OSTEOLOGY OF BARAPASAURUS TAGOREI
(DINOSAURIA: SAUROPODA) FROM THE EARLY
JURASSIC OF INDIA
by SASWATI BANDYOPADHYAY* , DAVID D. GILLETTE� ,
SANGHAMITRA RAY� and DHURJATI P. SENGUPTA**Geological Studies Unit, Indian Statistical Institute, 203 Barrackpore Trunk Road, Kolkata 700108, India; e-mails [email protected] and [email protected]
�Museum of Northern Arizona, 3101 N. Fort Valley Road, Flagstaff, Arizona 86001, USA; e-mail [email protected],
�Department of Geology & Geophysics, Indian Institute of Technology, Kharagpur 721302, India; e-mail [email protected]
Typescript received 15 October 2007; accepted in revised form 25 February 2009
Abstract: The sauropod dinosaur, Barapasaurus tagorei, is
known from the Early Jurassic Kota Formation (Sinemurian
to Pliensbachian) of India. The taxon is represented by c. 300
bones that were found associated with large fossilized tree
trunks and were collected from the interface of sandstone
and mudstone units covering an area of c. 276 m2. The col-
lection includes one partial skeleton; most of the remainder
of the bones were disarticulated, disassociated and dispersed,
but taphonomic analysis permits recognition of associated
elements comprising several individuals. Skeletal anatomy of
Barapasaurus includes several teeth, vertebrae from the caudal
cervicals rearward to the terminal caudals, and most elements
of the appendicular skeleton. Barapasaurus is characterized by
spoon-shaped teeth with bulbous bases and grooves on the
anterolabial and posterolingual sides of the crown, coarse
tubercles on the carina, acamerate cranial and dorsal verte-
brae, lateral laminae of the middle and caudal dorsal neural
spines composed of spinodiapophyseal and spinopostzygapo-
physeal laminae, neural canal of the mid-dorsal vertebrae
opens dorsally through a narrow slit into a large cavity and
sacrum with four co-ossified vertebrae. Phylogenetic analysis
reveals that Barapasaurus is basal in comparison with Vulcan-
odon and is removed from Eusauropoda.
Key words: Dinosaur, Sauropoda, Jurassic, Pranhita-Godav-
ari Basin, India.
Sauropod dinosaurs appeared during the Late Triassic
and diversified successfully in both the number of genera
and species throughout the Mesozoic but became extinct
by the end of Late Cretaceous. Fossil remains of these
gigantic, terrestrial, herbivorous dinosaurs with small
skull, simple teeth, long neck, long tail, quadrupedal gait
and upright limbs are found in almost all the continents
(McIntosh 1990; Upchurch et al. 2004). Nevertheless,
records of early sauropods of Late Triassic or Early
Jurassic age are still quite rare in comparison with their
Cretaceous successors and are known from a very few
places such as India, Thailand, China and Africa.
In India, early sauropod dinosaurs were first recorded
in 1962 with the discovery of about 300 bones occurring
just above the interface of sandstone and mudstone units
of the Kota Formation of peninsular India (Jain et al.
1962). The Early Jurassic Kota Formation occurs in the
Pranhita-Godavari basin, a Gondwana basin in Deccan,
India (Text-fig. 1). This is the only continental Early
Jurassic horizon in India and also one of the few in the
world producing rich terrestrial fauna including fishes,
reptiles, mammals, freshwater ostracodes, conchostrachans
and land insects (Jain 1980). After the initial discovery,
excavations in the following years produced a large num-
ber of bones including at least six skeletons in partial
association. Study of these collections led to the establish-
ment of a new sauropod dinosaur, Barapasaurus tagorei
(Jain et al. 1975, 1979), one of the oldest sauropod dino-
saurs of the world. The material included about six
individuals ranging in ontogenetic development from
juvenile to adult. Since the age of the Kota Formation is
generally considered as Early to early Middle Jurassic
(Bandyopadhyay and RoyChowdhury 1996; Bandyopad-
hyay and Sengupta 2006), this material constitutes an
early population of this sauropod in India. Jain et al.
(1975, 1979) presented preliminary osteological descrip-
tions of some of the diagnostic elements in the hypodigm,
concluding that certain aspects of the anatomy are primi-
tive for a sauropod, while other aspects resembled those
of the prosauropods.
According to the original diagnosis by Jain et al.
(1975), Barapasaurus tagorei is a large sauropod with
[Palaeontology, Vol. 53, Part 3, 2010, pp. 533–569]
ª The Palaeontological Association doi: 10.1111/j.1475-4983.2010.00933.x 533
slender limbs, spoon-shaped teeth with coarse denticles
on posterior and anterior keels, opisthocoelous cranial
dorsal vertebrae, sacrum consisting of four co-ossified
vertebrae and narrow width between the sacricostal yokes,
deep medial wall of the ilium, and shallow curvature of
the proximal part of the anterior border of the scapula. It
bears similarity with the prosauropod dinosaurs based on
slender limbs, distinctive fourth trochanter of the femur
and small pelvic basin (Jain et al. 1979).
Earlier description and diagnosis of Barapasaurus was
by Jain et al. (1975, 1979) during the time when sauro-
pods were much less well known. Subsequently, informa-
tion on several new basal sauropods came to light from
different parts of the world including two more basal
sauropods from the Pranhita-Godavari basin of India. Of
them, Kotasaurus yamanpalliensis Yadagiri, 1988 was also
recovered from the Early Jurassic Kota Formation of the
Pranhita-Godavari basin (Table 1) from a locality c.
40 km north of the type locality of B. tagorei (Text-
fig. 1). The other basal sauropod Lamplughsaura dharma-
ramensis Kutty, Chatterjee, Galton and Upchurch, 2004
has been recovered from the upper part of the early Early
Jurassic (Hettangian) Dharmaram Formation of the
Pranhita-Godavari basin (Table 1) and has recently been
described on the basis of a partially associated skeleton of
a nearly adult individual (Kutty et al. 2007).
The other known basal sauropods include Vulcanodon
karibaensis Raath, 1972 from the Early Jurassic of Zimba-
bwe, Isanosaurus attavipachi Buffetaut, Suteethorn, Cuny,
Tong, Le Loeuff, Khansubha, and Jongautchariyakul, 2000
from the northeastern Thailand, Antetonitrus ingenipes
Yates and Kitching, 2003 and Blikanasaurus cromptoni
Galton and Van Heerden, 1985 from the Late Triassic of
South Africa, Tazoudasaurus naimi Allain, Aquesbi, Dejax,
Meyer, Monbaron, Montenat, Richir, Rochid, Russell and
Taquet, 2004 from the Early Jurassic of the Moroccan
High Atlas, and ‘melanorosaurids’ from Africa and China
(Table 1). Apart from these Gondwanan basal sauropods,
TEXT -F IG . 1 . Type localities of Kota
sauropods Barapasaurus tagorei and
Kotasaurus yamanpalliensis, and the
geological map of the surrounding area
(modified after Rudra 1982;
Bandyopadhyay and Rudra 1985; Ghosh
1994; T. RoyChowdhury, pers. comm.
2003). Inset: Location of Pranhita-
Godavari basin in India.
534 P A L A E O N T O L O G Y , V O L U M E 5 3
TA
BL
E1
.A
syn
op
sis
of
the
kno
wn
bas
alsa
uro
po
ds.
Tax
on
Stra
tigr
aph
ico
ccu
rren
ces
Age
Mat
eria
l,ch
arac
teri
stic
feat
ure
sR
efer
ence
s
Kot
asau
rus
yam
anpa
llie
nsi
s
Lo
wer
par
to
fth
eK
ota
Fo
rmat
ion
,P
-Gb
asin
,In
dia
Ear
lyJu
rass
ic
(Sin
emu
rian
toP
lien
sbac
hia
n)
Ab
ou
t12
ind
ivid
ual
s;ex
cep
tte
eth
no
sku
llm
ater
ial;
sim
ple
do
rsal
vert
ebra
ew
ith
ou
tel
abo
rate
dsp
inal
lam
inae
;ab
sen
ceo
fp
neu
mat
oco
elo
n
the
bas
eo
fn
eura
lar
cho
pen
ing
into
neu
ral
can
al;
lon
gb
ut
low
ilia
cb
lad
e
wit
hst
raig
ht
do
rsal
bo
rder
;sc
apu
law
ith
nar
row
pro
xim
alsu
rfac
e;li
mb
bo
nes
rela
tive
lysl
end
er;
fem
ur
wit
hle
sser
tro
chan
ter,
v-sh
aped
chev
ron
s
wit
hw
ell-
dev
elo
ped
arti
cula
rfa
cets
on
the
do
rso
late
ral
corn
ers.
