Osteology of some catfishes
of the genus Glyptothorax (Teleostei: Siluriformes) of northeastern India
W. Vishwanath 1, A. Darshan 2 & N. Anganthoibi 3
1, 3 Department of Life Sciences, Manipur
University, Canchipur, Manipur 795003, India
2 Directorate of Coldwater Fisheries Research,
Bhimtal, Uttarakhand 263136, India
Email: 1 wvnath@gmail.com, 2 achom_darshan@yahoo.com,3 angannong@gmail.com
Date of publication (online): 26 October 2010
Date
of publication (print): 26 October 2010
ISSN
0974-7907 (online) | 0974-7893 (print)
Editor: Heok Hee Ng
Manuscript
details:
Ms # o1874
Received 24
October 2007
Final received
22 July 2010
Finally
accepted 03 September 2010
Citation: Vishwanath, W., A. Darshan & N. Anganthoibi
(2010). Osteology of some catfishes of the genus Glyptothorax (Teleostei:
Siluriformes) of northeastern India. Journal
of Threatened Taxa 2(11):
1245-1250.
Copyright: © W. Vishwanath, A. Darshan & N. Anganthoibi
2010. Creative Commons Attribution 3.0 Unported License.JoTT allows unrestricted use of this article in any medium for non-profit
purposes, reproduction and distribution by providing adequate credit to the authors
and the source of publication.
Author
Details: A.
Darshan is a Post-Doctoral
Fellow of Department of Biotechnology, Govt of India and is at present attached
to the Directorate of Coldwater Fisheries Research, Bhimtal, Uttarakhand.
He is working on the phylogeny of catfishes based on classical and molecular
techniques.
N. Anganthoibiis a Research scholar, registered for PhD degree in the Department of Life
Sciences, Manipur University. She is working on the inventory of the catfishes
of northeastern India and their phylogenetic analysis.
W. Vishwanath is
a Professor in the Department of Life Sciences, Manipur University. His field
of specialization is fish and fisheries. He is presently engaged in taxonomy
and systematics of freshwater fishes of northeastern India.
Author
Contribution: The study: AD
- survey, collection, morphometric and anatomic study of catfishes of
northeastern India and their phylogenetics; NA - survey, collection, morphometric
and anatomic study of catfishes of northeastern India and their phylogenetics;
WV - supervision of taxonomy and phylogeny of freshwater fishes of northeastern
India.
Current paper:
AD - detailed examination of collections, comparision with specimens in ZSI and
preparation of drawings; NA - detailed examination of the collections of Glyptothorax in MUMF and osteological studies; WV -
supervision of the work, interpretation of the results and comparison with
available literature.
Acknowledgements: The work is supported by National Bureau of
Fish Genetic Resources (an ICAR Institute), Lucknow, Projecton inventory and phylogeny of catfish superfamily Sisoroidea of Northeast India.
Abstract: The
morphology of the premaxilla, dentary, Weberian lamina, infraorbital series,
vomer and frontal bones were observed in eight species of Glyptothorax of northeastern India. In G.
botius, G. granulus, G.
manipurensis, G. ngapang, G. striatus and G.
ventrolineatus, the
premaxilla consists only of proximal and distal tooth plates, the anterior
portion of the dentary is slender and its dorsal surface bears villiform teeth,
the lateral extension of posterior portion of Weberian lamina terminates at the
level of the lateral margin of its anterior portion, and the frontal has a
shallow orbital notch. In G. cavia and G.
chindwinica, the premaxilla
consists of proximal, distal and posterior elements on the roof of the oral
cavity; the anterior portion of the dentary bears posterior extension of
dentary tooth-plate; the lateral extension of the
posterior portion of Weberian lamina extends almost to the distal tip of the
fifth parapophysis; there are nine or ten infraorbital bones with a longer and
broader body of the lacrimal; greatly enlarged vomerine head; and frontal bears
a deep orbital notch. The jaw
structure of G.
burmanicus is discussed.
Keywords: Glyptothorax,osteology, Sisorid
fish, tooth plates
Abbreviation: MUMF - Manipur University
Museum of Fishes; dp
- distal element; prp - proximal element; pop - posterior element; vt -
villiform teeth; Pdtp - posterior extension of dentary bony tooth-plate; wl -
Weberian lamina, ppwl-posterior portion of Weberian lamina; cc - complex
centrum; vc5 - fifth vertebral centrum; ns4 - fourth neural spine; sop - spine
of supraoccipital; lac- lacrimal (first infraorbital); ion- infraorbital1-10; Af - anterior cranial fontanelle; afp - articular facet for palatine; bo
- bassioccipital; e - mesethmoid; epo - epeotic; exo - exoccipital; f -
frontal; le - lateral ethmoid; n - nasal; on - orbital notch; pf - posterior
cranial fontanelle; po - prootic; pt - pterotic; pts - pterosphenoid; scl -
supracleithrum; so - supraoccipital; s - sphenotic; v - vomer.
