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Fishes of the Khuiala Formation (Early Eocene) of the Jaisalmer Basin,

Western Rajasthan, India

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Fishes of the Khuiala Formation
(Early Eocene) of the Jaisalmer Basin,
Western Rajasthan, India
Kishor Kumar1,*, Rajendra Singh Rana2 and
Hukam Singh3
1
Wadia Institute of Himalayan Geology, 33 General Mahadeo Singh
Road, Dehradun 248 001, India
2
Department of Geology, H.N.B. Garhwal University,
Srinagar (Garhwal) 246 174, India
3
Birbal Sahni Institute of Palaeobotany, 53 University Road,
Lucknow 226 007, India
A newly discovered assemblage of the Early Eocene
(Ypresian) fish is recorded from the lower part of the
Khuiala Formation of the Jaisalmer Basin, Western
Rajasthan. It consists of exceptionally well preserved
but mostly isolated teeth and some dermal denticles of
fairly diverse ichthyofauna. Based on isolated teeth, 14
taxa belonging to six elasmobranch and five teleostean
families have been identified, including a probably new
but unnamed species of the nurse shark, Ginglymo-
stoma. Other taxa comprise Galeorhinus sp., Hemiscyl-
lium sp., Rhinobatos sp., Gymnura sp., Heterotorpedo
sp., Dasyatis sp. 1, Dasyatis sp. 2, Coupatezia sp., Sphy-
raena sp., Eutrichiurides sp., Sparus sp., Stephanodus
lybicus and Diodon sp. Among these, Hemiscyllium and
Coupatezia are new to the subcontinent. Among the
Palaeocene–Eocene ichthyofaunas documented from
India, this Khuiala assemblage is closest to the Lower
Eocene fish assemblage from the subsurface Cambay
Shale of Vastan Lignite Mine, Surat District, Gujarat.
It is a dominantly shallow-water marine assemblage
typically lacking coastal freshwater elements.
Keywords: Early Eocene, ichthyofauna, Jaisalmer Basin,
Khuiala Formation, vertebrates.
T HE Rajasthan shelf lying to the west of the Aravalli
ranges comprises three major sedimentary basins, namely
Jaisalmer, Barmer and Bikaner-Nagaur (also referred to
as Palana-Ganganagar). All these basins have a good record
of coeval Palaeogene sequences that consist mainly of
shallow marine sediments and are fairly richly fossilifer-
ous1–5. Assemblages of foraminifers, ostracodes, molluscs
and palynofossils have been known from these sedi-
ments1,6–16 . However, studies of vertebrate faunas from
these sequences have picked up only recently, though their
occurrences were noticed much earlier 17,18 . The Jaisalmer
Basin, in particular, has remained poorly studied from the
viewpoint of vertebrate fauna, whereas some lower verte-
brate fossil assemblages have been documented from
Barmer and Bikaner-Nagaur basins in recent years19–22 .
To fill this lacuna, two of the authors (R.S.R. and H.S.)
recently undertook a preliminary sampling in the Khuiala
*For correspondence. (e-mail: kumark@wihg.res.in)
CURRENT SCIENCE, VOL. 93, NO. 4, 25 AUGUST 2007
RESEARCH COMMUNICATIONS
Formation for microvertebrate prospecting. Our initial
attempt has shown encouraging results and we have deve-
loped a small assemblage of lower vertebrates for this
contribution.
In Jaisalmer Basin, the Palaeogene succession is repre-
sented by the Sanu, the Khuiala and the Bandha forma-
tions in ascending order3–5. The Sanu Formation (Palaeo-
cene) unconformably overlies the Habur Formation of an
Early Cretaceous (Aptian) age4. It has a gradational contact
with the overlying Khuiala Formation (Early Eocene), which
has well documented existence in the surface (outcrops) as
well as subsurface3,4 . The Khuiala Formation comprises
four members, namely (in ascending order) Te Takkar, Lower
Khinsar, Sirhera and Upper Khinsar 3,4 . The Te Takkar
Member is a dominantly foraminiferal, brownish lime-
stone facies with intercalation of greenish clay and argil-
laceous limestone. The overlying Lower Khinsar Member is
represented by shale with occasional intercalation of marl.