Yad
agir
i
(198
8,20
01)
Lam
plu
ghsa
ura
dh
arm
aram
ensi
s
Up
per
par
to
fth
e
Dh
arm
aram
Fo
rmat
ion
,
P-G
bas
in,
Ind
ia
earl
yE
arly
Jura
ssic
(Het
tan
gian
)
Ah
eavi
lyb
uil
tq
uad
rup
edal
taxo
nw
ith
ab
od
yle
ngt
ho
f10
m;
teet
hw
ith
stro
ngl
yem
argi
nat
edd
ista
led
ge;
cau
dal
cerv
ical
neu
ral
spin
esb
eari
ng
a
tran
sver
sely
exp
and
edsp
ine
tab
lean
da
vert
ical
lyo
rien
ted
liga
men
tou
s
furr
ow
on
cran
ial
and
cau
dal
surf
aces
;cr
anio
do
rsal
lyd
irec
ted
spu
ro
nth
e
pro
xim
alca
ud
aln
eura
lsp
ines
wh
ile
ala
rge
pro
cess
on
mid
cau
dal
neu
ral
spin
e;ca
ud
aln
eura
lsp
ines
sho
rter
than
tran
sver
sep
roce
ssan
d
con
seq
uen
tly
lost
inta
il;
man
ual
un
gual
In
on
tren
chan
t.
Ku
tty
etal
.(2
007)
Vu
lcan
odon
kari
bien
sis
Vu
lcan
odon
Bed
s
(Mas
ho
bal
and
No
rth
),
Zim
bab
we
Ear
lyJu
rass
icSa
cru
mw
ith
?4fu
sed
vert
ebra
e;is
chiu
mlo
nge
rth
anp
ub
is,
pu
bes
form
ing
ap
rom
inen
tan
teri
or
faci
ng
pu
bic
apro
n,
iliu
mw
ith
lon
gp
ub
icp
edu
ncl
e,
amp
hic
oel
ou
sca
ud
als,
stra
igh
tco
lum
nar
fem
ur,
fore
lim
b78
%o
fth
e
len
gth
of
hin
dli
mb
,p
roxi
mal
end
of
met
atar
sal
IItr
ansv
erse
lyn
arro
wer
than
the
oth
erm
etat
arsa
ls,
hal
lux
un
gual
enla
rged
,d
ista
ln
on
un
gual
ped
al
ph
alan
ges
of
dig
itII
Ian
dIV
stu
bb
yan
dw
ider
,p
roxi
mal
arti
cula
r
surf
aces
of
ped
ald
igit
su
ngu
als
IIan
dII
Id
ors
ove
ntr
ally
com
pre
ssed
resu
ltin
gin
wid
erb
read
thth
anh
eigh
t.
Raa
th(1
972)
;
Co
op
er(1
984)
Isan
osau
rus
atta
vipa
chN
amP
ho
ng
Fo
rmat
ion
of
no
rth
east
ern
Th
aila
nd
Lat
eT
rias
sic
(No
rian
-Rh
aeti
an)
Ass
oci
ated
skel
eto
nw
ith
ap
rom
inen
tp
roce
ss(?
cran
ial
pro
cess
)p
roje
ctin
g
late
rall
yfr
om
the
pro
xim
alen
do
fth
efe
mu
r,m
akin
gth
ela
tera
lm
argi
n
of
the
fem
ora
lsh
aft
stro
ngl
yco
nca
vein
cran
ial
view
and
ave
ry
pro
min
ent,
acu
min
ate,
S-sh
aped
fou
rth
tro
chan
ter
inth
ep
roxi
mal
hal
fo
f
the
rob
ust
stra
igh
tfe
mu
r.
Bu
ffet
aut
etal
.(2
000)
An
teto
nit
rus
inge
nip
esL
ow
erE
llio
tF
orm
atio
n
of
Sou
thA
fric
a
Lat
eT
rias
sic
(No
rian
)P
arti
ald
isar
ticu
late
dsk
elet
on
;d
ors
aln
eura
lsp
ines
flar
edtr
ansv
erse
lyat
thei
rd
ista
len
d;
do
rsal
vert
ebra
ew
ith
bro
ad,
tria
ngu
lar,
hyp
osp
hen
ein
cau
dal
view
;ve
ntr
alri
dge
on
the
hyp
osp
hen
eo
fth
eca
ud
ald
ors
al
vert
ebra
e;a
dee
psu
lcu
sad
jace
nt
toth
ela
tera
ld
ista
lm
argi
no
fth
e
del
top
ecto
ral
cres
t;an
dan
extr
emel
ysh
ort
,b
road
met
acar
pal
I.
Yat
esan
d
Kit
chin
g(2
003)
Ch
insh
akia
ngo
sau
rus
chu
ngh
oen
sis
Lo
wer
par
to
fth
e
Fen
ghah
eF
orm
atio
no
f
Zh
on
ghe,
Yu
nn
anP
rovi
nce
of
Rep
ub
lic
of
Ch
ina
Ear
lyJu
rass
icO
ne
or
mo
rep
arti
alsk
elet
on
incl
ud
ing
a12
–13
-m-l
on
gin
div
idu
al;
the
ho
loty
pe
mat
eria
lis
ap
arti
alsk
elet
on
con
sist
ing
of
an
earl
yco
mp
lete
left
den
tary
,o
ne
cerv
ical
vert
ebra
,se
vera
ld
ors
alve
rteb
rae,
cran
ial
cau
dal
vert
ebra
e,b
oth
scap
ula
e,in
com
ple
tep
elvi
cgi
rdle
and
the
hin
dli
mb
s.
Do
ng
(199
2);
Up
chu
rch
etal
.(2
007b
)
Jin
gsh
anos
auru
sxi
nw
aen
sis
Lo
wer
Lu
fen
gSe
ries
(Yu
nn
an)
of
Peo
ple
’s
Rep
ub
lic
of
Ch
ina
earl
yE
arly
Jura
ssic
(Het
tan
gian
to
Pli
ensb
ach
ian
)
An
earl
yco
mp
lete
sku
llan
dp
ost
cran
ial
skel
eto
no
fan
adu
ltin
div
idu
al.
Zh
ang
and
Yan
g(1
994)
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 535
TA
BL
E1
.(C
onti
nu
ed).
Tax
on
Stra
tigr
aph
ico
ccu
rren
ces
Age
Mat
eria
l,ch
arac
teri
stic
feat
ure
sR
efer
ence
s
Gon
gxia
nos
auru
ssh
ibei
nsi
sZ
iliu
jin
gF
orm
atio
no
f
Sich
uan
Pro
vin
ceo
fC
hin
a.
Ear
lyJu
rass
icN
earl
yco
mp
lete
skel
eto
nex
cep
tth
esk
ull
;w
ell-
dev
elo
ped
po
ster
ior
pro
cess
inth
ep
rem
axil
la;
larg
e,sp
oo
n-s
hap
edte
eth
wit
ho
ut
any
serr
atio
nb
ut
wit
hth
ick
lon
gitu
din
alst
riat
ion
so
nli
ngu
alan
dla
bia
lsi
des
;
amp
hip
laty
anto
amp
hic
oel
ou
sve
rteb
rae;
thre
efu
sed
sacr
als;
den
se
inte
rnal
stru
ctu
res
incr
ania
lsa
cral
;ca
ud
alch
evro
ns
no
tb
ifu
rcat
ed.