For figures, images -- click here
Introduction
Blyth (1860) described the genus Glyptothorax, for which Bleeker (1862-63) designated Glyptosternon striatus McClelland as the type species. Glyptothorax is the most species-rich and
widely distributed genus of the family Sisoridae. The genus has as many as 81 valid species (Ferraris 2007;
Gopi 2007; Vishwanath & Linthoingambi 2007; Ng & Freyhof 2008; Ng &
Kottelat 2008; Ng & Rainboth 2008). de Pinna (1996) casts doubt on the monophyly of
the genus Glyptothorax, citing the lack of unambiguous synapomorphies
to diagnose it. However, his
phylogenetic analysis of the Sisoroidea still recovered a monophyletic
Sisoridae.
Although there are reports on the osteological
studies of Glyptothorax, viz., Gauba (1966) on G. cavia (Hamilton) and Diogo et al. (2002) on G. fukiensis (Rendahl), there are no reports of the
comparative accounts on the osteology of the genus.
This study attempts to identify the variation
of bones i.e., premaxilla, dentary, infraorbital series, frontal, vomer and
articulation of complex centrum with remaining vertebral column in eight
species of Glyptothoraxof north east India, viz. Glyptothorax botius (Hamilton), G.
cavia (Hamilton), G. chindwinica Vishwanath & Linthoingambi. G. granulus Vishwanath & Linthoingambi, G. manipurensis Menon,G. ngapang Vishwanath &
Linthoingambi, G.
striatus (McClelland), andG. ventrolineatus Vishwanath & Linthoingambi. The results are presented in this study
and discussed.
Material and
Methods
Fresh specimens of Glyptothorax were collected from different areas of
northeastern India. Measurements
of antero-posterior length and lateral-extent of upper jaw tooth band follow
Vishwanath & Linthoingambi (2007). Specimens were cleared and stained following Hollister (1934). Terminology and nomenclature of bones
follow Chen & Lundberg (1995) and de Pinna (1996). Components of premaxillary bones are
named as proximal, distal and posterior elements based on their positions.
Visible jaw structures of holotype of Glyptothorax burmanicus in ZSI Kolkata was also observed and compared.
RESULT
Premaxilla: In six
species, viz., Glyptothorax botius, G. granulus,G. manipurensis, G. ngapang,G. striatus, and G. ventrolineatus, the premaxilla (Figs. 1A & 1B) consists of
four bony elements, i.e., a pair of medially located proximal elements (prp)
and another pair of distal elements (dp), located lateral to the proximal
element. The
convex anterior margin of the distal element articulates with the corresponding
concavity along the posterolateral margin of the proximal element. The distal element is firmly united to
the proximal end by rigid connective tissue. However, in G.
cavia and G. chindwinica, the premaxilla comprises of numerous distal
(Fig. 1D: dp) and posterior elements (Fig. 1D: pop) in addition to a pair of
proximal elements (Fig. 1D: prp). The distal and posterior elements are attached respectively to the
lateral and the posterior parts of the proximal elements. All the subunits of the proximal,
distal and posterior elements are tightly fastened by connective tissue to form
the premaxilla. The sutures among
the elements that comprise the premaxilla are not visible ventrally because the
presence of numerous villiform teeth (Fig. 1C: vt) on this surface obscures
it. Ventrally the premaxilla is
seen as a single structure on the roof of the oral cavity (Fig. 1C).
Dentary: In the six
species, viz., Glyptothorax botius, G. granulus,G. manipurensis, G. ngapang,G. striatus, and G. ventrolineatus, the anterior portion of the dentary is slender,
with villiform teeth on its dorsal surface (Figs. 2A & 2B). However in G. cavia andG. chindwinica, the
dentary is a stout, long, curved bone with a very broad anterior two-third
portion. This broadness is
manifested by a posterior extension of the tooth-bearing surfaces on the
dentary (Fig. 2C & 2D: pdtp) which bears villiform
teeth at its dorsal surface. Gauba
(1966) also reported the tooth- bearing portion of the dentary to be very much
flattened in G. cavia. The dentary tooth plate of G.
burmanicus is also similar
to that of G. cavia and G.
chindwinica.
