COBITIS”, he also clearly saw the difference between Botia and the species he placed
in his new genus Syncrossus (something that is now, but only recently, accepted). The
description starts on page 165 and clearly states that it has a stout “forked” spine under
each eye, one of the spinous projections being longer than the other. He clearly states it
has 8 barbels. Under the subheading for Botia he says that their (including B. grandis =
B. almorhae) colours are “bright black and yellow”. On page 166 he goes on to state:
“Body imperfectly banded, shewing about seven irregular transverse bands, which are
double or dark only on their borders and more or less broken and confluent. Three
distinct lines of spots on the dorsal, besides its dusky tip: three also on the forked caudal,
besides the base and the tip: five in all on the pectorals; two on the ventrals; and two
(indistinct) on the anal. Colours black and gold in the recent specimen, as in the various
affined species; the markings tending to assume the spotted appearance proper to B.
GRANDIS.
The fin-rays are alike in both.
D. 10. – A. 6. – P. 13 or 14. – V. 8. – C. 19.
Length of specimen 5 ¼ in. Height of body 1 in; at base of tail nearly 5/8 in. From
Dorjiling: presented by the late Dr. Wallich.”
Day (1870) examined specimens in the ZSI in Calcutta and on page 549 (repeated in
1872, 1878, and 1889), and gives different fin ray counts for what he obviously
considered to be the same specimen that Blyth used. He gives the rays as:
“D. 3/12. P. 13. V. 8. A. 2/5. C. 17”
These counts are clearly different to Blyth’s and Day’s dorsal ray count appears too high
for most Botias (except B. dario) but the caudal fin ray count too low (19 in Botias). Day
also states that there are 6 barbels (Botia have 8, and Blyth said 8). Day states that the
sub-orbital spine was damaged (see below). Day gives a completely different explanation
of the colour pattern (notably mentioning a leaden band along the side, and an ocellus at
the upper margin of the base of the caudal fin); one that is reminiscent of what is
currently (see Part 3) being classed as Acanthocobitis botia. The combination of this, and
other factors led Hora (1922), Mukerji (1934) and then Menon (1987) to class B.
nebulosa as a junior synonym of A. botia. So, how come Blyth’s description differs
significantly to that of Day’s account of what is considered to be the holotype (Menon &
Yazdani, 1968)? I consider that this is because the specimen examined by Day and
subsequently by Hora, is not the same one that Blyth based his B. nebulosa on, and is
therefore not the holotype. Apart from the significant differences already mentioned, Day
states that the specimen is 4 ½ inches long. Blyth clearly states 5 ¼ inches. I considered
whether one was using SL and the other TL, but Day’s (1870) measurements of other
Blyth specimens tally with the ones given by Blyth in his 1860 work. Hora’s 1922
examination of the specimen shows that it does not have any suborbital spine at all,
showing that Day’s view that it was damaged was incorrect, as it just has the suborbital
flap present in some males of Acanthocobitis. Hora confirmed all the other characters
described by Day, but differed on the shape of the caudal fin as Hora had sight of an
earlier drawing of the specimen. Other characters such as the morphology of the
swimbladder, number and structure of the barbels, lips, jaws, and characters of the scales
showed that the specimen was not a Botia, but (what is currently accepted as) an
Acanthocobitis (Nemachilus [sic] of Hora, and Mukerji). As stated previously, I do not
consider that ZSI F2637/1 is the holotype of Botia nebulosa and as such do not consider
that B. nebulosa is a junior synonym of A. botia. Blyth’s description clearly relates to a
Botia, not an Acanthocobitis, and based on the purported locality, meristics, and
description of colour and pattern it is probably the same as B. dayi (and therefore possibly
B. rostrata). However until this is resolved with further searches for other specimens at
ZSI that could be the true holotype, or records of any other donations by Dr. Wallich to
the ZSI, Botia nebulosa should be a nomen dubium, even though it may turn out to be a
senior synonym of B. rostrata (unless B. dayi is proven not to be conspecific with B.
rostrata).
Botia dayi - described on the basis of 7 type specimens which Hora said were from the
“Mahanaddi River at the base of the Darjiling Himalayas”. I can find no Mahanaddi
River, but the Mahanadi River flows through the Indian states of Chhattisgarh and Orissa
and does not flow near the base of the Darjeeling Himalayas. There is however a
Mahananda River, which does flow from the base of the Darjeeling Himalayas, and I
consider that Hora meant the Mahananda, not the Mahanaddi/Mahanadi. Shaw &
Shebbeare (1937) also stated that they found specimens in the “Mahanadi” of their
sampling of Darjeeling and Jalpaiguri, but for the same reasons mentioned above, I class
this as a mistake (in fact the specimens they found appear to have been used by Hora for
his main type series for B. dayi). The Mahananda River flows through West Bengal, and
Bihar in India, then into Bangladesh where it joins the Ganges (or Padma). Hora also
used the specimen figured as Botia geto in Day (1878 & 1889). Day listed the locality of
this specimen as “a stream in the Sind hills”. Sind is what is now the province of Sindh in
Pakistan. If the locality data for this specimen is correct, it is very unlikely to be
conspecific with Hora’s West Bengal specimens due to the huge distance involved and
unlikely lack of recent4 hydrographic connection. If indeed it did come from Sindh, then
it is possible it represents a specimen of Botia birdi Chaudhuri, 1909 (there is a Sind
River in the Kashmir Valley which is a tributary of the Jhelum River); but its pattern does
resemble that of a B. rostrata. There is an alternative: the drawing is almost exactly the
same as the one in Menon (1993) at Fig. 4 on Plate 1. This drawing is remarkable in its
similarity with the one in Day. The drawing in Menon is captioned as that of a breeding
female of Botia almorhae (almost certainly of one of the specimens he reported on in
1949) from “Kosi R., Almorha” (he states it is from Uttar Pradesh but in 2000, the region
that Almorha [or Almora] is in, was renamed as the separate state of Uttarakhand). When
Menon says Kosi River at Almorha, he is not referring to the Kosi River system of Nepal,
and Bihar (India), he is referring to the Kosi [or Koshi] River that flows past Almorha in
the Kumaon [or Kumaun] region/division of Uttarakhand, and eventually joins the
Ganges. Based on this information, if the locality is correct, the specimen is likely to be
either a B. birdi or more likely a B. almorhae. It may even be that the specimen came
from the Sindh/Sind River (a tributary of the Yamuna River in Madhya Pradesh and Uttar
Pradesh in India – not to be confused with the Indus River which is also known by some
as the Sindh River, or the Sind River in the Kashmir Valley), and if so could represent a
B. almorhae as Menon (1993:23 & 33) gives the nearby and parallel Chambal River as a
known locality for B. almorhae. If the “Sind” specimen used by Hora does truly represent
a B. rostrata, then the locality given by Day must be incorrect, as Hora himself thought
possible. Hora said that in its long and pointed snout his B. dayi came close to B.
4
Rafique, (2000) discusses the previous connection of the Yamuna River of the Ganges system with that of
the Indus River system, but estimates are that this was around 4000 years ago.
rostrata. He used information sent to him by Mr Norman at the BMNH that said B.
rostrata could be differentiated from Day’s “Botia geto” by “the longer snout, longer
barbels on snout, and rather different type of colouration.” I have found that snout length
differs between sizes of specimens, but also by observing live specimens of several
species, that the genders also appear to have notably different sized / shaped snouts. Also,
barbel length may differ in different sized or gender specimens. Also, “colouration” will
differ based on level of preservation, gender, and also the known variability of these
species. Mr Norman went on to say “In rostrata the snout is distinctly longer than the
remaining part of the head, whereas in ‘geto’ it is about equal to or (generally) less than
this”. If Mr Norman was using the information from the Day specimen of ‘B. geto’
(which he didn’t have in his possession), he may be comparing B. rostrata to a B.
almorhae or B. birdi, based on what I have said above. However, I think that based on
how he phrased his last sentence Mr Norman was using specimens in the BMNH that are
logged as B. geto, and were donated by Day (Hora mentions their localities). If this is the
case, these are also flawed comparisons because I have found that the Day specimens of
B. geto in the BMNH actually represent B. almorhae and B. birdi, and perhaps sp. ‘Kosi’.
See discussion above about the possibility of B. dayi being synonymous with B. rostrata,
but that it is by no means definite.
Some specimens appear very similar to Botia kubotai Kottelat, 2004. My observations
would point to the different pattern when young (more bars in B. rostrata); and the
different zoogeography as pointers for classing them as different, and indeed I have put
them in different species complexes.
Most specimens in the hobby up until now that are called B. rostrata have been sp.
‘Upper Brahmaputra’ (see below). The accounts of B. rostrata in Kottelat & Chu (1987,
fig. 4 – specimens from Irrawaddy), Zhu (1993), Zhu (1995), Kuang, in Chu & Chen
(1990), Chen et al (1998), Vidthayanon et al (2005), Hoyer & Ott (2006) are in my
opinion the Irrawaddy form of Botia histrionica Blyth, 1860. In my opinion the
specimens known in the aquarium trade as ‘Botia sp. aff. rostrata’, or ‘ Botia cf.
histrionica’ are the Ataran River form of B. histrionica. The example (not figured) in
Kottelat & Chu (1987) from Salween basin is likely to be the Salween form of B.
histrionica.
Botia blythi Bleeker, 1863 nomen dubium
Botia grandis (nec Gray) McClelland, 1842
Syncrossus grandis Blyth, 1860
? Botia lohachata (nec Chaudhuri) Sen, 1995.
Meristics: McClelland gave them as “D 10, P 14, V 9, A 1/7, C19”
Bleeker gave the type locality as “Khasya mount[ains]”, based on McClelland’s account
from “Kasyah mountains” and Blyth’s use of that account as “Khásya hills”. All of these
localities are the same, which is the Khasi Hills of Meghalaya. It is not 100% clear if the
specimen that McClelland used was one collected by Griffith himself on his journey to
the Khasi Hills in 1837, or by one of his collectors who collected there in 1837-1840. I
have read the account of Griffith’s 1837 journey to the “Khasyah Mountains” (Griffith,
1847), and there is no specific mention of collecting the fish, but I consider that this is
primarily because his account was written from the point of view of mainly collecting
plants. It is made clear however that he sampled the parameters of the waterways he
visited. I hypothesise that he did collect the specimen in 1837. If you read the account of
his journey, it appears that he travelled from Dhaka, up the waters of the Meghna River,
then perhaps the Surma/Barak River, onto to “Moosmai” [Mawsmai], then to “Churra
Punjee” [Cherrapunjee/Cherapunji], then to “Moflong” [Maoflung], on towards
“Chillong” [Shillong], then returning south back towards Cherrapunjee and other
locations, then back north towards Assam. Looking at this route in the Khasi Hills region,
from Mawsmai, to Cherrapunjee, then to Shillong, (which closely matches a route he
took in 1835), it is possible that the specimen that would ultimately have B. blythi based
on it, came from the Shella River, as this river appears to lie along or near this course. If
it is not the Shella River, it will be one of the many tributaries of the Surma River. So far
the type specimen of B. blythi has not been located. I did wonder whether BMNH
1860.3.19.910 is the one that Griffith collected, and McClelland used, Blyth cited, and
then Bleeker based to describe B. blythi. It was deposited by the East India Company for
which McClelland worked. However, the fin ray counts don’t quite tie up, so this needs
further work.
Yazdani (1977 and 1985) did not list any Botia species from his survey of Khasi Hills.
However, clearly Sen (1995) did, and I know of definite catches there (Rao), so Botias
are present there.
In view of the fact that McClelland thought that the specimen he used represented Gray’s
Botia grandis (a junior synonym of Botia almorhae) it is possible that the specimen was
similar to B. almorhae in pattern and perhaps colour, and Sen (1995) has allegedly
encountered B. almorhae complex patterned juveniles in Khasi Hills. Because of this I
first thought that B. blythi may be a member of the B. almorhae complex. The
Brahmaputra and Ganges system in Bangladesh gives a hydrographic connection via the
Meghna and Surma rivers with other species of the B. almorhae complex. However, my
hypothesis was that it was a separate species from B. almorhae sensu stricto. The
topography and hydrography of the area and waters of the Surma River system compared
to those of B. almorhae, and the alleged differences in ventral fin ray count (McClelland,
and Sen, give ventral fin rays counts as 1/8 for B. blythi. A ventral fin ray count of 8
branched rays is outside the usual range for Botias; although it could be a miscount - see
under B. dario) were possible clues towards considering them as different species. When
I saw the photographs by Ken Childs of aquarium specimens of unknown locality I
thought they might represent B. blythi. These specimens possibly have the right colour
and pattern, and appear to have 8 branched ventral fin rays. Bleeker described B. blythi
by way of an indication. This means he referred to previous works (McClelland’s and
Blyth’s) which gave defining characters of the species (this was allowed by the ICZN for
descriptions prior to 1931 – Article 12). To see what the species looks like we therefore
have to go back to McClelland’s description in 1842 (because Blyth just repeated
McClelland’s words). McClelland said “Body and fins covered with irregular green spots
and streaks.” If Sen’s (1995) account is correct and reliable, it appears that the young of
this dubious species have the same pattern as B. almorhae and B. birdi. Sen used a
drawing of what he labelled as B. lohachata, but listed them as being from Khasi Hills
(probably Shella River?). Unfortunately the drawing that he used on figure 4 plate XXIV
to represent B. lohachata (= young B. blythi) is the one taken from Talwar & Jhingran
(1991) which was in turn taken from Sterba (1963), so although we can see the pattern, it
is not a drawing straight from Sen’s Khasi Hills specimens. So in essence he is stating
that the young, are very similar to young B. almorhae and B. birdi. I have also considered
if Botia sp. ‘Teesta’ is B. blythi as they have the Y and i pattern when young, and when
older have spots and streaks (see below). However, my thoughts changed when I received
information and images of Botia specimens from Khasi Hills, and specifically from the
Shella River system. These variable specimens match the drawing of Sen of B. rostrata in
Garo Hills, and also match the pattern in Günther for B. rostrata, and in Hora’s B. dayi
description. The larger specimens have anastomoses and disintegration, and in life (or
reasonably recently dead) and can have a greenish yellow tint to the spots and streaks.
The local fishermen told the collector that there were no differently patterned Botia that
they had seen, they were from the possible locality of B. blythi, and they have greenish
yellow spots and streaks. All this is evidence to possibly show that the specimen that B.
blythi is based on may be a B. rostrata, (although I cannot be certain at this stage that
Shella River B. rostrata are conspecific with Mahananda and/or Ganges B. rostrata).