He
etal
.(1
998)
Yu
nn
anos
auru
sro
bust
us
Lo
wer
Lu
fen
gF
orm
atio
n,
Yu
nan
Pro
vin
ceo
fC
hin
a
Ear
lyJu
rass
icA
do
rso
ven
tral
lysh
allo
wm
axil
law
ith
ten
pre
serv
edte
eth
inva
rio
us
stag
es
of
eru
pti
on
in16
alve
oli
wh
ich
grad
ual
lyd
ecre
ase
insi
zep
ost
erio
rly;
spat
ula
tete
eth
wit
hw
rin
kled
text
ure
inth
een
amel
;to
oth
cro
wn
s
lin
gual
lyco
nca
vean
dla
bia
lly
con
vex.
Yo
un
g(1
951)
;
Sim
mo
ns
(196
5);
Bar
rett
(199
9)
Taz
oud
asau
rus
nai
mi
Hig
hA
tlas
of
Mo
rocc
oE
arly
Jura
ssic
Ap
artl
yar
ticu
late
dsk
elet
on
;th
inb
on
yp
late
pre
sen
to
nth
ep
ost
ero
do
rsal
mar
gin
of
the
po
sto
rbit
al;
20d
enta
ryte
eth
wit
hd
enti
cula
tecr
ow
n
mar
gin
s;an
teri
or
end
of
den
tary
slig
htl
yex
pan
ded
inco
mp
aris
on
toth
e
dep
tho
fd
enta
ryat
mid
len
gth
;q
uad
rate
wit
ho
ut
any
po
ster
ior
foss
a;
fork
edch
evro
n;
pro
min
ent
cres
to
nth
ela
tera
lsu
rfac
eo
fth
ep
roxi
mal
end
of
the
fib
ula
;le
sser
tro
chan
ter
occ
urr
ing
late
rall
yo
nth
efe
mu
r;fl
at
pu
bic
apro
nan
dfl
atte
ned
pla
nte
rsu
rfac
eo
fp
edal
un
gual
sII
.
All
ain
etal
.(2
004)
;
All
ain
and
Aq
ues
bi
(200
8)
Bli
kan
asau
rus
crom
pton
iL
ow
erE
llio
tF
orm
atio
n
of
Sou
thA
fric
a
Lat
eT
rias
sic
(Ear
lyN
ori
an)
Par
tial
sto
cky
hin
dli
mb
con
sist
ing
of
asso
ciat
edle
ftti
bia
,fi
bu
la,
tars
us
and
pes
;th
ed
ista
len
do
fth
efi
bu
lafa
ces
stro
ngl
yve
ntr
om
edia
lly.
Ch
arig
etal
.(1
965)
;
Gal
ton
and
Van
Hee
rden
(198
5):
Gal
ton
etal
.(1
998)
‘mel
ano
rosa
uri
ds’
Lo
wer
Ell
iot
Fo
rmat
ion
of
Sou
thA
fric
a
Lat
eT
rias
sic
(Ear
lyN
ori
an)
Gal
ton
and
Up
chu
rch
(200
4)
Mel
anor
osau
rus
read
iL
ow
erE
llio
tF
orm
atio
n
of
Sou
thA
fric
a
Lat
eT
rias
sic
(Ear
lyN
ori
an)
An
earl
yco
mp
lete
skel
eto
nw
ith
ale
ngt
ho
f7Æ
5m
bu
tw
ith
ou
tan
ysk
ull
and
two
mo
rep
arti
alsk
elet
on
s
Hau
ghto
n(1
924)
Ver
teb
rae,
two
com
ple
tesa
cra,
scap
ula
e,a
hu
mer
us,
anu
lna,
mo
sto
fth
e
pel
vis
and
hin
dli
mb
Gal
ton
etal
.(2
005)
Sku
llan
dat
las-
axis
com
ple
xY
ates
(200
7a)
Mel
anor
osau
rus
thab
aen
siU
pp
erE
llio
tF
orm
atio
n
of
Les
oth
o
Ear
lyJu
rass
icF
emu
rG
auff
re(1
993)
‘Mel
ano
rosa
uri
ds’
Ch
ina
Ear
lyJu
rass
icG
alto
nan
dU
pch
urc
h
(200
4)
536 P A L A E O N T O L O G Y , V O L U M E 5 3
descriptions of four sauropod taxa from the People’s
Republic of China establish a Laurasian distribution of
basal sauropods in the Early Jurassic. These are Chinshak-
iangosaurus chunghoensis Ye vide Dong, 1992, Jingshano-
saurus xinwaensis Zhang and Yang, 1994, Gongxianosaurus
shibeinsis He, Wang, Liu, Zhou, Liu, Cai and Dai, 1998
from Sichuan and Yunnanosaurus robustus Young, 1951
from Yunnan Province (Table 1).
The above-mentioned early sauropod genera were
considered to belong to Eusauropoda (Upchurch 1995,
1998; Upchurch et al. 2004; Salgado et al. 1997; Wilson
2002; Wilson and Sereno 1998). Eusauropoda or the basal
Sauropoda are characterized by tall, broad dorsal neural
spines produced by bony laminae, columnar limbs with
reduced processes for muscle attachment, an elongated
forelimb, a deep radial fossa on proximal ulna, radius with
flat caudally facing ulnar facet at its distal end, semitubular
metacarpal, an elliptical femoral cross-section and a short-
ened, spreading pes (Upchurch 1998; Upchurch et al. 2004;
Wilson 2002; Wilson and Sereno 1998; Bonnan 2003).
However, Upchurch et al. (2007a) concluded that Bara-
pasaurus, Kotasaurus and Lamplughsaura from India were
basal Sauropoda and removed from the Eusauropoda,
although Allain and Aquesbi (2008) considered Barapa-
saurus as a Eusauropod. It is evident that the relation-
ships among all basal sauropods cannot be fully analysed
without additional elucidation of osteology in all these
genera. With the recent expansion of knowledge of early
sauropods (e.g. Vulcanodon, Kotasaurus, Chinshakiango-
saurus, Antetonitrus, Blikanasaurus, Tazaoudasaurus and
Jingshanosaurus), a more extensive description of Barapa-
saurus became imperative. Hence, this article reviews the
osteology of B. tagorei, with emphasis on specimens that
have not been previously described.
GEOLOGICAL SETTING
The Gondwana succession of the Pranhita-Godavari basin
(Text-fig. 1) occurs as a narrow, rectilinear outcrop
trending NNW–SSE and is bordered on both sides by
Proterozoic and ⁄ or Archaean rocks; the succession is
overlain by Deccan Trap basalt of Cretaceous to Palaeo-
cene age (69–63 Ma) (Pande 2002). The overall dip of the
succession is 5–12 degrees north and north-west, while
the general palaeocurrent direction is north (Sengupta
1970). The Gondwana sediments were deposited in gla-
cial, fluvioglacial, fluviatile and lacustrine conditions and
range in age from Permian to Cretaceous. The Kota For-
mation in the Upper Gondwana of the Pranhita-Godavari
basin overlies the Dharmaram Formation and shows a
more or less uniform lithology throughout the valley
(Table 2). Rudra (1982) divided the Kota Formation into
two lithological units. The lower unit includes 15–25 m
of thick, hard, compact and coarse sandstone, which is
pebbly in places. This sandstone becomes finer and grades
both laterally and vertically into finer siltstone and mud-
stones. The upper part of the Kota Formation includes
marl and 1–2 m thick limestone beds. This is followed
upward by mudstone and ferruginous shale interbedded
with sandstone. Rudra and Maulik (1994) suggested that
a meandering river system deposited the Lower Kota,
while a braided river system formed the upper part; the
limestone facies was interpreted to be a lacustrine deposit.
The Kota Formation has quite a rich vertebrate fauna
whose remains come from two successive stratigraphic
levels (Table 3). The mudstone of the dominantly fluvia-
tile lower unit has yielded two sauropod dinosaurs and
three mammals. Barapasaurus and Kotasaurus are the two
basal sauropod dinosaurs. The osteology of Barapasaurus
is treated in this article. Kotasaurus is characterized by
simple dorsal vertebrae (without spinal laminae), low iliac
blade, narrow proximal surface to the scapula, relatively
slender limb bones, femur with lesser trochanter and ‘v’-
shaped chevrons with well-developed articular facets on
the dorsolateral corners (Yadagiri 1988, 2001). Some the-
ropod teeth have also been recovered from this horizon.