Posterior portion of Weberian lamina: The transverse process or parapophysis of
complex centrum was named the Weberian lamina by de Pinna (1996). In the present study, all the examined
species of Glyptothorax(except, G. cavia andG. chindwinica) have an
extended posterior portion of Weberian lamina (Figs. 3B & 3C: ppwl) to the
level of lateral margin of its anterior portion. In G.
botius, there is a
large rounded space between the articulation of the fifth parapophysis and the
posterior portion of the Weberian lamina (Fig. 3D). Tight suturing is also observed at the adjoining parts of
the space. In G. cavia andG. chindwinica, the
lateral expansion of the posterior portion of Weberian lamina (Fig. 3A: ppwl)
extends beyond the lateral margin of the anterior portion of the lamina,
reaching almost to the distal tip of the parapophysis of the fifth vertebra.
Infraorbital: The number of infraorbital bones is variable. Glyptothorax botius has six infraorbital bones; of which the sixth
is longest while the fifth, the shortest. Both G.
ngapang and G. chindwinica have nine infra-orbital bones, while G. cavia has ten. In the remaining species, there are eight bones in the series. The thirdinfra-orbital in G.
ngapang (Fig. 4B: io3)
bears a broad ventral laminar process. Both G. cavia and G.
chindwinica have a larger and
broader body of the lacrimal (Fig. 4A: lac) when compared to other species
examined.
Orbital notch: The orbital notch of the Glyptothorax is formed at the lateral margin of frontal as a
shallow depression, forming an arc smaller than a semicircle (Fig. 5C:
on). G. honghensis Li (Zhou & Zhou 2005: Fig. 6B), G. fukiensis (Diogo et al. 2002: Figs.1 & 2) and G. major (de Pinna 1996: Fig. 13) have also shallow
orbital notches. However, in G. cavia and G.
chindwinica, the notch is
deep and forms an arc larger than a semicircle (Figs. 5A & 5B: on).
Vomer: The
head of the vomer of Glyptothorax is edentulous and extended laterally along
the entire length of the articular process of lateral ethmoid, reaching the
articular facet for palatine at the lateral tip of each lateral ethmoid. In all
the species examined (except in G.
cavia and G. chindwinica), the anterolateral margin of the head of the
vomer is concave resulting in the formation of a thin lateral process and
another sharply pointed medial anterior tip (Fig. 6 B & C). In G.
cavia and G. chindwinica, the head of vomer is very large
and broad with roughly convex anterior margin (Figs. 5A & 5B: v; 6A).
Discussion
Among the Siluriformes, the premaxilla of Glyptothorax (Tilak 1963; de Pinna 1996) and Bagarius Bleeker (Gauba 1962) is characteristic in
having separate distal elements connecting laterally to the proximal element.
Diogo et al. (2002) also reported the same structure in G.
fukiensis (Rendahl). A subdivided premaxilla has also been
reported in Glyptosternum
reticulatum McClelland
(Gauba 1969), and fragmentation of the distal element of
premaxilla into tripartite or multipartite structures have also been
reported in Euchiloglanis
kishinouyei Kimura (de Pinna
1996). This study shows that structure of premaxilla in most of the examined species havea pair of medially located proximal elements and another pair of distal
elements situated laterally. However,
the premaxilla in G.
cavia and G. chindwinica is markedly different, consisting of numerous
posterior elements and distal elements in addition to proximal elements. Gauba (1966) recorded the premaxilla ofG. cavia as being generally segmented or fused to form
an enormously broad band that extends a considerable distance posteriorly
across the palate. However, he
failed to notice the numerous individual tooth plates tightly attached by
connective tissue.
The holotype of Glyptothorax burmanicus Prashad & Mukherji (Image 1A) has been
examined. It has a central depression
in the thoracic adhesive apparatus (Image 1B) and a premaxilla in the form of a
broad patch with minute villiform sharp teeth and a dentary with broad
teeth-bearing plate (Image 1C), the characters, similar to those of G. cavia and G.
chindwinica.