However, as mentioned above there is counter information that B. blythi may be a
member of the B. almorhae complex, and may also have 8 branched ventral fin rays (the
fresh Shella specimens appear to have 6-7 total countable rays using naked eye). In view
of the conflicting information which I am unable to adequately resolve, I have decided to
list B. blythi as nomen dubium. If it is later proven that it is synonymous with B. rostrata,
the name B. blythi may not necessarily take precedence over B. rostrata (or B. nebulosa)
in view of Article 23.9 (my use of B. blythi in 2001 was in passing and I was not
identifying it with any taxon or claiming it to be valid).
Botia sp. ‘Upper Brahmaputra’
Botia dayi (nec Hora) Banarescu & Nalbant, 1968
Botia rostrata (nec Günther) Sen, 1985
Botia rostrata (nec Günther) Nath & Dey, 2000
Botia geto (nec Hamilton) Ott in Schmidt, 2002
Preserved specimens used: AMS I.31542.008, no locality data, presented 1991, 31mm
SL, identity uncertain.
Distribution: Brahmaputra system: Khwang River, near Khwang, Dibrugarh district,
Assam; Brahmaputra River near Guwahati town, Assam; Manas River, tributary of the
Brahmaputra, Assam, India (Rao, and Turner, personal communication); Dibrugarh,
Assam, India (Sen, 1985); Kaziranga, Mikir Hills, Assam (Banarescu & Nalbant, 1968);
Kameng, Subansiri, Dikrong, Pachin, Ranga, Siang, Dibang, Lohit, Noadhing, Buridhing,
Tirap rivers, Arunachal Pradesh, India (Nath & Dey, 2000). Therefore it is found in the
Brahmaputra River and its connected rivers, before the Brahmaputra becomes the
Jamuna, hence the name ‘Upper Brahmaputra’. The specimens that come into the trade
are usually collected at two different localities and this may account for the differences in
pattern (see images by Mark Duffill).
Meristics: Dii-iii 8-9; Pi 11-13; Vi 6-7; Ai-ii 5; C 19
Colour and pattern: Base colour is a beige, yellowish, or sand colour, usually with a
metallic sheen sometimes giving the pale spots a metallic gold colour. This species is
very variable, even in small specimens of the same size, but in small specimens the head
usually contains 3 bands (the first, which is sometimes broken, along the ridge of the
snout, to its tip; another which across the interorbital area is usually split in two,
connecting the eyes, then under the eye as one band; and another from the eye towards
the premaxillary), and the body 9-10 bars (albeit the first of these bars is predominantly
on the head and opercle). The bars are usually placed in pairs. In some specimens the bars
are almost fused together, but start to split to form separate bars as the fish grows.
However, even in specimens of the same size (5cm TL for example) and age and locality,
there is a difference in the outline of the bars, but also in the degree of disintegration.
Also, some specimens even at this small size have some or all of the bars disintegrated,
with the base colour showing through as spots, blotches and/or bars within the dark bars.
At this small (5 cm TL) size the bars also start to anastomose in some specimens,
whereas in some from the same catch the bars have not anastomosed. The bars and bands
are usually pale black to dark grey. When the bars are distinct, there is usually a bar over
the nape, which passes over the opercle. The next bar ends around the base of the pectoral
fin (mainly over the base of the rays). There are then two more similar bars. The next bar
usually has its vertical middle at the insertion of the leading ray of the dorsal fin. The
next bar has its vertical middle at the insertion of the base of the last dorsal fin ray(s); the
next 2 - 3 on the posterior half of the body, and the last being on the caudal peduncle and
base of the caudal fin rays. Some adults of 10cm SL + still have the bars showing but
anastomosed and disintegrated, whereas in some specimens of similar size the bars are
not discernable. The base of the dorsal fin usually has some markings where the fifth and
sixth dark body bar meet the fin, and there can be one crescentic stripe along the middle
to distal half of the rays. In semiadult to adults, there can be a black tip to the first 3 or 4
rays. In small to medium sized specimens, the pectoral fin usually has two dark bands;
one near or at the base, and one in the middle. In adults there are usually 3 to four. The
ventral and anal fins usually have two dark bands, one usually near the base and the other
in the distal half. The caudal fin usually has two to four dark vertical or crescentic bars
(the smaller the specimen, the fewer bars there are).
Discussion:
This is the fish that is sold as B. rostrata, in large numbers in the aquarium hobby. In its
pattern it is very similar to the true B. rostrata albeit it doesn’t appear to get as
anastomosed and disintegrated anteriorly as B. rostrata. In colour it is different. It doesn’t
have the bright base colours that can shimmer greenish yellow, or even bluish that can be
exhibited in true B. rostrata, in sp. ‘Upper Brahmaputra’ if the shimmer is present it is
like gold or copper. There are some minor differences in meristics also. It may be that
they are the same species and just be exhibiting differences due to water parameters or
habitat. Further study is required.
Rao (personal communication) states he has never encountered the adult “reticulated”
specimens in the wild, he has only seen them in aquaria. He thinks they could be found in
Arunachal Pradesh up in the hill streams as he states extra large B. almorhae complex
specimens are collected from dam sites in Nepal bordering India. There appears to be
some other evidence of this – see under B. lohachata. The largest specimen Nath & Dey
(2000) found in Arunachal Pradesh was 120mm and they comment on the variability of
the pattern (also they show a drawing and a photograph).
Botia geto – See information under B. dario, of which B. geto is the junior synonym. B.
sp. ‘Upper Brahmaputra’ is still sometimes sold as B. geto. I consider that this is due to
two reasons. The first is the locality. Hamilton described B. geto from the north eastern
parts of ‘Bengal’, but the type specimen was from Goalpara in Assam, and I have shown
that this species is caught in that region too. The other reason is due to the pattern and
colouration. The description and drawing in Hamilton could at first glance appear the
same as a young sp. ‘Upper Brahmaputra’. However, if you look at the drawing of B.
geto, the position, shape and angle of the head and body bars do not match that of any sp.
‘Upper Brahmaputra’ (or B. rostrata) that I have seen (and I have seen tanks full of live
young specimens of sp. ‘Upper Brahmaputra’), but do match that of young B. dario as I
have discussed above. Very small (under 31mm SL) sp. ‘Upper Brahmaputra’ are very
similar in pattern to similar sized B. dario, mainly because the bar on the caudal peduncle
in sp. ‘Upper Brahmaputra’ of this size is sometimes still fused with what becomes the
penultimate bar; however the position of the bars on the head and body is a factor which
differentiates them at this size. Also, when looking at specimens in life one can easily tell
them apart due to a combination of base and bar colour and sheen.
Possible hormone produced hybrid?
These are being sold in some aquarium outlets. We know that they are coming out of the
Czech Republic, and I think that they are hormone bred. My view is that they are a hybrid
between sp. ‘Upper Brahmaputra’and possibly B. dario and/or B. striata, although they
are being sold as B. almorhae (which they are not). In view of the confusion that already
exists about actual species, I consider that these (alleged) hybrids will only serve to
confuse matters further and should be frowned upon.
Botia almorhae Gray, 1831
Junior synonyms:
Botia grandis Gray, 1832
? Schistura maculata McClelland, 1839
Preserved specimens used: BMNH 2002.9.18.1 (holotype of B. almorhae), “Almorah,
Uttar Pradesh, northern India”, presented by General Hardwicke, 1 spec., 151mm SL;
BMNH 1889.2.1.1474-1476, “Delhi”, presented by F. Day, 2 spec., 29mm - 36mm SL;
BMNH 1889.2.1.1481, “Roorki” [Roorkee], presented by F. Day, 1 spec., 52mm SL;
BMNH 1889.2.1.1482-1491, “Jumna River”, presented by F. Day, 6 spec. of 7, c. 38mm
- 56mm SL; ANSP 79789, “Markunda River” [Markanda River], “India”, presented by
Rev. M Carleton, (1 spec. of 3), 34.1mm SL; ANSP 79786, same data as 79789, 43.4mm
SL; AMS B.7768, “Almorah”, presented by F. Day, 1 spec., 115mm SL; MCZ 4026,
“Gugger River and canal”, presented by Rev. M Carleton, 6 spec., c. 45mm – 92mm SL;
MCZ 4027, same details as MCZ 4026, 9 spec., c. 38mm – 58mm SL; MCZ 4028,
“Markunda River, washed into plains by storm”, presented by Rev. M Carleton, 3 spec.,
c. 36mm – 51mm SL; MCZ 4029, “Markunda River”, presented by Rev. M. Carleton, 2
spec., c. 32mm – 33mm SL.
Distribution: Ganges system in India: Almora (Kosi River, ?Sot River), Yamuna River,
Roorkee (Kosi River), Kosi River near Kashipur, Alaknanda River (Menon, 1993) Uttarakhand; Yamuna River (specimens, and Turner, personal communication) – Delhi;
Markanda River (a tributary of the Yamuna River) - Himachal Pradesh and/or Haryana;
Ghugger River – Himachal Pradesh; Kali River, Meerut and Kali River in Muzaffarnagar
district, and Bijnour district (Menon, 1993) – Uttar Pradesh; Jammarri Nadi, Gambhir
River at Sawai Madhapur, and Kota district, Chambal River drainage (Menon, 1993) –
Rajasthan.
Checks need to be made whether Chambal drainage specimens are conspecific with
topotypical B. almorhae.
As the holotype comes from Almora, it could have been either from the Sot River
(tributary of the Ganga River), or the Kosi (Kumaon) River.
Meristics: Div 9-11; Pi 13-15; Vi 7; Aiii 6; C 19. Further counts from B. almorhae sensu
stricto need to be made. Gray gave the counts of the holotype as “P 13. D. 11. V. 8. A. 7.
C. 25”; these appear to include all the rays, not just branched ones. Günther gave the
counts from the holotype and the East India Company one (which does not appear to be a
B. almorhae based on the shape of its bony anterior chamber of the swimbladder – it is
much narrower and bonier than the holotype of B. almorhae) as D12 [iii 9], A8 [iii 5], V8
[i 7]. Based on a radiograph of the holotype, the counts appear to be Div 10, Aiii 6, C19. I
have been unable to accurately count the pectoral rays and ventral rays on this specimen
due to its age and preservation, but it appears to be Pi 13, Vi 7. The AMS specimen
appears to have Div 11, C19. The dorsal fins of the MCZ specimens are either damaged
or would be if extended to count but there appears to be 9-10 branched rays.
Colour and pattern: When discussing the colour (and perhaps pattern) of this species, I
will here stick to the details for B. almorhae sensu stricto (see below). Gray described the
colour and pattern as “brownish with irregular pale spots; fins pale, three or four banded”.
The colour and pattern of B. almorhae is (in juveniles) is a yellowish base colour with
brown Y and i pattern on the body. 1 to 3 bands in the dorsal fin, number of bands in the
pectoral and caudal fins increase with the size of the specimen. In adults the body is
reticulated with brown, the yellow ground colour being broken up into yellow spots due
to anastomosis and disintegration, or in some specimens the remnants of the Y’s and i’s
can still be seen, some are anastomosed and some not, but all bars having disintegrated see original drawing of B. grandis; and the drawing in fig. 5 Plate CLIV in Day (1878) of
a specimen from “Almorah” (which based on information I have (Grant, 1999; Ferraris et
al, 2000) on other specimens at AMS, is probably a drawing of AMS B.7768); also the
drawings on Plate I and images on Plate II figs 1-3 of Menon (1993). Menon’s view is
that the males are the ones with full anastomosis, the females with the remnants of the
Ys. Based on images of live specimens of both genders (of unknown location), I consider
that this may not in fact be sexual dichromatism.
Discussion:
Gray described B. almorhae in 1831 based on a specimen in the BMNH. In 1832 he
published a drawing (reproduced here) of the same specimen and used the name B.
grandis, thus making B. grandis a junior synonym of B. almorhae.
This species has long been confused with two other described species: Botia lohachata
and B. birdi. The problem has been caused because juveniles of all three species exhibit
the Y and i pattern, and they come from nearby river systems. Thankfully we can more
easily class B. birdi as a separate species, and see under the account for B. birdi for a
discussion of the differences between these two species. The problem is more difficult for
B. lohachata, mainly because the river systems they were described from are
hydrographically connected, and appear to have been so for a considerable period of
time. Unfortunately I have not had access to freshly preserved topotypical specimens of
B. almorhae that would be needed to resolve this issue with more certainty. I have
however had access to specimens from the river system B. lohachata was described from,
and from nearby Himalayan systems. I have also studied the accounts of these species,
including their localities and appearance. I started firstly by comparing the appearance of
the specimens from the type locality of B. almorhae (see above) with that of specimens
from the river in which the type specimen of B. lohachata was found (Gandak River). As
mentioned above we know what B. almorhae sensu stricto looks like, but how does it
differ (if at all) from B. lohachata? The juvenile holotype of B. lohachata had black
markings, and white pectoral, ventral, and anal fins; but there was no mention of its base
colour. I then looked at any known specimens from Gandak River in India or Nepal. In
Menon (1993) on Plate 2 Figs. 7 and 8 he shows two males from the Kali Gandaki River
in Nepal (same river as the Gandak). These specimens appear to have a greenish or
greyish base colour (vs. yellowish) and the body bands more greyish green (vs.
brownish). I contacted Dr. David R. Edds, the collector of the Kali Gandaki specimens,
as he has done lots of collecting in that region. David kindly sent me images of live and
preserved specimens from the Kali Gandaki, the river system that the types of B.
lohachata were caught in. David also helped me access images of and information on
Nepalese specimens of the B. almorhae complex from the main river systems in Nepal
and northern India. The Kansas University kindly sent me quality images of their
specimens, but unfortunately could not send me x-rays in time for this publication. The
reason I asked for x-rays was that I have a feeling that despite Hora’s 1922 and 1935
comments that the bony swimbladder in B. almorhae is more encased in bone that in B.
lohachata, I have a feeling that if there is a difference between the two on this point it
will be B. lohachata that has a bonier anterior portion of the swimbladder. This is based
on the following:
•
•
•
The holotype of B. almorhae does not appear to have the anterior portion of the
swimbladder any bonier than or a different structure to almost all the Botia x-rays
I have seen.
BMNH 1860.3.19.910 does but this is different to B. almorhae holotype, and this
specimen could in fact be the nearby and similar B. lohachata.
If the structures are bonier than it would make sense for B. lohachata to be bonier
to give it the protection and different buoyancy needed in its waters of higher
altitude and rapidity.
See below for some morphometric comparisons between B. almorhae, B. birdi, B.
lohachata, possible B. lohachata, Birbhum (lower Ganges), and sp. ‘Kosi’.