On the basis of isolated teeth, three mammals have been
identified from this horizon – a kuehneotherid (Kota-
therium) (Datta 1981) and a probable amphilestid (Indo-
therium) (Yadagiri 1984). Datta and Das (2001) recently
described a molariform tooth Indozostrodon belonging to
Megazostrodontidae.
The fauna of the upper unit of the Kota Formation, a
limestone-dominated lacustrine deposit, includes three
semionontids (Lepidotes, Paradapedium and Tetragono-
lepis) (Jain 1959, 1973, 1983), a pholidophorid (Pholid-
ophorous) (Yadagiri and Prasad 1977) and a coelacanth
(Indocoelacanthus) (Jain 1974a). Of the ‘holostean’ fishes,
Lepidotes is the most common semionotid. Paradapedium,
though very close to Dapedium, morphologically differs in
skull and body proportions. Among the reptiles, there are
a pterosaur (Campylognathoides) (Jain 1974b), a mesosu-
chian crocodylomorph (Bandyopadhyay and Roychowdh-
ury 1996) and a cryptodiran turtle (Indochelys) (Datta
et al. 2000). The turtle, Indochelys, shows similarity with
the North American Kayentachelys. A scute and other
fragmentary limb bones indicate the presence of another
diapsid – a scelidosaurid dinosaur in this horizon (Ban-
dyopadhyay and Roychowdhury 1996).
Yadagiri (1986) reported the presence of a micro-verte-
brate assemblage comprising an elasmobranch (Lissodus),
a rhynchocephalian and fragments of a pleurodont denti-
tion, identified as Paikasisaurus indicus of uncertain affin-
ity. Later, Prasad and Arratia (2004) described two
elasmobranchs ?Polyacrodus and Lissodus along with some
other ‘holosteans’. Two sphenodontians, Rebbanasaurus
and Godavarisaurus, and three dentary fragments and a
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 537
partial maxilla of a probable pleurodont lepido-
sauromorph similar to basal rhynchocephalians have been
described by Evans et al. (2001). Subsequently, Evans
et al. (2002) described an acrodont lizard, Bharatagama
and two indeterminate agamid lizards. Among the micro-
mammals, two docodonts, Gondtherium and Godavari-
odon (Prasad 2003; Prasad and Manhas 2007) and a small
lower molar of uncertain affinities, Dyskritodon (Prasad
and Manhas 2002), two dryolestids and a probable ‘am-
philestid’ Paikasigudodon (Prasad and Manhas 1997,
2002) have been described from this horizon. Besides,
there are a therian (Trishulotherium) (Yadagiri 1984), and
a holotherian (Nakunodon) (Yadagiri 1985) of uncertain
familial affinities (Averianov 2002).
On the basis of fishes, the Kota Formation has long
been considered to be of Liassic age (King 1881; Robin-
son 1970). However, several workers noticed the similar-
ity of Kota fishes with the European Toarcian fishes
(Schaeffer and Patterson 1984). Patterson and Owen
(1991) suggested that the major marine transgression
during the Toarcian might have been instrumental for the
invasion of the European Liassic fishes in the circum-
tethyean continents including parts of Indian subconti-
nent. Recovery of Darwinula, an early Middle Jurassic
ostracod, led Govindan (1975) to suggest a Middle Juras-
sic age for the Kota Formation. Analysing the faunas of
the underlying Dharmaram Formation and of the lower
and upper units of the Kota Formation and comparing
them with faunas from coeval horizons, Bandyopadhyay
and Roychowdhury (1996) and Bandyopadhyay and Seng-
upta (2006) suggested that the Lower Kota Formation has
an age ranging from Sinemurian to Pliensbachian, while
the age of the upper Kota is Toarcian and may even be
extended to Middle Jurassic (?Aalenian).
TAPHONOMY OF THE BARAPASAURUSASSEMBLAGE
A large number of dinosaur bones were discovered as sur-
face finds from the Kota Formation of the P-G valley in
1958–59 (Jain et al. 1962). Proper excavation in 1960–61
at a site about 18 km south-east of Sironcha (18�51¢N,
79�58¢E) (Text-fig. 1), near a small village of Pochampalli
TABLE 2 . Gondwana stratigraphy of the Pranhita-Godavari basin.
Formations Main Lithologies Important Fossils Age
UPPER GONDWANA
Chikiala Highly ferruginous sandstone
and conglomerate
? ?
Gangapur Coarse, gritty sandstone, greyish
white to pinkish mudstone with
interbedded ferruginous sandstone
and concretions
Gleichenia, Pagiophyllum,
Ptilophyllum, Elatocladus
Early Cretaceous
-------------------------------------------------------------------- Unconformity -------------------------------------------------------------------
Kota Sandstone, siltstone and mudstone
with limestone bands
‘holosteans’, sauropods,
sphenodontians, lepidosaurids,
crocodylomorphs, cryptodire,
‘symmetrodonts’
Early to Middle Jurassic
Dharmaram Coarse sandstone and mudstone Sauropods, theropods,
sphenosuchians, phytosaurids,
aetosaurids
late Late Triassic to
early Early Jurassic
Maleri Mudstone, fine to medium
sandstone and calcirudite
Metoposaurids, chigutisaurids,
phytosaurids, rhynchosaur,
aetosaurids, theropods, cynodont,
prolacertid, therapsid, basal
saurischian, eosuchian
early Late Triassic
Bhimaram Coarse to fine sandstone,
ferruginous or calcareous at
places and mudstone
– late Middle Triassic
Yerrapalli Red and violet mudstone
and calcirudite
Capitosaurids, therapsids, cynodont,
rhynchosaur, prolacertid, rauisuchid,
erythrosuchid
early Middle Triassic
Kamthi Siltstone, ferruginous
sandstone – pebbly at places
Brachyopid, therapsid Early Triassic
After Bandyopadhyay and Sengupta (2006) and Kutty et al. (2007).
538 P A L A E O N T O L O G Y , V O L U M E 5 3
(18�44¢N, 80�05¢E) (Gadchiroli district, Maharashtra) led
to the recovery of a rich layer of sauropod bones, later
described as Barapasaurus tagorei (Jain et al. 1975, 1979).
These bones were found with fossilized large tree trunks
just above a sandstone–mudstone interface (Jain et al.
1962). The rich dinosaur bone layer occurred in an area
of about 276 m2. Only postcranial material and a few
teeth of adult individuals totalling c. 300 bones were
found from the site. On the basis of six left femora, it
is estimated that there were at least six individuals
(Text-fig. 2). Apart from one partially associated skeleton,
the bones were disarticulated, disassociated and dispersed.
However, the state of preservation of the majority of the
fossil bones is remarkably good. Although orientation of
the bones is polymodal (Bandyopadhyay et al. 2002,
fig. 14), two strong modes were identified. The majority
of the bones were oriented nearly NNW–SSE while
another set of bones along with logs were oriented NE–
SW. The disarticulated bones are all complete and well
preserved. The more or less good shape of the vertebrae
and the girdle bones indicate that the bones did not suffer
much distortion. Some of the bones show long, fine des-
iccation cracks indicating brief surface exposure before
burial, but do not show any other surface marks.
Bandyopadhyay et al. (2002) interpreted the bone
assemblage of B. tagorei as an accumulation from mass
mortality because of a catastrophic event. A flood might
have been the cause of death of this herd of Barapasaurus,
whose carcasses were transported by floodwater for a dis-
tance. The associated tree logs were uprooted in the flood
event and were transported together with Barapasaurus.
Subsequently, these carcasses became entangled with the
tree trunks, decomposed and disarticulated. The sauropod
skulls being fragile, light and with weak necks were frag-
mented and washed away, while the heavier postcranial
bones were left behind along with the tree trunks. This
bone assemblage was exposed on the surface for some
time and subsequently buried by silt and clay material.
MATERIAL
A major part of the material is in the mounted skeleton
of B. tagorei, displayed at the Geology Museum of ISI,
and the rest of the material is in the Palaeontological col-
lection of the Geological Studies Unit, ISI. Most of the
material (Material A) had been collected from the area
near the village Pochampalli, about 18 km southeast of
Sironcha (18�51¢N, 79�58¢E), Gadchiroli district, Maha-
rashtra, India.