Among the representatives of family Sisoridae,
the posterior portion of the Weberian lamina is reported to extend along the
anterior margin of the fifth vertebra in Bagarius, GagataBleeker, Glyptosternoids, Glyptothorax, Nangra (Day), Pseudecheneis Blyth and Sisor Hamilton (de Pinna 1996). This study indicates that the extension
of the porterior portion of the Weberian lamina is not equal within the genus Glyptothorax. In the species under study, the posterior portion of the Weberian lamina
is extended laterally to the level of the lateral margin of its anterior
portion, except in case of G.
cavia and G. chindwinica. A similar condition has been reported in G. major (de Pinna 1996: Fig. 26A) and in G. honghensis Li (Zhou & Zhou 2005: Fig. 6B). In G.
cavia and G. chindwinica, the lateral expansion of the posterior
portion of the Weberian lamina is long and reaches almost to the distal tip of
the parapophysis of the fifth vertebrae.
Large variations in osteological characters of
the Weberian lamina, infraorbital series, the shapes of the vomer and frontal
have been observed among the members of the genus. However, pending examination of more species of the genus,
it is not possible to establish the paraphyly of Glyptothorax. This study will help future workers to some extent in the study of
phylogenetic relationships within Glyptothorax.
Comparative Materials
Glyptothorax striatus: Uncat., 2 exs.,
79.7-83.0 mm SL, ICAR Complex for northeastern region, Barapani, Meghalaya,
India, coll. B.K. Mahapatra; 31.x.2005, 4 exs. 40.2-123 .9 mm SL, Siren River,
Rotung, East Siang District, Arunachal Pradesh, India, coll. K. Nebeshwar, MUMF
9040.
Glyptothorax cavia: 2 exs. 86.4-98.0 mm SL, left Bank of Kosi
River, two furlongs down the confluence with the Arun River at Tribeni, Nepal,
Kosi survey, F218/2; 2 exs., 82.8-80.3 mm SL, Same data, F219/2; 06.xi.2006, Uncat., 1
ex. 87.5 mm SL, Barak River, Tamenglong District, Manipur,
India, coll. Kingson.
Glyptothorax manipurensis: 10.xii.1998, 10 exs.,69.0-104.0 mm SL, Barak river, Vanchengphai, Manipur, India, coll. K.
Nebeshwar, MUMF 4029-4032.
Glyptothorax ventrolineatus: 15.i.2003, holotype, 85.8mm SL, Iril River,
Ukhrul District, coll. I. Linthoingambi, MUMF L0221; 5 exs.,Paratypes, 85.1-94.5 mm SL, data same as holotype, MUMF L0222/5; 10.iv.2003, 4
exs., 67.2-83.2 mm SL. Lokchao River, Moreh, Chandel District, Manipur, India,
coll. K. Nebeshwar and party, MUMF 4300/4.
Glyptothorax ngapang: 06.vi2001, holotype, 82.7mm SL, Iril River,
Bamonkampu, Manipur, India, coll. I. Linthoingambi, MUMF 6131; paratypes, 9
exs, 61.7-99.5mm SL, same data as holotype, MUMF 6132; 10.iv.2003, 65.0-98.5mm
SL, 10 exs., Lokchao River, Moreh (Indo-Myanmar
border), coll. W. Vishwanath, MUMF 6141.
Glyptothorax granulus: 10.i.2004, holotype, 76.6mm SL, Iril River,
Phungdhar, Ukhrul District, Manipur, India, coll. I. Linthoingambi, MUMF 6151;
06.vi.2003, paratypes, 10 exs., 61.7-76.6mm SL, same data as holotype, MUMF
6152; 12.xi.2003, 1ex., 96mm SL., Iril River, Urup, Manipur, India (Chindwin basin), coll. Linthoi, MUMF
9991; 03.iv.2004, 10exs., 80.5-89.8 mm SL Lokchao river, Moreh (Indo-Myanmar
border), coll. W. Vishwanath, MUMF 6156.
Glyptothorax chindwinica: 26.viii.2002, holotype, 145.4mm SL, Iril
River, Urup, Manipur, India, coll. I. Linthoingambi, MUMF 6366; 03.iv.2004,
115.6-145.5 mm SL, paratypes, 4 exs., Lokchao River,
Moreh (Indo-Myanmar border), Chandel District, Manipur, India, coll. W.
Vishwanath, MUMF 6368; 15.i.2004, 5 exs, 100.2-123.6 mm SL, Thoubal River,
Nongpok Keithelmanbi, Thoubal District, Manipur, India, coll. I. Linthoingambi,
MUMF 6369.
Glyptothorax burmanicus: holotype, 100.8mm SL, Sankha,
a large hill-stream, midway between Kamaing and Mogaung, Myitkyina District,
Upper Myanmar, coll. Dr. B.N. Chopra, ZSI F10877/1.
Glyptothorax botius: 16.iii.2006, 75.2mm SL, Dibru
River, Dibrugarh, Assam, India, coll. Santosh, MUMF 9520.