Body Depth in SL
Snout Length in SL
B. almorhae holotype
BMNH Delhi (2)
BMNH Roorkee (1)
BMNH Jumna (7)
MCZ Ghugger (11)
MCZ Markanda (4)
4.1
3.6 – 4.2
4.4
3.5 – 4.5
3.4 – 4.6
3.8 – 4.1
8.1 – 8.6
7.6 – 7.8
9.9
7.2 – 10.8
5.9 – 8.9
7.2 – 8.8
BMNH & CAS B. birdi (7)
BMNH Tawi River (1)
3.6 – 4.7
3.2
6.8 – 8.4
8.4
OSUS & KU Gandaki / Narayani B. lohachata (20) 3.6 – 6.1
KU Karnali / Ghaghra drainage, Ghomti (27)
3.6 – 5.5
5.4 – 7.7
5.7 – 8.3
KU sp. ‘Kosi’ (7)
BMNH Birbhum (3)
4.0 – 6.3
4.1 – 5.0
6.5 – 9.1
8.0 – 10.5
As you can see from the information above, based on the information I have, B.
lohachata appear to have specimens with smaller proportionate body depth and the
snouts are proportionately longer than B. almorhae.
Base and bar colour, distribution, and some morphometrics may not be deemed by some
to be differences to justify separate species, but I do consider that they are different (at
least enough for subspecies rank) and have listed them as valid species in their own right.
Further work is needed on the structure of the swimbladder, and if this proves
inconclusive, mitochondrial DNA analysis.
Botia lohachata Chaudhuri, 1912
Botia almorhae (nec Gray) Shrestha, 1994.
Botia almorhae (nec Gray) Shrestha in Petr & Swar, 2002.
Preserved specimens used: OSUS, Kali Gandaki River drainage, Nepal, collected by
David R. Edds, 9 spec., c. 58mm – 130mm SL; KU 29019, Kali Gandaki River at
Sumsaa, Nepal, collected by David R. Edds, 2 spec., 72mm – 79mm SL; KU 28879,
Narayani River, Downstream from Tribeni barrage at Pier No. 6, Nepal, collected by
David R. Edds, 1 spec., 35mm SL; KU 28633, Bagmati River, at Raj-Marg highway,
Nepal, collected by David R. Edds, 2 spec., 40mm – 41mm SL; KU 28867, Narayani
River at Narsai, Pier No. 12, Nepal, collected by David R. Edds, 8 spec., 42mm – 49mm
SL; KU 28858, Narayani River, just downstream from Tribeni barrage, Nepal, collected
by David R. Edds, 3 spec., 40mm – 51mm SL.
? - KU 28700, 3km W of Pipariya, Shuklaa Phataa Wildlife Reserve, Nepal, collected by
David R. Edds, 2 spec., 50mm – 57mm SL; KU 28723, At Raj-Marg highway,
9km E of Mahendranagar, Nepal, collected by David R. Edds, 5 spec., 48mm – 59mm
SL; KU 28754, At Chisapani, Nepal, collected by David R. Edds, 1 spec, 43mm SL; KU
28787, Five locations along about 25km stretch between Chisapani and
Kotiyaghat, Nepal, collected by David R. Edds, 10 spec., 44mm – 51mm SL; KU 28800,
Just downstream from irrigation project along Raj-Marg highway, Nepal, 8 spec., 44mm
– 65mm SL; KU 29085, At Khairenitar, Nepal, collected by David R. Edds, 1 spec.,
101mm SL; KU 29435, Just east of Katasi, Nepal, collected by David R. Edds, 2 spec.,
60mm – 67mm SL; KU 29457, At Gorangi, about 4km West of
Chisapani, Nepal, collected by David R. Edds, 1 spec., 55mm SL.
BMNH 1860.3.19.910, no locality registered at BMNH, East India Company Collection,
1 spec., 135mm SL. Identity uncertain.
Distribution: Ganges system in India - Chhoti Gandak (Srivastava, 2004), Gandak River
at Saran (Chaudhuri, 1912), Jamwari River, Saran (Menon, 1993), Gandak River near
Muzaffarpur (Rao, personal communication), Gandak River at Hazipur, Patna District
(Venkateswarlu, 1976), Ganges River near Patna? (Rao personal communication),
Balkhau River near Anhar District, and Shalkia River, Chapra? (Menon, 1993) – Bihar.
Rapti River, Gorakhpur (Menon, 1993), River Rohini, Gorakhpur (Srivastava, 1998) –
Uttar Pradesh, India.
Ganges drainage system in Nepal: Kali Gandaki River, Narayani River (two names of
same river), Kathmandu (Menon, 1993); Bheri (Rapti River drainage), Seti (Karnali /
Ghaghra River drainage), Janakpur (Bagmati River drainage), Narayani (Gandak River
drainage) zones (Shrestha, 1994); Narayani River at Sivaghat, Kharkhareghat, Koleghat;
Khoriyamohan and Deoghat and Narayanghat, and also Irrigation canal of eastern
Chitwan (Dhital & Jha, in Petr & Swar, 2002). Bardia National Park, and Shukla Phanta
Wildlife Reserve (Edds, 2007). Also see specimens listed above. The specimens listed
with a question mark and those from literature reported from the Seti (Karnali / Ghaghra
drainage) and Ghomti system need further study to see if they are conspecific with
topotypical B. lohachata.
It appears that the young inhabit the lower to middle courses of the rivers, and therefore
the lower to mid altitudes and the adults migrate to higher altitudes in the Himalayas, but
not to the highest (Shrestha, 1994; Menon 1993; Edds, 1993; Dhital & Jha, 2002; Edds,
2007).
Meristics: Di-ii 9, Pi 13-14, Vi 7, Ai-ii 5, C19 (Chaudhuri gives 20, which is one ray
more than normal so this may be a miscount).
Colour and pattern: See image by Dr David R. Edds of live freshly caught specimens. In
life when young they can exhibit a golden to silver sheen, when adults the base colour
can be greyish to greenish but the metallic sheen tends to fade. There are normally 3 Y’s
and some i’s and a bar at the caudal peduncle. The bars tend to have metallic sheen in the
young, and are dark greyish; when older they can be greyish to greenish. The bars can
disintegrate, fragment, and anastamose in varying degrees regardless of size of the
specimen. Some keep unconnected Y’s and i’s even when adult, whereas some adults do
not. The dorsal fin usually has 3 bands, sometimes 2, the first usually at the base of the
rays, the second near the middle, and the third near the tips of the rays. The caudal fin
usually has 3 to 4 diagonal bands; the anal fin usually 1 band at the base and sometimes
another in the distal half and dark markings at the tip; the ventral fins may have 1 to 4
bands (usually more as the specimen grows), the pectoral fins around 3.
Discussion:
This species was described on the basis of just one specimen, which is in the ZSI in India.
The specimen was a juvenile (measures “40mm”) and was from the Gandak River, in
Saran, Bihar. Because of this, the validity of this species has been in a state of flux since
its description. Some authors have classed it as valid (notably Shrestha, and Edds) and
others a junior synonym of B. almorhae (notably Menon).
Ng 2006 and Ng & Edds 2005 have shown that specimens of a catfish species that occur
in some of these rivers are actually distinct from other species in connected rivers. See
discussion above under B. almorhae. I do consider that B. lohachata is valid but further
study is needed to be definite.
As mentioned above, the specimens and records (including Edds, 2007) from the Seti
(Karnali / Ghaghra drainage) and Ghomti system need further study to see if they are
conspecific with topotypical B. lohachata. I have tried to look for any obvious recurring
differences in pattern in terms of degree of disintegration or fragmentation, or markings
in finnage but I have been unable to notice any on the specimens used.
Botia sp. ‘Kosi’
Botia birdi (nec Chaudhuri) Tekriwal & Rao, 1999.
? Botia lohachata (nec Chaudhuri) Ataur Rahman, 2005.
Preserved specimens used: KU 29574, Koshi River at Koshi barrage, Nepal, collected by
David R. Edds, 2 spec., 49mm – 75mm SL; KU 28539, Koshi River purchased at Koshi
barrage, Nepal, collected by David R. Edds, 3 spec., 43mm – 58mm SL; KU 29099,
Koshi River, purchased at Koshi barrage, Nepal, collected by David R. Edds, 3 spec.,
47mm – 59mm SL; AMS I.31134.-111, no locality data (aquarium specimen), presented
by N. Halliwell, 1 spec., 95mm SL (identity uncertain).
BMNH 1889.2.1.1477-1480, “Bheer-Bhoon” [Birbhum?], presented by F. Day, 3 spec.,
c. 39mm SL - 45mm SL (identity uncertain).
Distribution: Ganges system in India and Nepal: Kosi River, Nepal (KU specimens
above); Kosi River near border with India (Rao, personal communication) – Nepal;
Birbhum?, India. Hooghly River near Calcutta [Kolkata]? (Rao, personal communication)
- West Bengal, India. Kosi zone (Kosi or Sapt Kosi River drainage)? - Nepal (Shrestha,
1994).Ganges system in Bangladesh?: Faridpur and Dinajpur districts (Ataur Rahman,
2005).
Ng & Edds (2005) found that species of catfish that inhabited the river systems in Nepal
that were once thought to be one species were in fact two species. The specimens in the
Kosi system differed to those in the Narayani and Seti and Ghomti systems. This appears
to be the case too in Botias. The Kosi (possibly including Birbhum and Hooghly River –
although these may be different and a species in their own right) specimens appear to
have more yellowish to whiter base colour than B. almorhae and B. lohachata. The Kosi
specimens also appear to have the body not as deep in proportion to SL.
The specimen labelled as B. birdi in Tekriwal & Rao was not from Punjab as captioned, it
was from the Kosi River (Rao, personal communication).
Botia sp. ‘Teesta’
?Botia blythi
Botia almorhae (nec Gray) Ng, 2007 (in part).
Brahmaputra system: Tista (or Teesta) River Barrage, Gajoldoba, near Jalpaiguri, West
Bengal, India (Ng, and Rao, personal communications), and Ng (2007).
Meristics: From the images of specimens approximately 50mm SL and 60-70mm SL, the
following merisitics can be gleaned: Dii-iii 10, Pi 10, Vi 6, Aii 5, C 19. These specimens
are probably two from lot UMMZ 244844 as listed in Ng (2007).
Discussion:
This species has a Y and i pattern when young. Based on the origin of this species
(Brahmaputra system, rather than the Ganges system) and that its colour and particular
pattern is distinct from B. almorhae sensu stricto, I consider that this is a species in its
own right. The images by Heok Hee Ng are specimens from the Gojoldoba locality. The
images by Bogdan Janiczak are of unknown origin, but they appear to be adults of the
same species, based on images that Bogdan has provided of them at smaller sizes. As
discussed under B. rostrata, there is a possibility that sp. ‘Teesta’ is the species described
as B. blythi. Ng (2006) listed specimens of the catfish genus Pseudecheneis from the
upper regions of the Teesta River drainage as conspecific with those from Nong Priang
stream near Cherrapunji in Khasi Hills. It is therefore possible that the specimens listed
as B. lohachata by Sen (1995) are sp. ‘Teesta’ = B. blythi. Again, in view of the
confusion I list B. blythi as a nomen dubium, until this issue is resolved.
Botia birdi Chaudhuri, 1909
Junior synonym: Botia javedi Mirza & Syed, 1995
Botia lohachata (nec Chaudhuri) Yazdani, 1980.
Botia rostrata (nec Günther) Butt, 1992.
Preserved specimens used: BMNH 1956.4.12.1-2, “Chattabal, Srinagar, Kashmir”,
presented by Mr A. Datta, 2 spec., 74mm – 90.5mm SL; BMNH 1956.4.12.3-4, “Thal
Bel Nalla, Srinagar, Kashmir”, presented by Mr A. Datta, 2 spec., 86mm – 87mm SL;
BMNH 1889.2.1.1472-1473, “Jove River”, presented by F. Day, 2 spec., c. 40mm –
45mm SL; ANSP 97877, “Cashmere” [Kashmir], presented by Dr. F. Hallberg, 1 spec.,
51mm SL; CAS/SU 41285, “Srinagar, Kashmir, India”, presented to Dr. Herre, 5 spec., c.
76mm - 122mm SL.
BMNH 1983.7.11.30, “Taui [Tawi] River, Jammu, India”, presented by Jammu
University, 1 spec. c. 123mm SL – identity uncertain. This is logged as Botia dayi but is
not that species. I have included it under the account of B. birdi because of its locality
data, but its head and body profile seem to differ from the other specimens. When viewed
on x-ray the bony anterior portion of the swimbladder appears different, and the
dorsolateral profile of the skull appears different to the Jhelum river drainage specimens
in BMNH. It also has the long ‘hooked’ anal fin similar to the holotype of B. javedi. For
information purposes it has 29 vertebrae.
Distribution: Indus river system in India: Chattabal, Thal Bel Nalla, Dal Lake, and two
other sites, all Jhelum River or drainage, Kashmir (specimens above, Kullander et al,
1999, Menon, 1993); Nakera Khud, (Beas River?), 10km from Dehragopipur, Kangra,
Himachal Pradesh (Menon, 1993); Sirhind Circle (Ambala), at Rupar, where the Sirhind
Canal diverges from the Sutlej River, Punjab (Chaudhuri, 1909).
Tentative: Tawi River, tributary of Chenab River, Jammu (specimen above).
Indus river system in Pakistan: Michni, about 24km north of Peshawar, Kabul River
drainage, North West Frontier Province (Mirza & Syed, 1995); Jove River5, near Dera
5
The BMNH has numerous Day specimens of different genera with the locality “Jove River”. I was unable
to trace this river but a scholar in the Punjab region of India informs me that there is a Jove River near Dera
Ismail Khan in Pakistan. This would make it a tributary or distributary of the Indus River. All the non Botia
Ismail Khan, North West Frontier province (specimens above); (Ravi River?), Lahore,
Punjab (Menon, 1993); Jinnah Barrage area, Punjab (Rafique et al, 2003); Vale of
Peshawar (Rafique, 2000); (Sutlej River?), Ferozepore, Punjab (Menon, 1993); (Beas
River?) Dasuya [Dasua], Punjab (Menon, 1993); Baluchistan (Menon, 1993) and
(Rafique, 2000); Sangur [Sanghar], Sindh (Menon, 1993).
In view of the fact that B. almorhae or B. lohachata are not found in Pakistan, but B.
birdi is, the common name of Pakistani Loach should not be used for B. almorhae or B.
lohachata.
Meristics: Di-iv 8-11; Pi 13; Vi 6-7; Ai-iii 5-6; C 19
Colour and pattern: In juveniles they have the Y and i pattern of brown to blackish bars
on a yellow to beige base colour. Adults have a yellowish white lower half of the body,
the dorsal half being deeper and darker. The brownish black bars stop short of the ventral
surface and sometimes anastomose and/or disintegrate, and in large specimens (12cm SL
+) the bars can be absent. Dorsal (with 2-3) and caudal (with 3 to 4) fins have bands of
broad black or brown, on a yellowish white base. The pectoral, ventral, and anal fins can
have no markings, or can have around 2-3 bands. Snout can be reddish.