Material A. ISIR 50, sacrum (four co-ossified sacral vertebrae
with sacricostal yoke); ISIR 51, right ilium (mounted); ISIR 52,
right ilium; ISIR 53, right ischium (mounted); ISIR 54, left
ischium; ISIR 55, right pubis (mounted); ISIR 56, left pubis
(mounted); ISIR 57, right pubis; ISIR 58, right femur in two
TABLE 3 . The vertebrate fauna of the Kota Formation.
UPPER FAUNA
Fishes Semionotid Lepidotes deccanensis Toarcian to ?Aalenian
Paradapedon egertoni
Tetragonolepis oldhami
Pholidophorid Pholidophorus kingii, P. indicus
Coelacanthid Indocoelacanthus robustus
Reptiles Kyantachelyid Indochelys spatulata
Campylognathoidid Campylognathoides indicus
Sphenodontian Rebbanasaurus jaini
Godavarisaurus lateefi
Lepidosaur Bharatagama rebbanensis
Mammals Docodontid Gondtherium dattai,
Godavariodon denisei
Morganucodontid Indotherium pranhitai
‘Amphilestid’ ?Paikasigudodon yadagiri
Incertae sedis Dyskritodon indicus
Incertae sedis Nakunodon paikasiensis
Incertae sedis Trishulotherium kotaensis
LOWER FAUNA
Reptiles Sauropods Barapasaurus tagorei,
Kotasaurus yamanpalliensis
Sinemurian to
Pliensbachian
Mammals Kuehneotheriid Kotatherium haldanei
Amphilestid Indotherium pranhitai
Morganucodontid Indozostrodon simpsoni
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 539
pieces (mounted); ISIR 59, left femur (mounted); ISIR 60, left
femur in two pieces; ISIR 61, left tibia; ISIR 62, left tibia
(mounted); ISIR 63, only distal end of left tibia; ISIR 64, left fib-
ula; ISIR 68, left scapula (mounted); ISIR 69, left coracoid
(mounted); ISIR 70, left humerus; ISIR 71, left radius; ISIR 72,
left ulna; ISIR 74, twelfth dorsal vertebra (mounted); ISIR 79,
first dorsal vertebra (mounted); ISIR 80 third cranial dorsal ver-
tebra (mounted); ISIR 81, second dorsal vertebra (mounted);
ISIR 83, ungual phalanx (digit I) of right pes (mounted); ISIR
84, ungual phalanx of digit III of right pes, (mounted); ISIR 85,
right humerus; ISIR 86, proximal half of left humerus
(mounted); ISIR 87, left humerus; ISIR 88, right humerus
(mounted); ISIR 89, right radius; ISIR 90, right ulna; ISIR 91,
left ulna (mounted); ISIR 92, right scapulo-coracoid; ISIR 94,
metacarpal III; ISIR 95, metacarpal IV; ISIR 96, metacarpal V;
ISIR 97–98, left femora; ISIR 99–100, right femora; ISIR 101,
right tibia; ISIR 102, left tibia; ISIR 105, right fibula; ISIR 106,
left fibula; ISIR 108, left metacarpal of digit I; ISIR 110, ungual
phalanx of left digit I (juvenile); ISIR 111, large left ilium, ischi-
adic peduncle & iliac plate broken (mounted); ISIR 112, right
large ilium; ischiadic peduncle and iliac plate broken; ISIR 114,
left ischium (mounted); ISIR 115, right ischium; ISIR 116, left
ischium; ISIR 117, left pubis; ISIR 118, left pubis; ISIR 121,
tenth cervical vertebra (mounted); ISIR 122, fourth dorsal verte-
bra (mounted); ISIR 123, mid-dorsal vertebra (D5) (mounted);
ISIR 124, mid-dorsal vertebra (D6) (mounted); ISIR 125, mid-
dorsal vertebra (D7) (mounted); ISIR 126, mid-dorsal vertebra
(D8) (mounted); ISIR 127, mid-dorsal vertebra (D9) (mounted);
ISIR 128, mid-dorsal vertebra (D10) (mounted); ISIR 129, mid-
dorsal vertebra (D11) (mounted); ISIR 133, posterior caudal
vertebra: ISIR 134, posterior caudal (Ca43) (last in the mount);
ISIR 700, mid- to caudal dorsal vertebra; ISIR 701, third dorsal
vertebra; ISIR 702, isolated cervico-dorsal vertebra; ISIR 703,
second dorsal vertebra; ISIR 717, complete right upper tooth;
ISIR 718, small complete tooth; ISIR 719, tooth with complete
root but incomplete crown; ISIR 720, tooth crown only; ISIR
721, tooth crown only; ISIR 722, tooth crown only; ISIR 723,
distal chevron pair; ISIR 724, distal chevron pair; ISIR 725, sin-
gle chevron co-ossified with the proximal part of the right femur
of the mounted skeleton; ISIR 726, almost complete dorsal ver-
tebra (D13) (mounted); ISIR 727, almost complete mid- to cau-
dal dorsal vertebra with slit-like neural canal; ISIR 728, most
elongate cervical vertebra (mounted); ISIR 733, distal caudal ver-
tebra (Ca 34); ISIR 734, distal caudal vertebra (Ca 32); ISIR 737,
isolated sacral vertebra ISIR 739, distal caudal; ISIR 740, pubis
found near ISIR 739; ISIR 741, small right femur; ISIR 743, right
calcaneum; ISIR 745, anterior caudal vertebra (Ca 1 series); ISIR
746–747, mid-caudal (Ca 13–14); ISIR 748, distal caudal (Ca 33);
ISIR 749, large ungual phalanx of left digit I of manus; ISIR770,
almost complete posterior dorsal vertebra (D14) (mounted).
Besides the first excavation during 1960–61, another small
excavation was carried out in 1961–62 in north of the village
Krishnapur (19�13¢20¢¢N, 79�31¢18¢¢E), Adilabad district, Andhra
Pradesh, India (Material B).
Material B. ISIR 65, left metatarsal I (mounted); ISIR 66, left
metatarsal II (mounted); ISIR 67, left metatarsal IV (mounted);
ISIR 77, right tibia; ISIR 93, right humerus; ISIR 103, right tibia
(mounted); ISIR 104, right fibula (mounted); ISIR 107, right
astragalus.
A third excavation of very limited scope was attempted in
1964 in the same locality at Sironcha, and those collections
(Material C) are referred as ‘Colbert excavation’ in memory of
late Prof. Edwin H. Colbert’s participation (Colbert 1980, 1989).
TEXT -F IG . 2 . Bone distribution and bone association of Barapasaurus tagorei. The bones collected from the ‘Colbert excavation’ are
in the inset. The associated bones as occurred in the field are within the grey boundary and marked here as A, B, C, F, G, H and J. D
and E (with different shades) indicate two different associated vertebrae. This is not a complete plan; the overlapping and the surface
bones are omitted. Detailed information will be found in the text. Fossil tree logs are stippled.
540 P A L A E O N T O L O G Y , V O L U M E 5 3
Material C. Colbert collection. ISIR 113, left ilium; ISIR 120, cer-
vical vertebra; ISIR 135, right femur; ISIR 136–143, co-ossified
neural arches with spine; ISIR 144–149, caudal centra; ISIR 704,
caudal centrum.
There is no evidence that this material includes more than
one species.
Institutional Abbreviations. GSI, Geological Survey of India,
Kolkata, India; ISI, Indian Statistical Institute, Kolkata, India
(ISIR, prefix to specimen number indicating reptile collection).
Abbreviation of the vertebral laminae used in the text. cdl, ante-
rior centrodiapophyseal lamina; acpl, anterior centroparapo-
physeal lamina; cpol, centropostzygapophyeal lamina; cprl,
centroprezygapophyseal lamina; pcdl, posterior centrodiapophy-
seal lamina; pcpl, posterior centroparapophyseal lamina; podl,
postzygodiapophyseal lamina; posl, postspinal lamina; ppdl,
paradiapophyseal lamina; prdl, prezygodiapophyseal lamina; prsl,
prespinal lamina; spdl, spinodiapophyseal lamina; spol, spin-
opostzygapophyseal lamina; sprl, spinoprezygapophyseal lamina;
tpol, intrapostzygapophyseal lamina; tprl, intraprezygapophyseal
lamina.