REFERENCES
Bleeker, P.
(1862-63). Atlas ichthyologique des Indes Orientales Néêrlandaises,
publié sous les auspices du Gouvernement colonial néêrlandaises. Tome II. Siluroides, Chacoïdes et Hétérobranchoïdes. Amsterdam, 112pp.
Blyth, E. (1860). Reports on some fishes received chiefly from
the Sitang River and its tributary streams, Tenasserim Provinces. Journal of the Asiatic Soceity of
Bengal 29: 138-174.
Chen, X. & J.G. Lundberg (1995). Xiurenbagrus, a new genus of Amblycipitid catfishes
(Teleostei: Siluriformes), and phylogenetic relationships among the genera of
Amblycipitidae. Copeia 1995(4): 780-800.
de Pinna, M.C.C. (1996). A phylogenetic analysis of the Asian catfish
families Sisoridae, Akysidae, and Amblycipitidae, with a hypothesis on the
relationships of the Neotropical Aspredinidae (Teleostei: Ostariophysi). Fieldiana Zoology (New
series) 84: 1-83.
Diogo, R.,
M. Chardon & P. Vandewalle (2002). Osteology and mycology of the cephalic region and pectoral
girdle of Glyptothorax
fukiensis (Rendahl, 1925), comparison with other sisorids, and
comments on the synapomorphies of the Sisoridae (Teleostei: Siluriformes). Belgium Journal of
Zoology 132(2): 95-103.
Ferraris, C.J., Jr. (2007). Checklist of catfishes,
recent and fossil (Osteichthyes: Siluriformes), and catalogue of Siluriform
primary types. Zootaxa 1418: 1-628.
Gauba, R.K. (1962). The endoskeleton of Bagarius bagarius (Hamilton), part I- The skull. Agra
University Journal of Research11(1): 75-90.
Gauba, R.K. (1966). Studies on the osteology of
Indian sisorid catfishes II. The skull of Glyptothorax cavia. Copeia4: 802-810.
Gauba, R.K. (1969). The head skeleton of Glyptosternum reticulatumMcClelland & Grifith. Monitore Zoologico Italiano 3: 1-17.
Gopi, K.C.
(2007) Pisces,
pp.115-131. In: The
Director, Zoological Survey of India, Kolkata (ed.), Conservation Area Series, 32, Fauna of Kudremukh National Park (Karnataka). Zoological Survey of India, Kolkata.
Hollister, G. (1934). Clearing and dying fishes for bone study. Zoologica 12: 89-101.
Menon, A.G.K. (1999). Checklist- Freshwater fishes of India. Records of the Zoological Survey of India, Miscellaneous
Publications, Occasional Paper 175, xxviii+366pp.
Misra, K.S.
(1976). The Fauna
of India and The Adjacent Countries, Pisces 3. Teleostomi: Cypriniformes: Siluri, (2nd Ed.), Zoological
Survey of India, Calcutta, xxi+367pp.
Ng, H.H.
& J. Freyhof (2008). Two new
species of Glyptothorax(Teleostei: Sisoridae) from
central Vietnam. Zootaxa 1873: 11–25.
Ng, H.H. & M.
Kottelat (2008). Glyptothorax
rugimentum, a new species
of catfish from Myanmar and western Thailand (Teleostei: Sisoridae). The Raffles Bulletin of Zoology 56(1): 129-134.
Ng, H.H.
& W.J. Rainboth (2008). Glyptothorax
coracinus, a new species
of hillstream catfish from Cambodia (Teleostei: Sisoridae). Zootaxa 1839: 60–68.
Thomson,
A.W. & L.M. Page (2006). Genera of the Asian catfish families Sisoridae and
Erethistidae (Teleostei: Siluriformes). Zootaxa 1345: 1-96.
Tilak, R. (1963). The osteocranium and the Weberian apparatus of
the fishes of the family Sisoridae (Siluroidea): a study in adaptation and
taxonomy. Zeitschrift
für Wissenschaftliche Zoologie169: 281-320.
Vishwananth,
W. & I. Linthoingambi (2007). Fishes of the genus Glyptothorax Blyth
(Teleostei: Sisoridae) from Manipur, India, with description of three new
species. Zoos’
Print Journal 22(3):
2617-2626.
Zhou, W. & Y.W. Zhou (2005). Phylogeny of the genus Pseudecheneis (Sisoridae) with explanation of its
distribution pattern. Zoological
Studies 44: 417–433.