Discussion:
Botia javedi – described on the basis of 1 specimen (155mm SL) from Michni (about
24km north of Peshawar), Kabul river drainage, Northwest Frontier Province, Pakistan.
The anal fin on the holotype of B. javedi has a long ‘hooked’ anal fin, which the Tawi
River specimen also has, and also the holotype of B. almorhae has this too. Mirza & Syed
differentiated B. javedi from other species of Botia by its “peculiar body form, body
depth and the colour pattern.” When they refer to the ‘peculiar’ body form, it appears that
they are referring to the “concavity” of the head profile (predominantly the dorsolateral
profile of it, but also the ventrolateral profile). The specimen appears to be a large,
possibly gravid, female. The head profile is similar to smaller (but also gravid females) of
B. birdi from Jhelum River in the Kashmir Valley (Figs 5 and 6, Plate 7 in Menon, 1993),
although it is different to the (possible female) specimen from Tawi River in the BMNH
(tentatively identified). Hora (1922) said that in “ripe” B. birdi females the ventral profile
is greatly arched. I do not give much weight to the difference in head profile. In terms of
the body depth they state that the fact that the body depth is 31.6% of the SL is a
diagnostic character, but in smaller but deep bodied specimens that Menon studied, the
nearest percentage is 30.26%. I do not consider that a difference of 1.34%, when the
specimen is 40mm larger, is of any value when differentiating species. Hora (1922) said
that in ripe B. birdi females the depth of the body is considerably greater than the length
of the head. The differences in colour pattern are that B. javedi has the body devoid of
any bars, and its pectoral, ventral, and anal fin have markings. In an obviously adult and
old specimen the body bars could easily have faded. Although Chaudhuri’s B. birdi type
specimens did not have markings on the pectoral, anal, and ventral fins, other specimens
(BMNH and smaller CAS/SU) from Srinagar do. Having said this, based on the apparent
differences in the B. almorhae complex across its large range, it could be that all the
specimens in the whole Indus system are not all the same species (B. birdi). If this was
species names of the BMNH Jove River specimens are also either found in Pakistan, or fishes of the same
or similar genus are. Therefore I have tentatively used the BMNH Botia specimens as B. birdi.
the case the Jhelum, Chenab, Sutlej, Ravi, Beas, (?Tawi) rivers specimens may represent
B. birdi, and the Indus River (inc. Kabul River drainage and Jove river) specimens, B.
javedi. However, there is not enough information at this stage to support this, so for the
present time, B. javedi should be considered a junior synonym of B. birdi.
B. birdi is very similar to B. almorhae but since its description it has been considered as a
distinct species, but without any specific reasoning as to why. I also consider it valid and
distinct from B. almorhae. At this present time it appears that the Indus system is not
naturally connected to the Ganges system. However, it is hypothesised that the systems
have been connected in the past (around 4000 years ago) via the Yamuna River (see
Rafique, 2000:321-322) and it is possible that this is the reason that B. birdi and B.
almorhae are similar, despite occupying currently separate major river systems. I have
compared x-rays of B. birdi specimens (BMNH 1956.4.12.1-2, 1956.4.12.3-4, and
CAS/SU 41285) to all the BMNH specimens listed under my account of B. almorhae.
The 5 B. almorhae specimens have 27-30 vertebrae, the 9 B. birdi 30-31. I have also
compared the shape of the bony anterior chamber of the swimbladder on both species and
cannot identify any notable differences.
Menon used gave some characters to separate B. birdi from B. almorhae, notably that in
B. almorhae the ventral fins when laid flat reach or pass over the anal opening (vs. not
reaching), and the anal fin when laid flat almost reaches the caudal base (vs. not
reaching). If you look at the photographs of some of the specimens he has used, this does
appear to be the case. I have looked at the specimens of each species that I have used and
found that in almost all the B. almorhae the ventral fins do reach the anal opening (only
just though on the holotype, and 4 of the Ghugger River MCZ specimens don’t) and not
all of them have the caudal reaching or nearly to the base of the caudal (11 of the MCZ
specimens). In the B. birdi specimens (not including the Tawi river specimen), 5 (the
smallest) of the 12 have the ventral fins reaching the anal opening, and 5 (mixed sizes) of
the 12 have the anal fin reaching or almost reaching the caudal base. In his diagnosis for
each of the species, he also states that B. almorhae has a comparatively long ventral fin
and B. birdi a short ventral fin by expressing it as a proportion of SL. I have done the
same measurement in the adults of specimens of each species I have had access to and
found that the ranges overlap much more than the ranges given by Menon. Therefore as a
character to differentiate B. almorhae between B. birdi I consider that these characters are
unreliable. The morphometric and meristic figures I have (albeit limited due to numbers
of specimens) don’t appear to show any major differences either. This just leaves colour,
pattern and distribution. I have already discussed distribution; the colours and range of
disintegration in the pattern do appear subtly different but whether distribution, vertebral
counts, and subtle colour and pattern differences will be enough remains to be seen. Until
DNA analysis is carried out I consider that there is enough to render B. birdi as a distinct
species from B. almorhae.
The pictures by Bogdan have been used as B. birdi because of their colour and pattern.
However, because we do not know their locality, I cannot be certain that they are B. birdi.
Botia histrionica Blyth, 1860
As well as the ‘true’ B. histrionica, I consider there to be at least 3 (possibly 5) similar
but separate forms6, which I consider may (some of them) be described as new species in
the future. Having said this, the differences may just be down to the different conditions
in their habitat, such as water turbidity, substrate etc (Udomritthiruj, personal
communication), or that there is variability within each population or form to the degree
that no consistent differences can be noted. It may also be that each different form may
not just be restricted to one system, even if they are a different and new species (this was
the case for Pangio loaches as revealed recently by Britz & Maclaine, 2007). In view of
ongoing work by others on Myanmarian Botia and lack of access to many preserved
specimens, I will not cover this species, or the following ones, in as much detail.
So which is the ‘true’ B. histrionica? Blyth described B. histrionica from “Tenasserim”.
Kottelat (1989) and Ng & Kottelat (2005 & 2007) prefer to keep Blyth’s ‘Tenasserim’
distinct, as an historic collection locality, from the current Tanintharyi division, which is
the new name for just a part of the historic ‘Tenasserim’ region. Kottelat & Ng (2007)
appear to view Blyth’s Tenasserim locality as the Sittang River drainage, or the drainages
to the south of it (lower Salween basin etc), apparently based on most of Blyth’s
‘Tenasserim’ species being known to be caught there (Kottelat, 1989: 35). For some of
his species, Blyth gave the locality as “Tenasserim Provinces” rather than just
“Tenasserim” as he did for B. histrionica. One of the species (a Pangio) he described
from Tenasserim Provinces has actually been found in the Sittang, Salween and Ataran,
but it was noted that the Sittang specimens had much fewer vertebrae than the Salween
and Ataran ones, so it may be shown later that they are different (Britz & Maclaine,
2007). See Kottelat (2004: 7) for a discussion on the type locality, and on the possibility
of two forms, one from the Salween basin, and one from the Irrawaddy basin. My first
instinct was that if Blyth’s holotype did indeed come from ‘Tenasserim’, rather than Pegu
[Bago] as mentioned by Menon (1993) then it is likely to have come from the Salween
basin, rather than the Irrawaddy. If this were the case, this would mean that figures 3 a-c
in Kottelat (2004) would represent the true B. histrionica, and figure 3 d, another species
or sub species. The description of the pattern in Blyth is:
“Only five black bands on the body, the first of which encloses the gill covers and the
third descends from base of dorsal; another crosses the forehead and eyes; another, again,
passes from before each eye to the cleft of the mouth; and the medial portion of the face
is also black to the muzzle. The bands of the body are broad and subregular in shape,
each containing a pale round spot at the lateral line and another on the ridge of the back.
Dorsal fin with one broad interrupted black band, and some black also at base. Pectorals,
ventrals, and anal, each with two black bands; and the caudal also with two broad bands
and a black tip to each lobe.” The length was stated as 4 ½ inches.
This pattern seems to correspond with that of the fish in Kottelat’s figure 3 a-c, in that a
pale spot in the body bars is exhibited in the middle of the bar, and at the top. As the fish
grows the bars fragment. However, the title and introduction of the paper that Blyth
described B. histrionica in, clearly states that he was predominantly describing specimens
from the Sittang River and its tributaries, which would back up Day’s and Menon’s
6
In view of the ongoing work on Myanmarian Botia I have decided to use the term ‘form’, instead of
giving them a sp. name as in the species above. However, I am of the opinion that at least some of them
may be different species.
statements of Pegu (which if true would possibly mean the specimen was from the Pegu
(Bago) River, part of the Sittang system). The Sittang River is located in-between the
courses of the Irrawaddy and the Salween, and as mentioned previously it is theorised
that in the distant past that the lower course of the Irrawaddy occupied the valley. I have
access to images of live specimens of unknown location that when adult, are somewhat
intermediate between Irrawaddy and Salween B. histrionica. In adults they have the
whiter/greyer base colour of Salween form, but have a greyish to green tint to it, and the
disintegration of the bars is either absent or hardly noticeable (albeit each specimen
varies), and the bars are not split and only slightly anastomosed; but they don’t have the
base colour or anastomosis of the bars to the extent of Irrawaddy form either. These
specimens look similar to the deep bodied and cleanly barred and only slightly
disintegrated specimen in Day (1878 & 1889). Day said this specimen “came from Pegu,
and is in the Calcutta Museum” and stated that the base colour was olive. Perhaps these
live specimens are what adult Sittang River B. histrionica look like? I know for definite
that B. histrionica forms are present in the Sittang River and its tributaries/distributaries
as I have been told of preserved specimens from a tributary of the Sittang but have not
been allowed further information on them at this stage.
The base colour of Irrawaddy and Salween forms are not olive, whereas the base colour
of the adult mentioned above does have a greenish tinge. Kottelat theorises that the
specimen mentioned by Day may be the holotype of B. histrionica and if this is the case
this too poses some questions as to whether the Salween form is the true B. histrionica,
especially when one bears in mind that at 4 ½ inches some of the specimens theorised
here to come from the Sittang also have pale spots in the middle line and on the ridge of
the back.
Fig. 28 in Rendahl (1948), of a specimen from Rangoon [Yangon], possibly shows the
Sittang form as it could have come from the Pegu River, rather than the Irrawaddy system
(Myitmaka River), and it doesn’t appear to have the dark spot that tends to develop inbetween each dark bar in specimens of that size from the Irrawaddy.
We cannot be certain of the definite locality of the holotype of B. histrionica, but for the
time being my view is that we should accept the Sittang form as being the true B.
histrionica. It should not be ruled out however that the Salween form could be the ‘true’
B. histrionica.
Salween form
Fig. 666 in Zhu (1995) is very likely to be the Salween form.
Specimen KIZ 737003 (in Kottelat & Chu, 1987) from Nanka River (Salween basin),
Ximeng County, Yunnan, China is likely to be the Salween form (same locality is listed
in Kuang, in Chu & Chen, 1990).
The drawing in Chen & Zhang, in Chen (1998: fig. 51) is possibly the Salween form.
Irrawaddy basin form
If one looks at some specimens from the Irrawaddy basin and compares them with
Salween basin specimens, one can immediately see a difference in the pattern, but also
sometimes in the snout of some specimens (but possibly a sexual difference). The larger
specimens from the Chinese headwaters of the Irrawaddy seem particularly different. In
live Irrawaddy River B. histrionica, even in adults the base colour is more yellowish than
that of the usually paler, usually whiter, Salween form. As well as the specimen in
Kottelat’s fig. 3 d (and in Kottelat & Chu, 1987 – tentatively identified as B. rostrata, but
later classed as B. histrionica Irrawaddy basin by Kottelat, 2004), Zhu (1993) shows a
specimen (Number 108) from either the Irrawaddy River or Nujiang River (based on the
Nujiang River becoming the Salween, I hypothesize that the specimen is from the
Irrawaddy River). Fig. 667 in Zhu, 1995 (labelled as B. rostrata) will also probably be
the Irrawaddy form. The specimen (fig. 68) incorrectly labeled as Botia rostrata in
Kuang, in Chu & Chen (1990) also appears to show the Irrawaddy basin form, and is
obviously the same drawing as fig.4 in Kottelat & Chu (1987), which shows an adult
specimen from Daying Jiang River, Irrawaddy basin, Tuantian, Tengchong County,
Yunnan, China. The specimen in Kuang labelled as B. histrionica (fig. 67) appears to be
a juvenile, and an adult being labeled as B. rostrata. The distribution map for the “Botia
histrionica” specimens shows them coming from the Daying Jiang River (which is the
Irrawaddy), and/or the Longchuan Jiang River, which drains into the Irrawaddy. The
small specimen figured (70mm SL) shows the solid body bars, with no spots forming
within them as in Irrawaddy B. histrionica. The adult (135mm SL) also matches that of
fig. 3 d in Kottelat (2004). The body bars seem to sometimes fragment to become a
pattern of vertical bars of dark spots or small bars, then wavy bars.
Specimens CAS 88887 (2, from Myitkyina market, Kachin, Myanmar), and 88808 (1,
Mandalay fish market, Myanmar) appear to represent this form. Again, the body bars
show no pale spots within them, although they do fragment by branching off and forming
another smaller bar or spot.
The specimens captioned as Botia rostrata in Abbs. 1-3 in Hoyer & Ott (2006) were
caught in the Irrawaddy River (possibly near Mandalay). Thanks to Hoyer & Ott I have
been given permission to reproduce the images. The first specimen is approx. 73.5mm
SL. The middle body bar shows the typical spot at the top (forming the ‘Y’ shape) under
the base of the dorsal fin, and the bar before it also shows the spot where the bar similarly
turns into a ‘Y’ shape. However, there are no pale spots in the middle of the bars. The
caudal fin has 2 bands, the pectoral 1. Specimen 2 is approx. 52mm SL. It shows solid
pattern in the bars with no spots. Contrast this with the specimen of B. histrionica in
Kottelat’s fig. 3a, which has spots forming at the middle and top of the three central body
bars. The specimen has 1 band in the caudal fin, and 1 in the pectoral fin. Specimen 3
measures approx. 79.5mm SL. The body bars have gone thinner, the middle one
disintegrating. The caudal has 3 bands, the pectoral 2. As each of the three specimens
differs in size, the placement of the middle dorsal fin band becomes more central.
The specimen they caption as B. histrionica in Abb. 9 was caught at in the Irrawaddy
near Cain Kite Kwin, which is between Pyay to Pathein (not between Bagan and Pyay as
mentioned in the article – Hoyer, personal communication). The specimen was approx.