SYSTEMATIC PALAEONTOLOGY
DINOSAURIA Owen, 1842
SAURISCHIA Seeley, 1888
SAUROPODOMORHA von Huene, 1932
SAUROPODA Marsh, 1878
BARAPASAURUS Jain, Kutty, RoyChowdhury and Chatterjee,
1975
Barapasaurus tagorei Jain, Kutty, RoyChowdhury and
Chatterjee, 1975
Text-figures 3–16
Type specimen. Sacrum (ISIR 50).
Repository. Palaeontological collection, Geological Studies Unit,
Indian Statistical Institute.
Locality and horizon. Pochampalli (18�44¢N, 80�05¢E), Gadchir-
oli district, Maharashtra, India; Early to early Middle Jurassic
Kota Formation, Pranhita-Godavari basin, Deccan, India.
Remarks. The following amended diagnosis, representing a
unique combination of characters including at least one autapo-
morphy, the slit-like opening of the neural canal of the dorsal
vertebrae, distinguishes the species.
Amended diagnosis. Large sauropod with slender limbs; teeth
spoon-shaped with bulbous base and with grooves on anterolabi-
al and posterolingual sides of the crown; coarse tubercles mostly
on posterior carina; cervical and cranial dorsal vertebrae opistho-
coelous while others platycoelous; primitive acamerate vertebrae
morphology in cervical and cranial dorsal vertebrae; intra-
prezygapophyseal laminae joined the prezygapophyses on the
midline at the dorsal margin of the neural canal in the caudal
cervicals; hyposphene-hypantrum articulation well-developed in
middle and caudal dorsal vertebrae; lateral laminae of the middle
and caudal dorsal neural spines composed of spinodiapophyseal
and spinopostzygapophyseal laminae; dorsal neural spines flat-
tened craniocaudally but wide transversely; neural canal in the
mid-dorsal vertebrae open dorsally through a narrow slit-like
opening into a large cavity; sacrum with four co-ossified verte-
brae; sacral centra hour glass-shaped and amphiplatyan; sacral
neural spines high; sacricostal yoke set close together; distal cau-
dals spool-shaped with caudally inclined neural spine; y-shaped
chevrons with fused cranial and caudal projections; scapula with
tall narrow blade; coracoid subcircular with coracoid foramen;
humerus with prominent deltopectoral crest, expanded at both
ends; ulna with triradiate proximal end stouter than radius but
slender in shaft; ilium with prominent preacetabular process;
medial wall of acetabulum quite deep; ischiadic peduncle of ilium
short, while pubic peduncle long and directed downward and a
little cranially; pubis and ischium almost of same length; pubis
with a large obturator foramen and the pubic apron articulated
with its partner along nearly the full length of the midline result-
ing a narrow pelvic basin; ischium slender, straight and distally
moderately expanded; the symphyseal contact narrow; femur
long and slender with hemispherical femur head set at right angle
to the straight, slender shaft; well-developed fourth trochanter
projecting caudally a ridge-like process with an acuminate and
declined tip; short, robust tibia with well-developed cnemial
TEXT -F IG . 3 . Schematic drawing of left side of mounted skeleton of Barapasaurus tagorei in modern pose with elevated tail at the
Indian Statistical Institute.
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 541
crest; fibula slender with weakly developed lateral trochanter;
tibia articulating with prominently raised elliptical rugosity on
the lateral side of the fibula; subtriangular astragalus bearing
proximally a prominent ascending process and a medial depres-
sion; calcaneum quadrangular but proximally semi-trapezoidal;
distal articular surface for the metatarsals with a fine mediolateral
ridge.
OSTEOLOGY
Most of the elements of Barapasaurus were not found in
direct association. While examining the nature of the
association of some specimens, partial associations of the
bones could be established. The holotype sacrum (ISIR
50) of B. tagorei is associated with right and left ilium
(ISIR 51 and ISIR 111) indicating the presence of a med-
ium-sized individual (Text-fig. 2, Association A). One
partial skeleton (ISIR 113, ISIR 120, ISIR 135–149) con-
sisting of limb bones, pelvic bones and vertebrae is a
nearly mature adult as indicated by ossification of neural
arches. These were excavated in a condition of disarticula-
tion, but in close association from the ‘Colbert excava-
tion’. From this skeleton, proportions can be established
for certain limb elements, and some of the vertebrae can
be identified by region (Text-fig. 2, Association J).
Skeletal elements of a small individual are so far the
most complete association in the collection. This includes
left scapula (ISIR 68) and complete coracoid (ISIR 69),
left humerus (ISIR 70) and left radius and ulna (ISIR 71
and ISIR 72). In the same association were found the
right ilium (ISIR 52), right and left ischium (ISIR 115
and ISIR 54), right and left pubis (ISIR 57 and ISIR 117),
left femur (ISIR 60), left tibia and fibula (ISIR 62 and
ISIR 64) (Text-fig. 2, Association C).
Published descriptions of B. tagorei include Jain et al.
(1975, 1979). The following osteological descriptions are
based on all available material in the ISI collections. The
side view of the skeleton of B. tagorei (Text-fig. 3) sum-
marizes our present understanding of the anatomy of this
dinosaur in a contemporary interpretation of stance and
posture.
Skull and teeth
No skull bones are known. Several teeth were recovered with the
B. tagorei skeleton. Jain et al. (1975) described these teeth as
spoon-shaped, with anterior and posterior keels bearing coarse
denticles. This dentition (Text-fig. 4) includes three nearly com-
plete teeth and three crowns without roots.
tutu
tu
tu
tu
e
20 mm
A
H I J K L M N O
B C
D
F G
E
TEXT -F IG . 4 . Teeth of Barapasaurus tagorei. A–E, complete tooth ISIR 717 with A, root and crown in side view. B, other side view.
C, buccal aspect. D, close-up view of crown. E, close-up, oblique view of occlusal surface. F–G, ISIR 722, an isolated crown in two
views, orientation uncertain. H–I, ISIR 721, isolated crown in H, lingual and I, buccal views. J–K, ISIR 720, isolated crown in two
views, orientation uncertain. L–M, ISIR 719, a medium-sized complete tooth with root and crown in L, lingual and M, buccal views.
N–O, ISIR 718, a small complete tooth with root and crown in two views, orientation uncertain. D and E enlarged, all others at 2-cm
scale. Abbreviations used in the text-figures: a, acetabulum; as, astragalus; c, caudal vertebrae; ce, cervical vertebra; co, coracoid; c-fi,
fibular facet of calcaneum; cf, wing of fan suspended from ceiling; cl, ceiling light; e, enamel ridge of occlusal surface; f, femur; f-l, left
femur; f-r, right femur; f4t, fourth trochanter; fi, fibula; gl, glenoid cavity; h, humerus; il-l, left ilium; il-r, right ilium; is, ischium; is-l,
left ischium; is-r, right ischium; Mt1, metatarsal 1; Mt 2, metatarsal 2; Mt 3, metatarsal 3; Mt 4, metatarsal 4; Mt 5, metatarsal 5; nc-i,
interior cavity of neural canal; p, pubis; p-l, left pubis; p-r, right pubis; pp, pubic peduncle; sc, scapula; s, sacral vertebra; s-1, first
sacral vertebra; t, tibia; tu, tubercle.
542 P A L A E O N T O L O G Y , V O L U M E 5 3
The following description for the spoon-shaped teeth adopts
the orientation terminology of Calvo (1994). Moreover, the
outer ‘labial’ surface of the crown is described below as the distal
surface, and the inner ‘lingual’ surfaces of the crown as the
mesial surface. Grooves are situated on the anterolabial and pos-
terolingual sides of the crown. The largest tooth (ISIR 717, Text-
fig. 4A–E) consists of a nearly complete root, missing only its
tip, and nearly complete crown, missing parts of the apex. It is
58 mm tall (root tip to apex diameter). Crown dimensions are
15 mm maximum anterior-posterior (carina-carina) diameter,
and 24 mm maximum height. In overall shape, the tooth is only
slightly curved in anterior aspect and straight in lingual aspect
with slight asymmetrical bulge of the crown. The tapered root is
subcircular in cross-sectional aspect, and is slightly constricted at
its junction with the crown, which is also constricted at its base
and subcircular. By Calvo’s (1994) definition of anterior-poster-
ior, this tooth is right upper or left lower; its straight profile
suggests this tooth is maxillary rather than mandibular.