The Sirhera Member comprises alternations of shale and
argillaceous limestone. The topmost member of the Khuiala
Formation, i.e. the Upper Khinsar is argillaceous and
characterized by silty shale facies with thin intercalations
of marl and limestone. The Khuiala Formation is overlain
by the Bandha Formation of Middle Eocene (Lutetian) age.
Foraminifers and ostracodes are the most common and
well documented from the Khuiala Formation, though
bivalves, gastropods, cephalopods, echinoids, worms and
palynofossils are also known1,3,4,10–15.
Prior to this report, the Khuiala Formation was never
studied for its vertebrate faunal content, though notices of
unidentified fish teeth were mentioned by several workers
from its youngest member, i.e. the Upper Khinsar Shale3,4.
This contribution reports fish fauna from its oldest unit
(Te Takkar Member), and is a systematic record of Eocene
vertebrates from the Jaisalmer Basin.
The fossils were recovered by screen-washing of over
50 kg of rock matrix consisting of yellow to yellowish-
white calcareous shale/marl of the Te Takkar Member of
the Khuiala Formation exposed near a hill, about 4 km west
of Habur Village (lat. 27°10′N, long. 70°33′E) and 8 km
northeast of Khuiala Village (lat. 27°8′N, long. 70°26′E),
Jaisalmer District. At the fossiliferous site the base of the
Khuiala Formation is not exposed. The vertebrate fossil-
yielding horizon underlies the pinkish-white to buff
foraminiferal limestone that marks the top of the Te Takkar
Member, and also yields foraminifers besides vertebrates
(Figure 1).
The dental remains of fish described here are repre-
sented by the elasmobranch (sharks and rays) families
Triakidae (hound sharks), Ginglymostomatidae (nurse
sharks), Hemiscylliidae (bamboo sharks), Dasyatidae
(stingrays), Gymnuridae (butterfly rays), Rhinobatidae
(guitar fishes) and Torpedinidae (electric rays), and the
teleostean families Sphyraenidae (barracudas), Trichiuridae
(hairtails, cutlass and ribbon fishes), Sparidae (sea breams
or porgies) and Diodontidae (porcupine or burrfishes)
553
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Figure 1. Geological map of Khuiala–Sanu–Habur areas, Jaisalmer District showing location of verte-
brate fossil locality (a), and lithological log of the fossiliferous section (b). Geological map modified
after Bafna and Dhaka46.
and Trigonodontidae. A probably new but unnamed species
of nurse shark, Ginglymostoma and reports of Hemiscyllium
and Coupatezia from India are significant additions to the
ichthyofauna. Other taxa in the assemblage comprise
Galeorhinus sp., Rhinobatos sp., Gymnura sp., Hetero-
torpedo sp., Dasyatis sp. 1, Dasyatis sp. 2, Sphyraena sp.,
Eutrichiurides sp., Sparus sp., Diodon sp. and Stephanodus
lybicus. Many of these taxa have been earlier known from
the coeval beds in northwest sub-Himalaya, Gujarat and
other parts of Rajasthan (India), but not from the Jaisalmer
Basin.
All the specimens described and illustrated here are
registered (KV/R/GU Khuiala Vertebrates/Rajasthan/
Garhwal University) and housed in the Department of
Geology (collection of R. S. Rana), H.N.B. Garhwal Uni-
versity, Srinagar (Garhwal), India.
Ginglymostoma sp. (Figure 2: 1–12): Anterior teeth
(Figure 2: 1–4), fan-shaped, higher than wide; main cusp
high flanked by three to six pairs of low, sharp lateral
cusplets; heels oblique; labial side slightly convex, lingual
side flattened with a high protuberance; apron has a dis-
tinct central prolongation overhanging the root but not
reaching its base; root lobes unequal and slipper-shaped;
basal face of root flat and V-shaped; central foramen large
and elliptical; margino-lingual and labial foramina
smaller. Lateral teeth (Figure 2: 5–8) nearly as high as
broad with fewer lateral cusplets; root lobes obtusely spread
(as against V-shaped in anterior teeth). Posterior teeth
554
(Figure 2: 9–12), shorter than broad or as broad as high
with a small, posteriorly bent main cusp flanked by lateral
cusplets; apron broad and roundish; medio-lingual protu-
berance moderately high; root lobes roundish and thin.