8cm TL, and Abb. 10 showed the locality 2 years after the specimen was caught. This
again shows the solid bar pattern, with just the middle body bar showing the ‘Y’ at the
base of the dorsal fin.
Vidthayanon et al (2005) shows on page 83 a specimen allegedly from Irrawaddy basin
and on page 81 incorrectly lists as B. rostrata, what is allegedly a specimen from
Salween or Tenasserim basin.
Manipur form(s)
Hora (1921) reported specimens from Manipur, India, and Menon figured 2 large
specimens from Manipur. In Menon’s figured specimens, apart from the spot at the top of
the central body bar (which tends to show in most specimens), the bars have no spots, and
have not disintegrated. Hora recorded 2 specimens from Amambi stream of 121mm and
93mm SL. In the larger specimen he states that there are black dots between the vertical
body bars. At first I thought that these specimens may belong to the Irrawaddy form in
view of the spots inbetween the bars reported by Hora and the hydrographic connection
of some waters of Manipur, with that of the Irrawaddy, via the Chindwin River of
Myanmar. However, upon looking at Vishwanath (2002) my view changed. Fig. 89 in
Vishwanath shows a juvenile or sub adult pattern that is more similar to Sittang form than
Irrawaddy form. Plate I fig. 13 shows an adult (10.5cm SL) Manipur form. Vishwanath
states this is maximum size for adults that so far he has only found in the Chindwin
drainage at lower altitude. He gives the colour and pattern description as “Yellowish
ochre with 5 dark vertical bands on body and 2 in head. Each band broken up into 2 in
larger specimens. Fins with 2 broad brown bars.” In the adult the pattern is more similar
to Salween form, but the base colour is more similar to Irrawaddy form. Some of the
body bars have disintegrated near the ventral edge, in the middle, and along the ridge of
the back, one has split to almost become 2 bands. There are no spots or marks inbetween
the bands, which are normally present in similar sized Irrawaddy form. The patterning on
the head is different to the other forms. It may be that specimens from the Imphal River,
Manipur River and its tributaries are different to the more easterly Chindwin drainages.
More study is needed.
The record (not backed up by any specimens) of Sen (1995) of B. histrionica in Garo
Hills, Meghalaya, India, is likely to be in error. Sen (1985) did not report any B.
histrionica from Assam.
Ataran form
This form is caught in the Ataran River, along with Botia kubotai.
Kamphol discovered this species some years back. According to Kamphol the specimens
are always larger (up to 125mm), and deeper bodied than the B. kubotai he catches from
the same river. The bars disintegrate and anastomose, but do not do so to the same degree
as B. kubotai, but are more variable and anastomosed and disintegrated than that of
Salween B. histrionica. In this regard they are somewhat intermediate between Salween
B. histrionica, and B. kubotai. At first I queried if these were females, the ‘male’ being
kubotai, but this does not appear to be the case. This form has sometimes been given the
name ‘Botia sp. aff. rostrata’, or ‘ Botia cf. histrionica’ in the aquatic trade.
Hormone produced hybrid?
? Botia sp. ‘lineomaculata’ Baensch & Evers, 2002
Botia sp. ‘pseudorostrata’ Ott in Schmidt, 2002
The following fish are being sold in some aquarium outlets. We know that they are
coming out of the Czech Republic, and I think that they are hormone bred. My view is
that they are a cross between Chromobotia macracanthus (Bleeker, 1852) and a form of
B. histrionica. Baensch & Evers (2002) give a locality of the Mekong River in Laos.
They are pictured in Baensch & Evers and in Ott in Schmidt (2002) by Dr. Alexander
Kochetov, who has in the past described a species of catfish which was based on the
results of hormone breeding (Grant, 2003).
Because of the different forms (which are possibly different species), aquarists can be
quite confused when looking at specimens and trying to identify the species. It has led to
some thinking they have a young B. kubotai rather than a B. histrionica because of the
markings, but in fact may have one of the other forms of B. histrionica. Because of this,
when trying to determine the identity of (particularly) young specimens, one should try
and find out their origin: are they from Irrawaddy River system, Sittang River, Salween
River system, Manipur, or Ataran River? Are they actually from wild stock at all (if not,
then they are possibly the hybrid). The appearance of hybrids can only serve to confuse
(for aquarists) what is already an incomplete picture. From my observations of live
specimens, and all the collated information, B. kubotai appear to have a higher degree of
disintegration at smaller sizes (see fig 1 in Kottelat, 2004) than that of the forms of B.
histrionica.
Botia udomritthiruji Ng, 2007
Preserved specimens used: BMNH 1992.11.16.12, Tenasserim River, Myanmar, 1 spec. –
Manuscript type specimen of Tyson R. Roberts.
In view of the recent description it is not necessary to describe the pattern in any detail.
However, as you can see from Kamphol’s images, very small specimens of this species
start off with distinct dark bars on the head and body. As the fish grows the 4 body bars
completely split to form two separate bars, making 8 body bars, sometimes with
fragmented spots or blotches in-between. The solid colour of the bars is not as distinct in
ontogeny, and the bands and stripes in the fins become less distinct (see original
description). In larger specimens the bars can anastomose or almost fuse back together.
The marks on the caudal fin become higher in number.
Botia kubotai Kottelat, 2004
In view of the recent description of this species (Kottelat, 2004), I consider it superfluous
to describe it further other than to say that there is variation within the markings. Some
have large disintegrated pale spots, some small. I have seen an image of one specimen
that had the bands where they were only disintegrated in pairs, not connected with other
pairs. Sexual dichromatism may be in play here, as in B. almorhae. Some aquarists have
debated about whether this means that there is more than one species involved. I
personally consider that providing someone isn’t getting mixed up with Ataran B.
histrionica, if a specimen has come from the Ataran or rivers around it, it is unlikely that
they will have a different species just because there is some slight variation in the
markings. It is found in the Salween system, in Myanmar and Thailand (border region).
Found in the Ataran River, and Suriya River (Panitvong, 2006).
Botia sp. ‘Ladder’
This undescribed species or variant possibly came in as a contaminant with B. kubotai
(based on what its owner Dave Speed can remember). I have seen images of specimens in
aquariums but it appears to be rarely obtained. The number of bars and their placement
and disintegration seems to show this as a member of the B. rostrata complex. This may
mean that the specimen originally came from the Brahmaputra or Ganges system rather
than Myanmar or Thailand.
PART 2
Mainland fishes of the genus Syncrossus Blyth, 1860
(Teleostei: Botiidae)
Key words: Botiidae, loaches, Syncrossus, Hymenphysa, Hymenophysa
Introduction
Whilst planning and working on Part 1 I found myself and Andy Rushworth trying to
work out the identification and validity of the Syncrossus loaches of the mainland. I say
mainland because the identities and validity of the Syncrossus of the archipelago
(Indonesia, Bornean Malaysia, and Brunei) appear settled.
As with Part 1, this work is not an attempt to resolve the issues as far more specimens
would need to be utilised and actually examined. Also I understand that an indepth
scientific study is ongoing. This work is a discussion on the various species and ‘forms’
(some which appear to be undescribed species), and is meant to highlight them formally
in writing (users of Petfrd and Loaches Online forums have long discussed these
problems); I have however used some previously synonymised names if I consider that
they are sufficiently different. Readers should treat these with caution though as it may
well be shown in the future when they are studied in depth that although they have
differences, they may not be considered sufficient enough to warrant separate species.
Images of live specimens with known locality data (predominantly by Kamphol
Udomritthiruj, and some by Rainer Hoyer) have been extremely useful. There are
numerous specimens of each species and form in some of the museums that have allowed
access to specimens, but unfortunately I have not been able to utilise them all. Also,
caution needs to be taken as some of them are misidentified.
As well as the abbreviations for museum collection used in Part 1, I have also utilised a
specimen in IHCAS (Museum of Institute of Hydrobiology, Chinese Academy of
Sciences, Wuhan, China).
Syncrossus, Hymenphysa, or Hymenophysa?
Nalbant (2004) argues that Hymenophysa Bleeker, 18587 is not an incorrect subsequent
spelling of Hymenphysa McClelland, 1839, and that Bleeker intended to describe it as
7
In Bleeker, 1863, the date of the publication often quoted as 1858-59 (following Weber & de Beaufort,
1911) is given as 19 August 1958, and the name of a meeting mentioned. It is not clear if the paper was just
discussed there, submitted to the meeting on that date, or whether it has been published and handed out
new, or is at least an unjustified emendation and is in prevailing usage; and therefore it is
the valid name for the group Grant (2001) and Kottelat (2004) grouped under Syncrossus.
I do not agree that Hymenophysa is in prevailing usage. Although Fang (1936) (but he
clearly attributed it to McClelland), Nalbant (1963, 2002, 2004) and Taki (1972) used
Hymenophysa; me (2001), and Kottelat (2004), used the genus Syncrossus; Roberts
(1989) classed Hymenophysa as an unwarranted emendation of Hymenphysa, and
Eschmeyer (1990) classed Hymenophysa as an incorrect subsequent spelling. The Code
does not appear to back up Nalbant’s view (glossary explaining prevailing usage, and
also Article 33.5 about what view takes precedence in a dispute of this kind). Also, in
what appears to be the same publication as Nalbant takes Hymenophysa (pages 302-304
of Bleeker, 1858), Bleeker at page 358 changes his Cobitis hymenophysa to
Hymenophysa macclellandi (to unnecessarily avoid tautonymy). This to me shows he was
linking his use of Hymenophysa, with McClelland’s Hymenphysa, and not proposing a
new genus of his own. More importantly, in addition to this, Bleeker himself (1863: 34,
and 36) makes it clear that Hymenophysa is the same as McClelland’s Hymenphysa: to
quote him (roughly translated) at page 34:“I also note that the name of Hymenphysa or
Hymenophysa McClelland like that of Schistura McClelland being posterior to that of
Botia, proposes by Mr. Gray for a species of the same kind, will be able to take seat only
among the synonyms.” On page 36 he writes “Botia Gr. = Hymenphysa McCl. =
Hymenophysa McCl….”. Also in Bleeker (1863-64:3 & 5) he repeats “Botia Gr. =
Hymenphysa McCl. = Hymenophysa McCl…..”.
Therefore, as in 2001, and in agreement with Kottelat (2004), I use the name Syncrossus
as the valid generic name for this group of fishes, and view Hymenophysa as an
unjustified emendation that is not in prevailing usage. See Kottelat 2004:13 for a
diagnosis of the genus.
Type species
Whilst researching for Part 1, in May 2007 I found an earlier type species designation for
Syncrossus. The currently accepted earliest designation is by Jordan (1919), but Jordan &
Fowler (1903) designated Syncrossus berdmorei Blyth, 1860 as the type by their use of
parentheses for type species. This was a method used on a regular basis by Jordan and his
co workers Gilbert, Fowler etc (see Eschmeyer et al, 1998; this method has been accepted
as establishing a type species for numerous genera). Jordan & Fowler’s designation of a
type species for Hymenphysa (spelled Hymenophysa by them but clearly referring to
McClelland) of Cobitis hymenophysa Bleeker, 1852 is not valid as C. hymenophysa was
not an originally included species in McClelland’s Hymenphysa.
Differentiating characters
As I have discussed in Part 1, pattern and colour appears to play an important role in
Botia differentiation. This is also the case in Syncrossus, and in fact Kottelat (1984) could
there (Bleeker had a habit of sometimes putting the date he finshed the paper or submitted it). In view of
this uncertainty, and Article 9.9 of the Code, and the fact that workers have been unable to determine if the
separate articles written by Bleeker were published in one go in Natuurkd. Tijdschr. Neder. Indië, and that
either that one publication has an uncertain date; or if they were printed separately and they have uncertain
dates: we may at the moment be stuck with 1858-59. However, Nalbant, and Kottelat use 1859, but I will
use 1858 in view of Article 21, until the publication date is settled.
not find any morphometric or meristic differences between two species, but found the
colour pattern diagnostic and having a taxonomic value. As in Botia, what has also been
useful is the zoogeography of the species and forms. I would have preferred to have had
x-rays of all the specimens here and the numerous others in the institutions, but this has
not been possible. Again, as in Botia I have found morphometrics to be of little or no
value, but perhaps those who work on these fish in the future will have larger samples
which may show some useful differences.
There are currently only 8 described taxa for fishes that belong in Syncrossus (plus one
that is a replacement name). Only 5 of these 8 have been recognised (Kottelat, 2004), but
there appears to be around 12 or more forms that could in fact be different species when
studied further.
Distribution
In addition to the Salween, Irrawaddy, Sittang, and Ataran river systems discussed in Part
1, Part 2 discusses fishes from the following systems:
Chao Phraya
This is a major river system in Thailand. The Mae Nam Ping (sometimes referred to in
specimen localities as the Meping) and the Mae Nam Yom converge to make the Chao
Phraya. It is hypothesised that in the past the Mae Nam Ping was the lower course of the
Salween, and the Mae Nam Yom the lower course of the Mekong (see Kottelat, 1989).
Mekong
This major river system starts in China’s Qinghai Province, and runs through Tibet (as
the Dza Chu) and Yunnan (as the Lancang Jiang). It forms the border between Myanmar
and Laos and is met by its tributary the Ruak River, which marks the division between
Upper and Lower Mekong. The river also divides Laos and Thailand in parts. It passes
through Cambodia, then through southern Vietnam, where it splits into two branches, the
Tien Giang and the Hau Giang, which both empty into the sea. The Mae Nam Chi (Chi
River) and Mae Nam Mun of Thailand are connected to the Mekong. See Rainboth
(1996) for a detailed discussion.
Mae Klong
A river and system in western Thailand, not to be confused with the Mae Nam Khong, a
Lao and Thai name for the Mekong. The Khwae Noi River (Kwai) starts near the border
with Myanmar. The Khwae Yai River (which also starts near the border) eventually
merges with the Khwae Noi to form the Mae Klong.
Pahang
A river system in Peninsular Malaysia; where it is called the Sungai Pahang. The Tasek
Bera swamp drains into the Pahang via the Bera River. Approximately 10,000 years ago
there was a land and freshwater connection between the Pahang and Chao Phraya and
Mekong to the north, and to the Sarawak area of the island of Borneo to the east (see
Kottelat, 1985; Ng, 2002).
Syncrossus helodes (Sauvage, 1876) – 2 or 3 forms
Botia hymenophysa (nec Bleeker) Fowler (1934, 1937), Smith (1945), Suvatti (1981),
Mai et al (1992).
Preserved specimens used: Chao Phraya system: USNM 109708, “Meping R. At
Chiengmai, N. Siam”, 1 spec., c. 95mm SL; USNM 109707, “Siam: Northern, Meping R.