Most of the enamel on the crown is weakly wrinkled, visible
only under magnification, as in almost all basal sauropods (Bar-
rett and Upchurch 2007). Along its apicobasal axis (root tip to
apex), the labial surface of the crown is convex; the lingual surface
along the same axis is weakly sigmoid, producing a spatulate pro-
file in lateral or mesial aspect. Its labial surface is convex in over-
all cross-sectional shape, but with a weakly developed groove on
the apical one-third of the crown near the anterior carina. The
lingual surface is likewise convex in cross-section, but with a
weakly developed groove near the opposite (posterior) carina.
These labial and lingual grooves are primitively present in eusau-
ropods (Barrett and Upchurch 2007). This tooth is missing part
of the apex, but its bulbous base retains anterior and posterior
carinae, with three tubercles on the posterior carina. These tuber-
cles are coarse, each with an overall scalloped, asymmetrical out-
line, shallow profile proximally and steep profile apically.
Upchurch et al. (2007a) reported in the discussion of their char-
acter state C86 that Barapasaurus possesses coarse denticles with a
A
co
h
sc
1
1
C C co
2 3 411
12 13 14 15
2 3 45 6 7 8 9 10 il-r
il-l
il-l
8
9
10
11
12
13
14
il-ril-l
il-l
S-1pp
il-r
14
13
12
11
15
15
cf
clf
isp
11 12 13 14 15
B
C D
E F
TEXT -F IG . 5 . Cervical and dorsal
vertebrae of Barapasaurus tagorei,
mounted skeleton. A, cervicals and
cranial dorsals, left side. B, middle and
caudal dorsals, left side. C, close-up of
caudal cervicals and cranial dorsals, left
side. D, close-up of caudal dorsals, left
side. E, caudal dorsals, cranial sacral and
cranial pelvis, ventral aspect. F, caudal
dorsals, postero-oblique aspect, numbers
placed on tips of neural spines; 1–14,
dorsal vertebrae.
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 543
45 degree orientation on the basis of a published figure (Kutty
et al. 2007, fig. 8). However, a close look at ISIR 717 reveals that
the denticles are actually set at higher angles with the long axis;
the angle of the individual denticle varies from c. 60–70 degrees
to the long axis (Text-fig. 4C–D). This tooth does not show evi-
dence of an apical wear facet, perhaps because part of the apex is
missing. The crenulated enamel in the labial and lingual grooves
lacks any indication of occlusal wear.
The three crowns (ISIR 720–722) (Text-fig. 4F–K) are roughly
the same sizes as ISIR 717 and possess nearly identical anatomy,
with only slight variation. In all three, the carinae are incom-
plete. These teeth are spatulate, with bulbous base, and tapering
profile in lingual and labial aspect. ISIR 721 has two weakly
developed tubercles on its posterior carina. These tubercles,
which are slightly raised surfaces of the enamel, extend onto the
lingual surface as weakly expressed, linear wrinkles in the crenu-
lations, but show no evidence of striation or wear. ISIR 720 has
a slightly narrowed crown, but nearly complete apex, demon-
strating the complete shape of the crown; it lacks evidence of
wear facets, perhaps because the carinae are incomplete.
Tooth ISIR 719 (Text-fig. 4L–M) is approximately two-thirds
the size of the previously described teeth. Most of its root and
crown are intact, but the tooth is poorly preserved. A much
smaller tooth (ISIR 718) (Text-fig. 4N–O) is nearly complete,
with elongate root and reduced crown. In overall shape, this
tooth resembles a mammalian incisor. The bulbous crown is
truncated by a beveled surface, apparently a lingually inclined
wear facet. This is probably the tooth of a juvenile individual.
In the two teeth that have tubercles, they occur only on the
posterior carina. None have tubercles on the anterior carina, a
condition that might be attributed to incomplete preservation.
Presacral vertebrae
Partial vertebral associations and a number of isolated vertebrae
were collected. Three pairs of dorsal vertebrae, ISIR 123 and ISIR
124, ISIR 726 and ISIR 770 (Text-fig. 2, Association D) and ISIR
127 & ISIR 128 (Text-fig. 2, Association E), which are very likely
to be adjacent ones, belong to such associations. Of the other
100 mm
A B C D
EF
K
L
JGH
I
nc-i
TEXT -F IG . 6 . Dorsal vertebrae of Barapasaurus tagorei, isolated elements. A–B, ISIR 701, posterior cervicodorsal in A, left and B,
posterior aspects. C–D, ISIR 703, dorsal vertebra in C, anterior and D, left aspects. E–F, ISIR 700, dorsal vertebra in E, anterior and F,
left aspects. G–H, ISIR 702, dorsal vertebra in G, anterior and H, left aspects. I–L, ISIR 727, dorsal vertebra in I, anterior; J, left; K,
dorsal and L, anterodorsal oblique aspects; tipped to provide perspective with respect to J and K.
544 P A L A E O N T O L O G Y , V O L U M E 5 3
two, one consists of eight articulated neural arches (ISIR 136,
ISIR 137, ISIR 138, ISIR 139, ISIR 140, ISIR 141, ISIR 142 and
ISIR 143) of a small individual from ‘Colbert excavation’ and a
series of caudal centra (ISIR 144–ISIR 149) found detached from
the neural arches but lying close to them (Text-fig. 2, Association
J). These cover the region of the caudal cervicals and cranial dor-
sals. The third association includes five vertebrae (ISIR 728, ISIR
79, ISIR 81, ISIR 80 and ISIR 122) of a medium sized individual,
disarticulated but lying close together and belongs to the cervico-
dorsal region (Text-fig. 2, Association B).
Precise count of presacral vertebrae (cervicals and dorsals) is
not possible to determine, as there was no completely associated
vertebral column. However, it is assumed by analogy with other
related forms (sauropods and prosauropods) that there were 26
presacral vertebrae. Except for the associated caudal cervical and
the cranial dorsals, there are very few duplicate specimens in the
collection of the presacral vertebrae. The material in the collec-
tion suggests that B. tagorei had at least 14 dorsal vertebrae as
indicated by the cervico-dorsal transition determined on the
basis of the available associations of this region. Moreover, the
caudal cervicals show certain modification from the typical cer-
vical structure. The available material does not allow a complete
cervical count and it is assumed that there were at least eight
cervical vertebrae posterior to the atlas-axis. Thus, this set of
eight, the atlas-axis, and the two caudal cervicals give a mini-
mum count of 12. We identify the next vertebra as the first dor-
sal, D1 rather than C13. According to Wilson and Sereno (1998),
Upchurch (1995) and Upchurch et al. (2004) basal sauropods
had 13 cervicals, 13 dorsals, and a minimum count of 26 presac-
rals. Because this count is not inconsistent with the typical pre-
sacral count in basal sauropods, we assume that B. tagorei had
12 cervicals and 14 dorsals making up the presacral region.
The arranging of the presacrals in their proper order is essen-
tially ordering of the dorsals. There is no atlas or axis in the
C B
100 mm
16 15 14 13 12 11 109
8
403530
25
20
15
109
87
654321
43
A
D
TEXT -F IG . 7 . Tail of Barapasaurus tagorei. A, tail of mounted skeleton, left side in photomosaic. B, photograph showing detail of
articulated mid-caudals, right side. C, isolated anterior chevron affixed to right femur on mounted skeleton, posteroproximal
extremity, in photograph and line drawing interpretation. D, ISIR 723, isolated posterior chevron in dorsal, lateral-oblique, and ventral
aspects, as photographs and line drawing interpretations. Numbers indicate caudal position.