The anterior teeth have general morphological resem-
blance with several known species of Ginglymostoma23–25,
but differ in being higher than wide, having a high and
straight main cusp and three to six pairs of thin lateral
cusplets, and likely represent a new species. Posterior
teeth are similar to those of G. sokotoense and G.
maghrebianum from Thanetian of Nigeria and Ypresian
of Morocco and Tunisia 23,24,26 , but differ in size. Teeth
differ from Ginglymostoma sp. of the Late Palaeocene–
Early Eocene, Akli Formation of Barmer (Rajasthan) in
having fewer lateral cusplets and in less prominent lin-
gual protuberance19 . Ginglymostoma is widely known
from Cretaceous to Miocene of Europe, North America,
India and Africa; presently it thrives in tropical Atlantic
and Indo-Pacific Sea 25 .
Hemiscyllium sp. (Figures 2: 13–16): Teeth wider than
high with a large and stout main cusp. In the smaller of
the two teeth in the collection (Figure 2: 13, 14), the main
cusp is short and more conical with well differentiated
lateral heels, each having a pair of massive but divergent
lateral cusplets well separated from each other. In the
other tooth (Figure 2: 15, 16), the main cusp is higher
without well-differentiated heels and there are faintly de-
veloped blunt lateral cusplets. Lingual surface is convex with
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Figure 2. 1–12: Ginglymostoma sp. – 1, 2: KV/R/GU 501; 3, 4: 502, anterior teeth in labial (1, 3) and lingual (2, 4) views; 5: 503; 6: 504;
7, 8: 505, lateral teeth in lingual (5, 8) and labial (6, 7) views; 9, 10: 506; 11, 12: 507, posterior teeth in labial (9, 11) and lingual (10, 12)
views; 13–16: Hemiscyllium sp. – 13, 14: 508; 15, 16: 509, teeth in occlusal (13), lingual (14, 15) and labial (16) views; 17–26: Galeo-
rhinus sp. – 17, 18: 510; 19, 20: 511; 21: 512; 22: 513; 23, 24: 514; 25, 26: 515; lateral (17, 18), posterior (19–22) and anterior (23–26)
teeth in lingual (17, 19, 21, 23, 25) and labial (18, 20, 22, 24, 26) views; 27–32: Rhinobatos sp. – 27, 28: 516; 29, 30: 517; 31, 32: 518,
teeth in lingual (27, 29, 31) and labial (28, 30, 32) views; 33–42: Dasyatis sp. 1 – 33–35: 519; 36, 37: 520; 38: 521; 39: 522, teeth of
female individuals in lingual (33, 37–39) and labial (34–36) views; 40, 41: 523; 42, 524, teeth of male individuals in lingual (40, 42) and
labial (41) views. Scale bar in all figures equals 1 mm.

a distinct but short and wide apron having a central notch
as in H. bruxelliensis. The notch is more pronounced in
the tooth with lateral cusplets; lingual protuberance is so
prominent that its upper part is produced like an enameloid
rod and horizontal section of the root has a trilobed ap-
CURRENT SCIENCE, VOL. 93, NO. 4, 25 AUGUST 2007
pearance; root short and triangular with flattened basal
face; central foramen and margino-lingual foramina present.
Teeth have close similarities with H. bruxelliensis from
the Upper Ypresian of Belgium27 . Prior to this there was
no record of Hemiscyllium from India.
555
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Galeorhinus sp. (Figure 2: 17–26): Lateral teeth (Figure
2: 17, 18) nearly as high as wide; posterior teeth (Figure
2: 19–22) wider than high with posteriorly inclined cusp;
mesial cutting edge of posterior and lateral teeth long,
slightly sigmoid due to upturning of cusp (one tooth shown
in Figure 2: 19, 20 has the main cusp downturned like in
Galeocerdo); lower two-thirds of mesial cutting edge
finely serrated; distal edge shorter and smooth; distal heel
has one to three cusplets; labial face of crown slightly
convex with a moderately pronounced, nearly horizontal
ridge overhanging the root; lingual face of crown convex
with a broad, bilobed root having flat (e.g. Figure 2: 17,
19, 22) or slightly convex (e.g. Figure 2: 24, 25) base and
well-differentiated groove between the two lobes. Anterior
teeth (Figures 2: 23–26) have nearly vertical main cusp
with smooth cutting edges; mesial heel has one or two
cusplets; root massive. Anterior teeth look specifically
similar to Galeorhinus sp. from the Early Eocene of Gu-
jarat 28 and Rajasthan 19,20 . Teeth of G. minutissimus, G.
gomphorhiza and G. latus from the Tertiary of Africa24,29,
Belgium 30–32 , and USA33 are also similar to those of the
Khuiala species, but differ in possessing more erect and
higher main cusp.