At Chiengmai”, 2 spec., c. 105mm – 120mm SL, ANSP 56659, “Doi Angka, Chieng Mai,
Thailand”, 1 spec., 166mm SL; MCZ 35536, “Chiang Mai, Thailand”, 3 spec., c. 146mm
– 156mm SL; USNM 271701, “Thailand, 14 33 N, 100 46 E”, 2 of 7 spec., c. 84mm 90mm SL;
Mekong system: USNM 271732, “Lam Chi, 1.5 km Below Hwy 23 Bridge, 4 km W of
Selaphum, Roi Et Province, NE Thailand. 16 01 N, 103 52 E”, 1 spec., c. 111mm SL;
USNM 271744, “Morning Market Muang Khong, Sithandone Province, Laos. 14 06 30
N, 105 51 30 E”, 2 of 6 spec., c. 107mm SL – 118mm SL.
Distribution: Chao Phraya system, Mae Khlong system, Mekong system in Thailand
(Fowler, Smith, Suvatti, 1981, Vidthayanon, 2004). Mekong system in Cambodia
(Rainboth, 1996), Laos (Kottelat, 2001), and southern Vietnam (Mai et al, 1992).
Merisitics: Dii-vi 12-15, Pi 12-14, Vi 7-8, Aii-iii 5, C17. (Nguyen (2005) states there can
be 11 branched dorsal rays, but this may be a miscount as the 12th ray is small and usually
clamped against the penultimate ray). I have found that when counting non branched
dorsal fin rays in preserved specimens using the naked eye, you usually get a count of iiiii. However, I have looked at x-rays of 3 specimens (USNM 109707, 109708) and
counted between v and vi, as 2 to 3 of them are usually hidden under the skin. The first 4
tend to share a pterygiophore.
Discussion: Sauvage described S. helodes from “Tma-Kré” which is in present day
Cambodia. I have been unable to find this spelling on any map but there is a Thmâ Kre at
the side of the Mekong River, just north of Krâcheh [Kratié] city, Krâcheh Province,
Cambodia. This is probably the same locality as the type locality given by Sauvage.
Until Kottelat resolved the issue in 1984, S. helodes was usually wrongly identified as
“Botia hymenophysa”. Pictures of Mekong specimens are shown in Rainboth (1996) and
possibly Kottelat (2001) (I’m assuming this is a Mekong specimen). Mai (1978 & 1985)
found no S. helodes in northern Vietnam, and Serov et al (2006) did not in central
Vietnam, but Mai et al (1992) did in southern Vietnamese drainages of the Mekong. See
details of specimens above for Mae Nam Ping and Mekong specimens and other Chao
Phraya specimens. See Vidthayanon (2004) for a picture of a Thai specimen. Suvatti
(1981) mentions Chao Phraya and Mekong (in the Mae Nam Chi) occurrences.
I have tried to check for any meristical differences between Chao Phraya specimens and
Mekong specimens. The USNM and MCZ Chao Phraya specimens above have 13
branched dorsal fin rays, the same number given by Kottelat (1984) for the holotype of S.
helodes. Mai et al (1992) give 12, and Nguyen 11-12 (2005). Kottelat (2001) gives a
range up to 15 (he gave 14 ½ as he classes the last ray as ½ a ray). Images by Terry
Warren and Ian Baird of specimens from the Mekong appear to show 13, and the Muang
Hong USNM specimens appear to have 14. So this means we have Chao Phraya
specimens with 13, Mekong with 12-15 (11 if you believe Nguyen). I do not know if all
Chao Phraya specimens always have 13, and if Mekong ones vary depending on their
location. Therefore I cannot at this stage assign any importance to this. I cannot see any
other morphometric or meristical differences of note. The Chao Phraya specimens appear
to have yellow ventral and anal fins (Smith, 1945, and Fowler, 1934), the Mekong
specimen in Rainboth (1996) and the Muan Khong USNM specimens have them red, the
description by Sauvage of his Mekong holotype describes them as blackish (from a
preserved specimen), the Mekong specimens pictured by Warren and Baird show them
orange, and Kottelat (2001) shows them yellow (not sure if this is actually a Mekong
specimen or Chao Phraya?). Even if it was shown that Chao Phraya ventral and anal fins
were always yellow, and Mekong always red or orange, this alone is probably not
significant anyway. However, it is possible that the Mekong form has darker body bars
than the Chao Phraya form (which appear bluish). Fowler (1934, fig. 51) shows a Chao
Phraya specimen, and in 1937 (fig. 64) a Mekong specimen of about the same size. The
patterns do look different, and the shapes of the outer edge of the dorsal fins look
different. However the Mekong specimens pictured by Baird and Warren have the dorsal
fin the same shape as Fowler’s Chao Phraya specimen; I have also been told that
Fowler’s figures were not always anatomically accurate. As discussed earlier it is
hypothesised that parts of the two systems were connected in the past so this explains
them being present in both systems. Further study (perhaps utilising x-rays of both forms
as I only have them of Chao Phraya form, and DNA analysis) is needed as the Chao
Phraya form (possibly including the Mae Khlong form?) may be a different species or
subspecies?
Syncrossus hymenophysa (Bleeker, 1852)
Junior synonym and unneeded replacement name: Hymenophysa macclellandi Bleeker,
1858.
Preserved specimens used: BMNH 1932.5.19.26, “Telom River, Pahang, Malaysia”,
collected by Capt. F. A. B. Holloway, 1 spec., c. 113mm SL; BMNH 1922.5.19.91-93,
“Tembeling River, Malay Peninsula”, presented by C Kloss, 3 spec., c. 112mm-118mm
SL; BMNH 1906.2.28.64, “Kuala Teku, Pahang”, 1 spec., c. 122mm SL.
Distribution on mainland: Only found in Peninsular Malaysia, so far recorded from the
Pahang system, and Perak system.
Sungai Tahan, at Kuala Tahan, Pahang system (Tweedie, 1956); Taman Negara National
Park, Pahang system (Hora, 1941); Tasek Bera, Pahang system (Hora, 1941); see above
for 5 Pahang system specimens from BMNH; Tasik Chini, Pahang system (Khan et al,
1996).
Lake Chenderoh, Sungai Perak, Perak (Mohsin & Ambak, 1983).
I cannot find any reliable reports of it from Singapore.
Meristics (for peninsular Malaysian specimens): “D 13, P 15, V 8, A 7”. These meristics
are taken from Mohsin & Ambak (1983) for one specimen, and they appear to be for the
branched and non branched rays combined. The specimens in the BMNH appear to have
Diii-v 13-14, P 15, A i-ii 7, C 17.
Discussion: Bleeker described S. hymenophysa based on a specimen from Palembang,
Sumatra, Indonesia. Bleeker (1863) and Weber & de Beaufort (1916) give details and
figures, and Kottelat (1984), Roberts (1989) and Kottelat et al (1993) have recently
discussed archipelagic occurrences of S. hymenophysa. Based on these works it can be
seen that the archipelagic distribution (including the type locality) is the islands of
Sumatra and Borneo (it may occur in Java but the record there of Weber & de Beaufort
may relate to Syncrossus reversa Roberts, 1989). Roberts (1989) said that the records
from mainland Asia (from the Malay Peninsula) needed to be re-examined to confirm
their identity. Although I personally have not been able to examine specimens, James
Maclaine of BMNH has kindly sent me images and x-rays of the BMNH specimens from
the Malay Peninsula. I can confirm that (in my opinion) they are conspecific with
archipelagic and topotypical S. hymenophysa. The pattern is exactly the same, and the
dorsal fin ray count range is exactly the same as given by Roberts (1989). Mohsin &
Ambak (1983) list the species as endangered. I would hazard an opinion that this species
may now be critical or extinct in the Malay Peninsula. Mohsin & Ambak found only one
specimen, and the last recorded finding of them before that was in 1955 (Tweedie, 1956).
All recorded collections that I have been able to find have been a few specimens at a time
only. Mizuri et al (1982), Lim et al (1990), Ng & Tan (1999), Sim (2002) did not report
any specimens. I have not seen Khan et al (1996) but I have been informed that no
specific details of specimens are listed, but that it is classed as “Not common”.
Syncrossus berdmorei Blyth, 1860 – various forms
Preserved specimens used:
Sittang system: AMS.B 7786, (syntype of S. berdmorei), “Burma” (see comments below,
not definitely from the Sittang), 1 spec., 100mm SL; USNM 378413, “Kayin Stream near
the Sittallang River, east of Taungoo, Myanmar”, collected by R. Britz, 1 spec., c.
58.5mm SL.
Manipur: BMNH 1975.11.21.15-16, “Manipur, India”, donated by Sarkas, 1 of 2 spec., c.
40mm SL.
Irrawaddy system: USNM 372227, “Myitkyina Market, Kachin State, Myanmar”,
collected by C. Ferraris, Tun Shwe & Mya Than Tun, 1 spec., c. 94mm SL; USNM
378418, “254 feet. (Shwe Sa Kayth), Myanmar. 22 20 22 N, 96 59 36 E”, collected by R.
Britz, 2 spec., c. 68.4mm SL – 71.6mm SL; USNM 372220, “Myanmar, Yangon
Division, Hliang River, Insein Township. 16 53 41 N 96 05 28 E”, collected by C.
Ferraris, Mya Than Tun, Yin Shein & Local Fishermen, 1 spec., c. 46.5mm SL.
Distribution: See under discussion but appears to be India (Manipur), Myanmar
(Irrawaddy, Sittang, Salween, Ataran).
Discussion: Blyth described S. berdmorei from “Tenasserim provinces”. This gives us a
problem in establishing the locality of the type specimen(s). Blyth did not say how many
specimens he had but the way on which he describes the pattern, shows he had more than
one specimen, but were they all from different localities, hence “provinces” instead of
‘province’? Day (1870) lists 5 specimens and obviously viewed them all as syntypes.
Menon and Yazdani (1968) list ZSI F2636/1 as syntypes but only list 4 specimens, and
Eschmeyer et al list AMS B.7786 as a possible syntype, probably based on the AMS
listing it as such. This situation is similar to that discussed in Part 1 for Blyth’s Botia
histrionica, not only about the issue of the type locality but because S. berdmorei forms
are found in the Manipur River (Chindwin drainage), Irrawaddy, Sittang, Salween,
Ataran. So if the different forms do represent different species (which I consider that they
do), which is the true S. berdmorei? I do not have access to the ZSI specimens but Day
(1870) and Menon (1993) did and did not list a more accurate locality. I have had
information and images sent of the AMS specimen. According to its database the AMS
specimen has a locality of “Madras”, but there are no species of Syncrossus there, the
only place in India I am aware of that has Syncrossus is Manipur. Some of the putative
types in the AMSs Day collection have turned out to be types but some have not, and
some of them being the specimens that Day used (and sometimes illustrated) in his Fishes
of India (Grant, 1999; Ferraris et al, 2000). I asked the AMS to look into their records
and upon checking Sally Reader (personal communication) has confirmed that the AMS
Annual Report of 1884 lists the specimen as from “Burma”, so Madras has been inputted
in error. There is a note in the report that states “one of Blyth’s types”. I understand from
Ferraris et al that Day’s collection included some of Blyth’s specimens from species he
described. I considered whether the AMS specimen is the specimen that Day discussed
and figured in his Fishes of India (page 607, Plate CLIV fig. 3). The specimen was said to
be life sized, making it approximately 80mm SL and 100mm TL. I have been informed
that the AMS specimen is 100mm SL, and 123mm TL which means it is too large to be
the specimen figured by Day. The specimen in Fishes of India was said to be from
Mandalay in Myanmar, which means this would not be a syntype of Blyth’s S. berdmorei
as Mandalay is not part of the historic Tenasserim Provinces region. Eschmeyer et al list
the AMS specimen as from Moulmein [Mawlamyaing] which would tie in with the
historic Tenasserim Provinces region, and would make the specimen either from the
Salween, the Ataran, or the Gyaing. However, I cannot see where the locality of
Moulmein has been taken from and is possibly an error (AMS B.7529 of
Lepidocephalichthys berdmorei is from Moulmein). In view of the fact that Day said
there were 5 specimens, and Menon & Yazdani only listed 4 specimens, and the AMS
Annual Report lists it as being one of Blyth’s types, and the size fitting in with the range
given by Blyth, I consider that AMS B. 7786 is a syntype of Syncrossus berdmorei. As
discussed in Part 1, most of Blyth’s species were allegedly caught in the Sittang River
and its tributaries, as with B. histrionica I have been informed of the existence of
specimens of a form of S. berdmorei that were definitely caught in the Sittang system,
and have used a USNM specimen from the Sittang. Because of the unresolved issue of
the type specimens, and them not having accurate riverine localities, Blyth’s description
of the colour and pattern of his species is important as specimens collected of each form
sometimes have slightly differing patterns and/or colours to those from some other rivers.
The problem is that although they are different, the patterns change in ontogeny (as the
fish gets bigger, more spots tend to form in the body), and also they are similar, and it is
usually colours that are different. All but perhaps one of the forms can fit the description
of the pattern described by Blyth in terms of number of bars in the body, fins, and lines
on the head etc. The only possible exception to this is the form pictured here by Kamphol
as “Central Myanmar”, as this form has spots even in the posteriormost body bars even in
small specimen, and this doesn’t appear to be the case in the AMS syntype. Because of
this problem of identifying which form is the true S. berdmorei, I have decided just to
picture each one until the work that I know is ongoing can decide for definite which river
systems the syntypes came from, and whether the different forms are indeed different
species or subspecies. If it is shown that they are not then where the syntypes were caught
wouldn’t be that important.
The other issue that hasn’t been adequately resolved yet is what is the difference between
a S. berdmorei form, and a S. beauforti? This is important because S. beauforti ‘forms’
are found in the Ataran, and so are S. berdmorei forms. The main visual differences
appear to be:
•
•
The number of branched dorsal fin rays: up to 9 or 10 in S. beauforti ‘forms’, and
up to 14 in S. berdmorei forms
In S. berdmorei forms the young usually always start off with distinct body bars
and as the fish grows the spots start in the region behind the opercle (and on the
‘Central Myanmar’ form within all the bars), whereas in S. beauforti ‘forms’ even
very small specimens have the horizontal lines of spots running across the body
from the snout and head region.
This needs further study.
In the aquarium trade there have recently been imports of very small specimens of
Syncrossus with yellow fins that were reported to be from Laos, but that exact locality
details were being withheld. The specimens look very similar to a S. berdmorei form, and
have the right number of dorsal fin rays (11 branched), but do not appear to be the correct
body shape and pattern for a S. helodes. Based on what I can glean from numerous
images of their development provided by Emma Turner on Loaches Online, and
comparing this with images of Irrawaddy and Ataran forms of S. berdmorei and also with
other Syncrossus species and forms, my opinion is that they are Myanmarian Ataran S.
berdmorei form and are not from Laos. Only time and further study will tell.