Rhinobatos sp. (Figure 2: 27–32): Lateral teeth wider
than long; occlusal surface smooth with a distinct trans-
verse ridge; three uvulae (enamel prolongations) overhang
the root on lingual face of teeth; central uvula longer and
globular; lateral uvulae shorter but well-separated from
central one; root massive and displaced lingually; root
lobes subtriangular, becoming narrower lingually. Teeth
are closely similar to Rhinobatos sp. from Lower Eocene of
Gujarat28 and Rhinobatos sp. 1 from Rajasthan20. Rhinobatos
sp. recorded from the Lower Eocene beds of the Subathu
Formation, Himachal Pradesh has more convex occlusal
surface with a more prominent transverse ridge34 . R.
casieri from Cretaceous of New Jersey has teeth with
pointed central uvula35 .
Dasyatis sp. 1 (Figure 2: 33–42): Teeth of females
(Figure 2: 33–39) slightly longer than wide with distinct
visors and a transverse crest; labial zone of crown slightly
convex with scallop ornamentation and a slight ridge par-
allel to labial edge in some teeth; lingual zone smooth
with a distinct median visor hanging over the root; median
lingual ridge lacking and marginal faces without depres-
sions; marginal angles well marked; root shallow and
bilobed with a wide groove; basal face of root lobes tri-
angular and subequal; central foramen pronounced and
situated on labial side of root. Teeth of male individuals
(Figure 2: 40–42) differ in having a prominent projection
or cusp on lingual side of crown. Teeth are closely similar
to those described as Dasyatis sp. 1, from the Early Eocene
Cambay Shale, Gujarat and as Dasyatis sp. from the Ka-
purdi Formation, Barmer, Rajasthan 20,28 ; but in the latter
species crown is more convex and slightly bigger in size.
Dasyatis sp. from the Late Eocene deposits of the Paris
Basi36 has morphologically similar but considerably larger
teeth.
556
Dasyatis sp. 2 (Figure 3: 1–8): Teeth differ from Da-
syatis sp. 1 in having a labial zone with an elliptical and
smooth central depression and a crenulated transverse
ridge parallel to the labial edge. At least three teeth in the
collection are smaller and much wider than long with
elliptical oral face (Figure 3: 4–8), and one of these has a
deep transverse groove between the lingual edge of the
oral face and the lip (Figure 3: 7, 8).
Gymnura sp. (Figure 3: 9–12): Teeth have moderately
high, broad and more or less smooth crown with a lin-
gually directed cusp having marginal crests meeting at
lateral edges; transverse crest semicircular, joins lateral
angles, which are roundish but acute and labially directed
forming prominent margino-labial protuberances; labial
contour of crown concave in occlusal view, also having a
small median protuberance; lingual face of crown broad
with lateral depressions and slightly notched lower margin;
root bilobed with a deep groove. Gymnura sp. from Palaeo-
cene–Early Eocene Akli Formation of Rajasthan differs
from Khuiala species in having larger teeth with much
more concave labial contour of crown, a prominent trans-
verse crest and in lacking a median protuberance19 . The
Khuiala species differs from Gymnura sp. of the Thanetian,
Morocco25,37 in possessing a median protuberance in addi-
tion to two margino-labial protuberances on the labial
contour of the crown.
Heterotorpedo sp. (Figure 3: 13–17): Teeth strongly
cuspidate, labial edge concave in the middle with two
labially directed lateral expansions corresponding to lateral
angles; occlusal face of crown has irregular, scalloped
ornamentation on labial side and marked by a labio-
lingual groove; transverse crest sharp and limited to basal
part of cusp; root low, lingually displaced and divided by
a prominent groove. The teeth look to be specifically similar
to teeth described as Heterotorpedo sp. from the Cambay
Shale, Surat, Gujarat and from the Kapurdi Formation,
Barmer District, Rajasthan 20,28 . They also have close
morphological affinities with H. fowleri from the Palaeo-
gene of Sussex38 .