Syncrossus beauforti (Smith, 1931)
Junior synonym?: Botia lucasbahi Fowler, 1937 (originally lucas-bahi)
Botia lucas-bahi Fowler, 1937 & 1939
Preserved specimens used: Chao Phraya system: USNM 107853, “Mechem R. Trib. of
Meping, N. Siam”, 2 of 5 spec., c. 41mm – 46mm SL; ANSP 68005 (holotype of B.
lucasbahi), “Tachin, Siam”, 1 spec., 56.7mm SL. ANSP 56843 (paratype of B.
lucasbahi), “Chieng Mai, on Me Nam Ping, North Siam”, 1 spec., 69.5mm SL.
Identity uncertain: ANSP 60087, “Chieng Mai, North Siam”, 1 spec., c. 78mm SL.
Distribution:
Peninsular Thailand: Tadi Stream, Ban Kiriwong, Amphoe Lan Saka, Nakhon Si
Thammarat Province; Trang, Trang Province.
Chao Phraya system, Thailand (it may also be present in the Mae Khlong).
Ataran River basin, Myanmar.
Discussion:
The first thing I felt I needed to do was to try and find the paratype of B. lucasbahi, as
Fowler (1937) clearly designated one. Prior to the 9 October 2007 version of the Catalog
of Fishes, only the holotype was mentioned, which is in ANSP. I checked ANSP and
USNM collection databases and the paratype wasn’t listed. My view was that either it
had been lost, or likely been misplaced. I checked with ANSP and they had no record of
it in the collection. I started going through ANSP and USNMs collection database to see
if any Chieng Mai specimens collected in 1932 had perhaps been left in the name of S.
hymenophysa, or perhaps changed to S. beauforti. There were three candidates in ANSP
(60092, 60094, and 56843). I asked Mark Sabaj if he could check the sizes of the
specimens to see if any matched with that given by Fowler. Mark confirmed that 56843
matched exactly the size and pattern given in Fowler (1937) and he agreed that this was
the ‘lost’ paratype (and the 9 October 2007 Catalog of Fishes now shows the paratype).
However, when I was kindly sent an image of 56843, it did not appear to match the
drawing in Fowler (1934) fig. 52, which according to the footnote in Fowler (1937) is the
paratype (Smith, 1945:288 also lists it as such). 56843 looked just like the holotype of B.
lucasbahi and not that 1934 figured specimen which had a different pattern and unusually
had 7 dorsal fin rays (very low for a Syncrossus, even 8 being low). I queried this and
Mark kindly replied with the following:
“Fowler's (1937:154) designation of a paratype for Botia lucasbahi appears to correspond
to 2 specimens: ANSP 56843 and 60087.
Fowler's (1937:154) paratype designation refers to a specimen previously reported in
Fowler (1934:101) as Botia hymenophysa (Bleeker) collected from Chieng Mai in Dec
1932 and having length (TL) 90 mm. This corresponds best with ANSP 56843, TL 89
mm, a specimen that has 9 branched dorsal rays (consistent with Fowler's count for the
holotype of B. lucasbahi, ANSP 68005), pigmentation pattern consistent with Fowler's
(1934:101) description, and collection data as Chieng Mai, Dec 1932. ANSP 56843,
however, does not match Fowler's (1934:100) fig. 52 with respect to TL (89 mm vs. 96.5
mm for figured specimen), branched dorsal rays (9 vs. 7 in figure) and pigmentation
pattern (many small dark spots overlying fainter bars on body vs. small dark spots absent
in figure).
The specimen figured by Fowler (1934:100, fig. 52) measured about 96.5 mm TL based
on scale bar and corresponds best with ANSP 60087, 96.3 mm TL. ANSP 60087 was
collected from Chieng Mai by R.M. DeSchauensee 8 Feb 1933, has 7 branched dorsal
rays as does specimen in fig. 52, and agrees well with figured pattern of spots in dorsal
fin and vertical bars on body (except figured specimen has one fewer bars on posterior
body). ANSP 60087 would not be considered the paratype except for Fowler's (1937:154)
footnote which refers specifically to Fowler's (1934) fig. 52. This footnote confuses the
paratype designation because Fowler's (1934:101, 1937:154) text references point to a
different specimen, ANSP 56843, currently treated as the "true" paratype.”
Gladly, this has now cleared up the confusion about what the paratype looks like, but
leaves us with the question: What is ANSP 60087? It appears to have the pattern and a
similar body shape of a Syncrossus, but also has traits of a Yasuhikotakia in its body
shape and proportions and its dorsal fin ray count (although looking at an x-ray of it there
appears to be 8 branched dorsal fin rays if you count the penultimate one, not 7 as
reported by Fowler). I show it here so that you can draw your own conclusions. It may be
a Syncrossus yunnanensis based on its pattern, and if so, its presence in the Chao Phraya
can be explained by the fact that the Upper Mekong was once connected to the Chao
Phraya. Further work on and examination of the specimen is required. The image by
Kamphol of a specimen from the Chao Phraya may represent this species.
Unfortunately the holotype of S. beauforti (USNM 90285, from “Tadi Stream, Ban
Kiriwong, Amphoe Lan Saka, Nakhon Si Thammarat Province, Malay Peninsula,
Thailand”) is currently missing. It was not present in the USNM in 1980 when they did
an inventory, and is still missing (Raredon, personal communication). Therefore I have
not had specimens from peninsular Thailand to compare with those from Chao Phraya
and Mekong and therefore the decision that B. lucasbahi is a synonym of S. beauforti is
tentative and is based on pattern and meristics from the original description by Smith, and
the specimen in Fowler (1939). The two appear very similar to me and I have struggled to
ascertain any difference. However, more and fresher specimens from the type localities
may reveal subtle differences in colour and pattern, and other differences. In view of the
findings on Botias, it may well be that the tentative view put forward here that the same
species is present in the Ataran, Chao Phraya, Peninsular Thailand, and possibly the Mae
Khlong is incorrect.
Syncrossus yunnanensis (Chen, 1980)
? Botia hymenophysa (nec Bleeker) Fowler (in part), 1934
? Botia beauforti (nec Smith) Kottelat & Chu, 1987
Botia beauforti (nec Smith) Kuang, in Chu & Chen, 1990
Botia yunnanensis Kuang, in Chu & Chen, 1990
Botia beauforti (nec Smith) Zhu, 1995
Botia yunnanensis Zhu, 1995
Preserved specimens used: IHCAS 638040 (holotype of S. yunnanensis), “Jinghong,
Yunnan, China”, 1 spec, 120mm SL.
Meristics: Div 9, Pi 12, Aiii 5, Vi 6
Distribution: Upper Mekong system in southern Yunnan (Jinghong & Menghan), China.
It may be present in northern Laos in the Upper Mekong. If ANSP 60087 represents S.
yunnanensis then it is also present in the Chao Phraya.
I tentatively listed this species as valid. This is based on the fact that the holotype was
found in the Upper Mekong, and it appears to differ from Mekong Form S. beauforti
(which should be called Syncrossus formosus if a valid species in its own right) in that it
appears to have no or very few spots in the body, even at adult sizes. Also, according to
the original description there are 34 total vertebrae in the holotype. In 4 Mekong S.
formosus x-rays (inc. some syntypes of S. formosus) I count 32, 32, 32, 33.
Syncrossus formosus (Pellegrin & Fang, 1940)
Botia beauforti (nec Smith) Smith, 1945 (in part, specimen from Pakjong, Mae Nam
Mun, Thailand).
Botia beauforti (nec Smith) Mai et al, 1992
Botia beauforti (nec Smith) Rainboth, 1996.
Botia beauforti (nec Smith) Kottelat, 2001.
Botia beauforti (nec Smith) Nguyen, 2005.
Botia beauforti (nec Smith) Serov et al, 2006.
Preserved specimens used: MNHN 1939-0218-0220 (3 of the 5 syntypes of B. formosa),
“Ban Nam Khueng, Mekong, 30 km north-east of Ban Houei Sai, Laos”, 3 spec.; BMNH
2000.6.10.518, “Mekong River, Dondou village, Hinboun district, Khammouane
province, Laos”, collected by villagers, 1 spec., c. 84.7mm SL; USNM 229289, “Menam
Kok (Mekong Basin) At Thaton, Myanmar”, collected by Tyson Roberts, 1 of 2 spec., c.
61.5mm SL; USNM 109741, “Menam Mun at Pakjong, E. Siam”, collected by H. Smith,
1 spec., c. 156mm SL.
Distribution: Lower Mekong system in Thailand, Laos, Cambodia, and southern
Vietnam.
Discussion: I have tentatively listed this species as valid. Live pictures of specimens of
known Mekong system locality by Kamphol, and by Ian Baird, coupled with images of
the preserved specimens above appear to be much more spotted than Chao Phraya (S.
beauforti) or Upper Mekong (S. yunnanensis) specimens (live and preserved). I cannot
ascertain any notable meristical differences but pattern combined with distribution has
led to me to tentatively conclude this as a valid species.
Further work is needed.
Other comparative material
Yasuhikotakia modesta?: USNM 107853, “Mechem R. Trib. of Meping, N. Siam”, 3 of 5
spec., c. 21mm – 24mm SL.
PART 3
A new subgenus of Acanthocobitis Peters, 1861
(Teleostei: Nemacheilidae)
Key words: Nemacheilidae, loaches, Nemacheilus, Schistura, Acanthocobitis, Cobitis
pavonacea, Cobitis zonalternans, Cobitis botia, Acanthocobitis longipinnis
Whilst researching for Parts 1 and 2 I came across the original description of Cobitis
pavonacea McClelland, 1839. The description and figure of the specimen matched that of
a live specimen that had been exported from the type locality. It became obvious to me
that this species had characteristics that made it sufficiently different to warrant its own
subgenus, but a subgenus of what genus? Day (1878) classed Acanthocobitis longipinnis
Peters, 1861 as a synonym of C. pavonacea, and in fact Peters himself put after the new
name he was proposing “(? Cobitis pavonacea M. Cl)”. However, subsequent authors
(Menon, 1987; Talwar & Jhingran, 1991) have classed A. longipinnis as a junior
synonym of Cobitis botia Hamilton, 1822, and considered C. pavonacea as a valid
species of Acanthocobitis, or Nemacheilus (or the nomen nudum Noemacheilus – see
Kottelat, 1987, and 1990). Cobitis botia is currently generally accepted as a member of
the genus Acanthocobitis, using the synonymy of Acanthocobitis longipinnis (type
species of Acanthocobitis) = a junior synonym of Cobitis botia.
Banarescu & Nalbant (1968: 332) and Menon (1987:152) both recognised that C.
pavonacea is different to members of the genera they used. Banarescu & Nalbant and
Menon stated that it was a taxonomically isolated species and not closely related to any
of the other members of the genus Acanthocobitis (they used it as a subgenus).
Rather than accepting these synonymies and raising a subgenus for C. pavonacea, I
decided to check the descriptions and type specimens (where they exist) for each species.
I have been able to access the descriptions for all species, the holotype of A. longipinnis
(ZMB8 4795, which is circa. 156mm SL), and the syntypes of C. pavonacea (SMF 68,
9070, 9090, 90919, which are circa 102mm - 120mm SL); no types are known to exist for
A. botia.
McClelland described C. pavonacea from specimens from Assam in India. The drawing
in McClelland of one of the specimens clearly shows the species to have a long based
dorsal fin, and a characteristic pointed caudal fin. Thanks to Horst Zetzsche I have also
been able to access images of the syntypes and can confirm that SMF 68 still show the
remnants of this characteristic shape (in SMF 9070, 9090-91 the rays are not as well
preserved). The drawing, the description, and the images of the syntypes match exactly
with that of the (unpreserved) specimen shown live here. This specimen came from Dibru
River, Guijan, around 12 km from Tinsukia town, Assam, India (Rao, personal
communication). Its meristics are Div 18 (last ray not branched), Pi 12 (last ray not
branched), Vi 7 (last ray not branched), Ai 6, C 19 or 20? These counts match or are very
close to those given by McClelland “D.17 : P.13 : V.9 : C.20”, bearing in mind that he
will have represented the formula of branched and unbranched rays differently. On one
side the lateral line is damaged due to previous fighting so it appears broken from around
parallel with the 7th branched dorsal ray and the penultimate branched dorsal ray. On the
other side of the body the lateral line is complete (as it is in the syntypes). There is a
small suborbital flap (not visible in SMF 68, but is in SMF 9070, 9090-91). The cephalic
lateral line pores are distinctly visible (they exactly match the syntypes). The anterior
nares are tubed; there are 3 pairs of barbels. The mouth, jaws (process dentiformis
present) and lips match those of the syntypes, which in turn are similar to other
Acanthocobitis, albeit the lips do not appear as papillated when compared to the figures
in Kottelat (1990). These fish have in the past been sold in the hobby as Panther (or
sometimes Puma) Loaches, however, McClelland compared the ocellus on the caudal
peduncle with that of a Peacock (‘pavonacea’ or ‘pavonaceus’ is from Latin: of or
belonging to a peacock (Lewis & Short, 1879); and the generic name of the Asian
Peacock birds is Pavo); so the name in the trade should be the Peacock Loach. A.
pavonacea was incorrectly captioned as A. rubidipinnis in Tekriwal & Rao (1999:89).
Day (1878) gives the fin ray counts as Di-iii 14-18, P 14, V 8, Aii 5, C19; Peters gives
the meristics of the holotype of A. longipinnis as Div 18, P 18, Vi 7, Aiii 6. The drawing
in Day on Plate CLIII fig 12 (and his key on page 612) shows the caudal slightly
emarginate. Menon (1987) reproduces this drawing. It is not clear if this drawing is based
on a specimen or not but this appears to be in error, and Day’s drawing may have been
taken from one of the damaged syntypes. The drawing in McClelland and SMF 68 clearly
show the caudal fin pointed.
8
Universitat Humboldt, Museum fur Naturkunde, Berlin, Germany.
Natur-Museum und Forschungs-Institut Senckenberg, Frankfurt-am-Main, Germany. These specimens are
listed as syntypes by Rüppell (1852). The SMF database lists SMF 68 as lectotype and the rest as
paralectotypes due to a selection made by W. Klausewitz, but this was never published officially and they
remain syntypes.
9
Thankfully, Dr Peter Bartsch was able to provide images of the holotype of A. longipinnis
(which was from the “Ganges River, India”). It became clear that it was the same species
as C. pavonacea. It matches C. pavonacea in every aspect, noticeably in the high dorsal
ray count, the long cylindrical body, the relative position of the ventral fins to the anus
and the anal fin, the shape of the head, the cephalic lateral line pores, and the pointed /
spear shaped caudal fin. The only character of note is that in the holotype of A.
longipinnis the suborbital flap is greatly enlarged and is a rigid structure, although it is at
the same place and angle as in C. pavonacea. This is probably because it is a large
sexually mature male (which also accounts for its slenderer body than SMF 68; and all of
the syntypes are at least around 35mm smaller than the holotype of A. longipinnis). I
consider that Acanthocobitis longipinnis is a junior synonym of Cobitis pavonacea, not of
Cobitis botia, but C. pavonacea does not belong in Cobitis, it does warrant its own genus,
which by synonymy of type species is Acanthocobitis, therefore Cobitis pavonacea
McClelland, 1839 become Acanthocobitis pavonacea (McClelland, 1839).
Kottelat (1990) queried why Peters would name his genus Acanthocobitis when the non
Indian specimens Kottelat had of A. botia did not have a spine under the eye, but had a
suborbital flap and was Peters referring to this? Peters described it (roughly translated
from German) as having ‘no mobile forked infraorbital spine’, but the infraorbital bone
having a “blunt thorn”. Incidentally, according to Hamilton’s description the holotype
(not preserved) of A. botia had “strong and undivided” “prickle” below each eye (it
would have been a flap if subsequent authors identification of A. botia is correct).
However, the other Hamilton species that are currently classed as junior synonyms of A.
botia, (Cobitis turio and Cobitis bilturio) did not have any “prickle”, but it could be that
these were females and as such didn’t have the suborbital flap. There were no drawings
of C. botia, C. turio, or C. bilturio published by Hamilton but the drawings on Plate LI
fig. 4. and fig. 6 in McClelland of Cobitis bimucronata and C. ocellata are (according to
McClelland, and later Hora) the drawings of C. botia and C bilturio respectively,
prepared but not used by Hamilton. These drawing show the typical shape and merisitics
of an A. botia. The drawing in McClelland on Plate LII fig. 7 of Cobitis gibbosa is also
supposedly the drawing of Hamilton’s Cobitis turio. This drawing shows a fish that is
similar to what is currently accepted to be an A. botia, but has only 8 dorsal fin rays. I
know that these matters are ones that have been covered by ichthyologists some years ago
but I consider that further investigation is warranted, bearing in mind the alleged wide
distribution of A. botia (Kottelat, 1990, has already mentioned that this is needed).
Based on the fact that I consider A. longipinnis a junior synonym of C. pavonacea, and
that A. longipinnis is the type species of Acanthocobitis, obviously it is the other species
currently assigned to Acanthocobitis that would need a separate subgenus, not the other
way round.
Paracanthocobitis new subgenus of Acanthocobitis Peters, 1861
Type species: Cobitis zonalternans Blyth, 1860
Gender: Feminine
Etymology: From the Greek para (shortened to par when combined with Acanthocobitis)
meaning ‘alongside’ or ‘beside’, meaning in this context similar to the genus
Acanthocobitis.
Diagnosis: A Nemacheilidae subgenus distinguished from Acanthocobitis sensu stricto
(meaning the nominotypical subgenus) by the following characters: Suborbital flap
horizontal or flap not present (a groove instead) Vs. in Acanthocobitis sensu stricto flap
placed diagonally to almost vertically; flap not greatly enlarged in any specimens Vs. in
Acanthocobitis sensu stricto greatly enlarged in some specimens; caudal fin rounded,
truncate or slightly emarginate Vs. in Acanthocobitis sensu stricto median and some rays
above, extended, giving a pointed or spear shaped appearance; anus closer to insertion of
leading ray of anal fin than to insertion of leading ray of ventral fins Vs. in
Acanthocobitis sensu stricto closer to insertion of leading ray of ventral fin than to
leading ray of anal fin; body comparatively more slender and elongated in Acanthocobitis
sensu stricto.
So far, all the species listed as valid by Kottelat (1990) under Acanthocobitis appear to
fall into the subgenus Paracanthocobitis.
Acknowledgements
A big thank you to James Maclaine, Natural History Museum, London, who patiently provided
much information and assistance. Thank you to Patrice Pruvost & Claude Ferrara, Museum
National d'Histoire Naturelle, Paris; Jon Fong, California Academy of Sciences; Mark Sabaj and
Kyle Luckenbill, Academy of Natural Sciences, Philadelphia; Mark McGrouther, Sally Reader,
Amanda Hay and James King, Australian Museum; Andrew Williston, Museum of Comparative
Zoology, Harvard; David R. Edds, Emporia State University, Kansas; Sandra Raredon,
Smithsonian Institution, Washington; Andrew Bentley, Kansas University; Dr. Zuogang Peng,
Museum of Institute of Hydrobiology, Chinese Academy of Sciences, Wuhan; Dr Peter Bartsch,
Museum fuer Naturkunde
der Humboldt-Universitaet zu Berlin; Dr. Horst Zetzsche & Friedhelm Krupp, Natur-Museum
und Forschungs-Institut Senckenberg, Frankfurt-am-Main, Germany; Bogdan Janiczak; Mark
Duffill; Kamphol Udomritthiruj; Andy Rushworth; J. C. Daniel, S. R. Sane, B. F. Chhapgar, and
Varad Giri of Bombay Natural History Society; Erik Åhlander, Swedish Museum of Natural
History, Sweden; Emma Turner of Maidenhead Aquatics, Peterborough; Andrew Arunava Rao of
Malabar Tropicals; Dr Maurice Kottelat; Rainer Hoyer & Gerhard Ott; Heok Hee Ng; Tyson R.
Roberts; Prof. Waikhom Vishwanath, Department of Life Sciences, Manipur University, India;
Ian G. Baird; Terry Warren; Ola Åhlander; Ken Childs; Martin Thoene; Graeme Robson; Helen
Nightingale; Dave Speed; Choy Heng-Wah; Phil Hurst, Natural History Museum, London. Allan
James for kindly creating the PDF.
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01 Botia dario juvenile – image by Ola Åhlander
02 Botia dario juvenile with unusual markings – image by Emma Turner
03 Botia dario semi-adult – image by Graeme Robson
04 Botia dario adult – image by Graeme Robson
05 Botia dario BMNH 1870.11.30.47, “North East Bengal” – image by Phil Hurst &
James Maclaine
06 Botia dario MNHN 1984-0344, a paratype of Botia macrolineata – image by
Claude Ferrara
07 Botia striata juvenile – image by Emma Turner
08 Botia striata adult – image by Emma Turner
09 Botia striata with unusual markings – image by Helen Nightingale
10 Botia striata NHSB uncatalogued specimen from Panchganga River, Kolhapur,
India – image by Varad Giri
11 Botia rostrata BMNH 1855.12.26.694, syntype of Botia rostrata – image by Phil
Hurst & James Maclaine
12 Botia rostrata BMNH 1860.3.19.114, syntype of Botia rostrata – image by Phil
Hurst & James Maclaine
13 Botia rostrata non preserved aquarium specimen, apparently from the Ganges
system – image by Emma Turner
14 Botia rostrata? from Shella, Meghalaya – image courtesy of Andrew Rao
15 Botia rostrata? from Shella, Meghalaya – image courtesy of Andrew Rao
16 Botia rostrata? from Shella, Meghalaya – image courtesy of Andrew Rao
17 Botia rostrata? from Shella, Meghalaya – image courtesy of Andrew Rao
18 Botia rostrata? from Shella, Meghalaya – image courtesy of Andrew Rao
19 Botia rostrata? from Shella, Meghalaya – image courtesy of Andrew Rao
20 Botia blythi? – image by Ken Childs
21 Botia sp. ‘Upper Brahmaputra’, juvenile – image by Emma Turner
22 Botia sp. ‘Upper Brahmaputra’, juveniles – image by Emma Turner
23 Botia sp. ‘Upper Brahmaputra’, approx. 12 months old – image by Bogdan
Janiczak
24 Botia sp. ‘Upper Brahmaputra’, adult, possibly female – image by Bogdan
Janiczak
25 Botia sp. ‘Upper Brahmaputra’, adult, possibly male – image by Bogdan Janiczak
26 Botia sp. ‘Upper Brahmaputra’, juveniles – image by Mark Duffill
27 Botia sp. ‘Upper Brahmaputra’, 2 of the juveniles pictured earlier after approx. 12
months – image by Mark Duffill
28 Botia sp. ‘Upper Brahmaputra’ Czech Republic produced hybrid with another
species? – image by Emma Turner
29 Botia sp. ‘Upper Brahmaputra’ Czech Republic produced hybrid with another
species? – image by Emma Turner
30 Botia almorhae BMNH 2002.9.18.1, holotype - image by Phil Hurst & James
Maclaine
31 Original drawing of Botia grandis = Botia almorhae
32 Botia almorhae BMNH specimen from Delhi - image by Phil Hurst & James
Maclaine
33 Botia almorhae BMNH specimen from Roorkee - image by Phil Hurst & James
Maclaine
34 Botia almorhae BMNH specimen from Jumna River - image by Phil Hurst &
James Maclaine
35 Botia lohachata OSUS specimens, Kali Gandaki River drainage – image by
David R. Edds
36 Botia lohachata live specimens, Kali Gandaki River drainage – image by David
R. Edds
37 Botia lohachata KU 28867, Narayani River, Nepal – image by Andrew Bentley
38 Botia lohachata? KU 29085, Seti River, Nepal – image by Andrew Bentley
39 Botia lohachata? KU 28754, Karnali River, Nepal, - image by Andrew Bentley
40 Botia lohachata? Adult aquarium specimen – image by Bogdan Janiczak
41 Botia sp. ‘Kosi’ KU 29099, Koshi River, Koshi Barrage, Nepal - image by
Andrew Bentley
42 Botia sp. ‘Kosi’ KU 29574, Koshi River, Koshi Barrage, Nepal - image by
Andrew Bentley
43 Botia sp. ‘Kosi’? BMNH specimen from Birbhum, India – image by Phil Hurst
and James Maclaine
44 Botia sp. ‘Teesta’ juvenile – image by Heok Hee Ng
45 Botia sp. ‘Teesta’ juvenile – image by Heok Hee Ng
46 Botia sp. ‘Teesta’? adult males? – image by Bogdan Janiczak
47 Botia birdi BMNH adult specimen from Chattabal, Srinagar, Kashmir – image by
Phil Hurst and James Maclaine
48 Botia birdi? BMNH juvenile specimen from Jove River - image by Phil Hurst and
James Maclaine
49 Botia birdi? BMNH adult specimen from Tawi River – image by Phil Hurst and
James Maclaine
50 Botia birdi? adult aquarium specimen – image by Bogdan Janiczak
51 Botia birdi? adult aquarium specimen – image by Bogdan Janiczak
52 Botia histrionica Sittang Form?, adult – image by Bogdan Janiczak
53 Botia histrionica Salween Form, adult (only fish on left) – image by Emma
Turner
54 Botia histrionica Irrawaddy Form CAS 88887 from Mandalay fish market,
Myanmar – image by Jon Fong
55 Botia histrionica Irrawaddy Form caught in the Irrawaddy River – image by
Rainer Hoyer
56 Botia histrionica Irrawaddy Form caught in the Irrawaddy River – image by
Rainer Hoyer
57 Botia histrionica Irrawaddy Form caught in the Irrawaddy River – image by
Rainer Hoyer
58 Botia histrionica Ataran Form – image by Kamphol Udomritthiruj
59 Botia histrionica Ataran Form – image by Kamphol Udomritthiruj
60 Botia histrionica x Chromobotia macracanthus Czech Republic juvenile hybrid –
image by Martin Thoene
61 Botia histrionica x Chromobotia macracanthus Czech Republic juvenile semiadult – image by Martin Thoene
62 Botia udomritthiruji juvenile - image by Kamphol Udomritthiruj
63 Botia udomritthiruji adult - image by Kamphol Udomritthiruj
64 Botia kubotai – image by Kamphol Udomritthiruj
65 Botia sp. ‘Ladder’ – image by Dave Speed
66 Syncrossus helodes Chao Phraya Form ANSP 56659 adult specimen from Doi
Angka, Chieng Mai, Thailand – image by Mark Sabaj
67 Syncrossus helodes Chao Phraya Form USNM 271701 juvenile specimens from
Thailand – image by Sandra Raredon
68
Syncrossus helodes Mekong Form – image by Ian Baird
69 Syncrossus helodes Mekong Form adult – image by Terry Warren
70 Syncrossus hymenophysa BMNH 1932.5.19.26 adult from Telom River, Pahang,
Peninsular Malaysia – image by Phil Hurst and James Maclaine
71 Syncrossus berdmorei AMS.B 7786, syntype – image by James King
72 Syncrossus berdmorei Irrawaddy Form juvenile caught in the Irrawaddy River –
image by Rainer Hoyer
73 Syncrossus berdmorei Irrawaddy Form semi-adult caught in the Irrawaddy River
– image by Rainer Hoyer
74 Syncrossus berdmorei ‘Central Myanmar’ – image by Kamphol Udomritthiruj
75 Syncrossus berdmorei Bagan Market - image by Kamphol Udomritthiruj
76 Syncrossus berdmorei Ataran Form, approx. 5 inches - Kamphol Udomritthiruj
77 Syncrossus berdmorei Ataran Form, approx. 7 inches – image by Kamphol
Udomritthiruj
78 Syncrossus berdmorei Ataran Form, approx. 7 inches – image by Kamphol
Udomritthiruj
79 Syncrossus sp.? ANSP 60087 from Chieng Mai, Thailand – image by Kyle
Luckenbill
80 Syncrossus beauforti ANSP 68005 holotype of B. lucasbahi – image by Mark
Sabaj
81 Syncrossus beauforti ANSP 56843 paratype of B. lucasbahi
– image by Mark Sabaj
82 Syncrossus beauforti? Specimen from Chao Phraya, possibly live specimen of
same species as fig. 79 - image by Kamphol Udomritthiruj
83 Syncrossus beauforti from Ataran River, Myanmar - image by Kamphol
Udomritthiruj
84 Syncrossus yunnanensis MIHW 638040 holotype – image by Zuogang Peng
85 Syncrossus formosus from Laos – image by Ian Baird
86 Holotype of Acanthocobitis longipinnis ZMB 4795 – image by Peter Bartsch
87 Holotype of Acanthocobitis longipinnis ZMB 4795, close-up of head – image by
Peter Bartsch
88 Acanthocobitis pavonacea – image by the author
89 Acanthocobitis pavonacea – image by the author
90 SMF 68, syntype of Cobitis pavonacea – image courtesy of Horst Zetzsche
91 SMF 9091, syntype of Cobitis pavonacea, close-up of head - image courtesy of
Horst Zetzsche
92 SMF 9092, syntype of Cobitis pavonacea, underside of head - image courtesy of
Horst Zetzsche