Coupatezia sp. (Figure 3: 18–21): Lateral teeth of females
transversely elongated with elliptical crown more than
twice broader than long; labial face bears a transverse crest
sub-parallel to a grooved labial crown edge; transverse
crest finely crenulated, laterally joins the labial edge;
marginal angle blunt; lingual face steeper, slightly higher
than labial face; occlusal surface concave and smooth;
root moderately high, bilobed, slightly shifted lingually;
with a moderately deep groove and a small foramina; basal
face of root triangular and nearly flat. This is the first re-
port of Coupatezia from India. C. woutersi from the Mid-
dle Eocene of Belgium 25,39 has larger teeth with less
length/width ratio and widely notched labial contour.
Sphyraena sp. (Figure 3: 22–23): Teeth labio-lingually
compressed with acute apex and distinct pulp cavity; sur-
face and edges smooth; lingual side flattened, labial side
convex, more so in lower part of tooth. S. fayumensis from
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Figure 3. 1–8: Dasyatis sp. 2 – 1: 525; 2: 526; 3: 527; 4, 5: 528; 6: 529, 7, 8: 530, teeth of female individuals in lingual (1–4, 7) and
labial (5, 6, 8) views; 9–12: Gymnura sp. – 9, 10: 531; 11, 12: 532, teeth in lingual (9, 12) and labial (10, 11) views; 13–17: Heterotorpedo
sp. – 13–15: 533; 16, 17: 534, teeth in occlusal (13, 14, 17) and basal (15, 16) views; 18–21: Coupatezia sp. – 18, 19: 535; 20, 21: 536,
teeth in lingual (18, 20) and labial (19, 21) views; 22, 23: Sphyraena sp. – 22: 537; 23: 538, teeth in lateral views; 24–29: Sparus sp. – 24:
539; 25: 540; 26: 541; 27: 542; 28: 543; 29: 544, teeth in lateral views; 30–34: Eutrichiurides sp. – 30: 545; 31: 546; 32: 547; 33: 548; 34:
549; 35, 36: Stephanodus lybicus – 35: 550; 36: 551, pharyngeal teeth in lateral views; 37, 38: Diodon sp. – 37: 552; 38: 553, fragments of
dental plates in lateral views. Scale bar in all figures equals 1 mm.

Lutetian of northern Africa 24 and Sphyraena sp. from
Cambay Basin 28 and Kutch 40 have similar teeth but with
serrated edges.
Sparus sp. (Figure 3: 24–29): Teeth vary in size and
shape; generally short and conical, slightly curved; apex
small, pointed to blunt with smooth basal surface; some
globular teeth with flat and smooth occlusal surface (Figure
3: 29) are comparable to Egertonia reported from Guja-
rat 28 . Sparus sp. from the Lower Eocene of Morocco has
similar teeth 24 .
Eutrichiurides sp. (Figure 3: 30–34): Teeth conical,
some laterally compressed, more or less smooth; apex

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about one-tenth of tooth height and pointed; basal part
nearly rounded with fine vertical lines; some teeth have
smooth surface. Eutrichiurides species from the Lutetian
of northern Africa 24 and Palaeocene Cannonball Forma-
tion of South Dakota have some resemblance with Khui-
ala teeth, but differ in size41 . Eutrichiurides teeth are also
known from the Early Eocene Cambay Shale of Gujarat28.
They differ from the Khuiala teeth in being slightly laterally
compressed near the base and in having distinct vertical
ridges and grooves.
Stephanodus lybicus (Figure 3: 35, 36): Pharyngeal
teeth have a large, hook-like principal cusp and a deep
root. Secondary cusps are lacking. S. lybicus is widely
known in Late Cretaceous–Middle Eocene sediments of
India and has been reported from Infra and Intertrappean
(Cretaceous–Palaeocene) beds, Fatehgarh Formation (Pa-
laeocene) of Rajasthan, Cambay Shale (Early Eocene) of
Gujarat and from the Kakara–Subathu succession (Pa-
laeocene–Middle Eocene) of sub-Himalaya 21,28,34,42–44 .
Diodon sp. (Figure 3: 37, 38): Fragments of dental
plate consist of unequal sized lamellae closely pressed
together. They are similar to dental plates of Diodon
known from the Early Eocene Kapurdi Formation of Barmer,
Rajasthan, Cambay Shale of Gujarat and Subathu Forma-
tion of Himachal Pradesh 20,28,34 .
The Khuiala fish assemblage has several taxa in com-
mon with those recorded from the Lower Eocene of Akli,
Kapurdi and Cambay Shale formations of the Barmer and
Cambay basins of Rajasthan and Gujarat respectively, as
well as from the Kakara and Subathu formations of the
northwest sub-Himalaya, Himachal Pradesh. Like the
Akli and Cambay fish assemblages, the presently identi-
fied Khuiala assemblage is generally devoid of oceanic
fish but it also lacks enchodontids and freshwater ele-
ments like lepisosteids and osteoglossids, which are quite
common, especially osteoglossids in both the Akli (from
Giral Lignite Mine, Barmer District) and Cambay Shale
(from Vastan Lignite Mine, Surat District) formations.
This may be related to the fact that during the Lower Eo-
cene the strandline was closer to Vastan and Akli than to
westerly Khuiala, Jaisalmer, and therefore, there may
have been little or no influence of coastal riverine fauna
around Khuiala. Possibility of the absence of osteoglos-
sids and lepisosteids being an artefact because the Khui-
ala assemblage as known today is based on our first
collection from screen-washing of a small sample is mini-
mal, because teeth and scales of both these taxa are gen-
erally among the first to be picked from screen-washed
matrix, but it cannot be ruled out completely because
there is definitely a scope for developing a larger assem-
blage from Khuiala. Among the Paleocene–Eocene fish
faunas known from India, this Khuiala assemblage ap-
pears closest to the Vastan fish assemblage known from
the subsurface Cambay Shale, Surat District, Gujarat
since out of its 14 taxa 8 are common with Vastan. Notable
among Vastan fishes that are not present in this Khuiala
558
ichthyofauna are Myliobatis, Enchodus and osteoglossids.
This Khuiala assemblage differs from the Akli fish as-
semblage in lacking Apateodus, Myliobatis, Jaekelotodus,
Squatina and Arius and in possessing Galeorhinus, Rhi-
nobatos, Coupatezia, Sphyraena, Eutrichiurides, Sparus,
Diodon and Stephanodus. It also differs from the Kapurdi
assemblage mainly in lacking Galeocerdo, Notorhynchus,
Myliobatis and Enchodus. An exclusively Paleocene fish
assemblage known from the Fatehgarh Formation, Bar-
mer District, Rajasthan differs from Khuiala ichthyofauna
mainly in having Apateodus, a typical Cretaceous–Paleo-
cene marker, besides Myliobatis, Lepisosteus, Enchodus
and osteoglossids21 . The Paleocene–Eocene ichthyofaunas
becoming known from several localities of western India
are broadly similar and part of the same bioprovince that
extended into the sub-Himalaya.
The age of the Khuiala fish assemblage is constrained
by the presence of index larger foraminifer Nummulites
burdigalensis in the Lower Khinsar Shale and Sirhera
members that overlie the vertebrate-yielding Te Takkar
Member 3,4 . N. burdigalensis corresponds to the SBZ
(Shallow Benthic Zone) 10 and Planktic Zone P8 and in-
dicates a Ypresian age45 . The ichthyofauna documented
here also supports a Ypresian age. The ostracode, bivalve
and other biotic assemblages previously reported from the
Khuiala Formation are also consistent with this age as-
signment1–4,10–15 . The modern relatives of the Khuiala fish
are mainly confined to tropical, subtropical, essentially
shallow near-shore habitats, suggesting a rather protected
shallow marine environment (inner–middle shelf zone),
not widely exposed to the oceanic realm, and probably
without any influx of coastal freshwater. This is also sub-
stantiated by the occurrence of serpulid worm tubes and
other invertebrate fauna known from the Khuiala Forma-
tion 1,3,4,10–15 .
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ACKNOWLEDGEMENTS. We thank the Director, Wadia Institute
of Himalayan Geology, Dehradun for providing the Scanning Electron
Microscope and other necessary facilities. R. Smith and J. Herman
(Belgium) helped in identifying some of the material. This work was
partly funded by a grant to R.S.R. by the Department of Science and
Technology, New Delhi.
Received 1 December 2006; accepted 27 April 2007
559