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 545
collection. The last cervical vertebra (C12) is ISIR 728 (Text-
fig. 5, labeled ce-12), and the first four dorsals (D1–D4) are ISIR
79, ISIR 81, ISIR 80 and ISIR 122, respectively (Text-fig. 5,
labeled 1–4). Lack of sufficient cranial cervical and mid-cervical
vertebrae precludes proper ordering in this region. Among the
collected cervicals, ISIR 121 has been separated as C10 from C12
(ISIR 728) but it is not possible to distinguish it from the other
cranial or mid-cervicals. For the ordering of dorsals D5–D14, ver-
tebrae from this region were first placed into groups on the basis
of some broad characters. For example, on the basis of the struc-
ture of the neural canal and associated features on the neural
arch ISIR 126 (D8), ISIR 127 (D9), ISIR 128 (D10) and ISIR 129
(D11) are placed in one group and ISIR 74 (D12), ISIR 726
(D13), ISIR770 (D14) in another. The latter is placed as the cau-
dal set of dorsals, i.e., D12–D14, on the basis of its similarities
with the vertebrae in the sacrum. ISIR123 and ISIR 124 are
placed as D5 and D6 as indicated by their similarities with D4,
especially in the position of the zygapophyses. Similarly, ISIR
125 (D7) based on its similarities with D6 on one hand and the
ISIR 126 (D8) group on the other. Further ordering within the
groups is based on details of individual vertebrae. In the follow-
ing description of the vertebrae, the abbreviated form of the
nomenclature of the vertebral laminae as proposed by Wilson
(1999) have been used (for details, please see above).
Cervical vertebrae. Jain et al. (1975, 1979) described the cervical
vertebrae as opisthocoelous, with centra probably a little less
than twice the length of dorsal centra. The caudal cervicals
increase in length rearward. Because only the caudal cervicals are
known, this proportion applies only to the caudal-most cervical
of Barapasaurus. (The exact positions of other cervical vertebrae
in the collection are not clear). On the mounted skeleton, the
caudal cervicals increase in length rearward.
Cervical vertebrae are deeply opisthocoelous with markedly
convex cranial facets. The centra are elongate, longer than tall.
They have a weakly developed double-keel construction ven-
trally, producing a flattened ventral surface and slightly squared
cranial ⁄ caudal profiles of the articular facets. In lateral aspect,
the vertebrae are constricted at mid-centrum, and a well-defined
ridge is present along the ventrolateral edges; the diapophysis
position is low. Facets for cervical ribs are indistinct.
A
C
B
D
sc
sccogf
sc
coco
sc
scco
co sc
co
gf
100 mm
E
TEXT -F IG . 8 . Scapula and coracoid of Barapasaurus tagorei. A–C, mounted skeleton from three viewing angles of the left scapula-
coracoid complex in overall dorsolateral aspect. A, a slightly anterior perspective. B, lateral aspect of scapula with lateral perspective of
the coracoid in articulation. C, anterodorsal aspect, cranial surface of coracoid and cranial edge of scapula in articulation. D–E, line
drawing interpretation based on stylized reconstruction with restoration of missing parts after left scapula on mounted skeleton; D, in
medial aspect, with internal surface of coracoid curving upward towards viewer from plane of scapula; E, line drawing of the
conjoined right scapula and coracoid in lateral aspect. 10-cm scale bar applies only to D and E.
546 P A L A E O N T O L O G Y , V O L U M E 5 3
Jain et al. (1979, pl. 98A) illustrated the largest cervical verte-
bra (ISIR 728), which we assign to position C12. Its centrum is
approximately two times longer than tall. In lateral aspect, the
cranial and caudal facets are not parallel; instead their extended
outlines converge ventrally (Text-fig. 5C). This orientation indi-
cates an arched profile in the caudal cervical series. Cervicodor-
sals display a similar condition, described below. The acdl
extends from the diapophysis to the cranial part of neurocentral
suture, whereas the pcdl connects the diapophysis to the caudal
part of the neurocentral junction.
A B C D
F
E
100 mm
G H I JL
M N O P
R
Q
K
TEXT -F IG . 9 . Forelimb of Barapasaurus tagorei. A–F, ISIR 70, left humerus in A, anterior, B, lateral, C, posterior, D, medial, E,
proximal, and F, distal aspects. G–L, ISIR 72, left ulna in G, anterior, H, lateral, I, posterior, J, medial, K, proximal, and L, distal
aspects. M–R, ISIR 71, left radius in M, anterior, N, lateral, O, posterior, P, medial, Q, proximal, and R, distal aspects.
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 547
The parapophyses of the caudal cervicals occur on an outward
and downward projecting buttress of the ventrolateral ridge just
behind the rim of the cranial face of the centrum. The parap-
ophyses on C12 (ISIR 728) are larger, project downward and
outward and originate on the lateral sides of the dorsal to the
ventrolateral ridge. The prezygapophyseal facets are projected
more forward than outward from the cranial margin. The prezy-
gapophyses are subcircular and fairly highly tilted. The cprl
extend from the anterolateral margin of the centrum to the pre-
zygapophyses. The prdl (for details, please see above) connect
the lateral surfaces of the prezygapophyses with the cranial part
of the transverse process. The transverse processes originate from
anterior position low on the lateral surface of the neural arch.
The postzygapophyseal facets are shortened but not very dis-
tinct. The podl connect the diapophyses posterodorsally to the
postzygapophyses and are almost parallel to the sprl which con-
nect the prezygapophysis with the neural spine.
Part of the neural spine and neural arch is preserved in the
tenth vertebra (ISIR 121). The neural spine appears to be short,
craniocaudally elongated and laterally compressed near its base.
At the mid-length, the spine expands upward, giving it a dia-
mond-shaped outline for the top of the spine. The prespinal
cavities are deeper than postspinal cavities; however, the depth
reduces caudally. The cranial and caudal openings of the neural
canal are triangular and low in position, with a relatively broad
base in outline. The cranial and caudal faces of the neural arch
are concave on both sides.
Cranial dorsal vertebrae (D1–D3). Jain et al. (1979) identified
two cranial dorsal vertebrae as opisthocoelous. In the mounted
skeleton, these two vertebrae (ISIR 79 and ISIR 81) are immedi-
ately posterior to the longest cervical (Text-fig. 5A, C). The cra-
nial dorsal centra are the longest in the dorsal series. Their rib
facets are situated high on the neural arch, and their neural
spines are transversely expanded, indicating without doubt that
they are dorsal vertebrae. Because their anatomy is distinctive,
these two vertebrae represent separate vertebral positions, D1
and D2 respectively, as mounted.
The ventral surface of the opisthocoelous centrum of the cra-
nialmost of the two mounted cranial dorsal vertebrae (ISIR 79)
is craniocaudally concave, gently convex transversely, weakly
double-keeled like the condition in the caudal cervical described
above, and in contrast to succeeding dorsal vertebrae in which
the ventral surfaces of the centra are flat. In lateral aspect, the
projected planes of the centrum in this vertebra, like those of
the caudal cervical described above, converge ventrally, contin-
uing the arch of the cranio-dorsal region. Accordingly, this ver-
tebra is assigned to the cranialmost position (D1) among the
cranial dorsals. The centrum has deep cavities on the lateral sur-
faces; neither penetrates to the interior of the centrum; these
depressions are not pleurocoels. Britt (1993; cf. Wedel et al.
2000) termed these lateral excavations as pneumatic fossae.
Wedel et al. (2000) characterized these pneumatic fossae to be
broad in contour but not enclosed by ostial margin to form a
foramen. They suggested that Barapasaurus has primitive aca-
merate vertebrae morphology in which pneumatic fossae are
present but do not significantly invade the centrum.
The parapophyses are small and originate on the lateral face of
the centrum with the ventral border still on the ventrolateral
ridge as in the caudal cervicals. One sharp, well-defined dorsal
A B C
D
100 mm
M
LKJI N O P Q
G
H
R
FE
TEXT -F IG . 10 . Manus of Barapasaurus tagorei. A–D, ISIR 108, left metacarpal I in A, anterior, B, lateral, C, posterior, and D,
proximal aspects. E–H, ISIR 94, left metacarpal III in E, anterior, F, lateral, G, posterior, and H, proximal aspects. I–M, ISIR 96, left
metacarpal IV in I, anterior, J, lateral, K, posterior, L, medial, and M, proximal aspects. N–Q, ISIR 749, ungual phalanx, left digit I
in N, anterior, O, lateral, P, medial, and Q, proximal aspects. R, ISIR 110, ungual phalanx, left digit I of a young individual, in medial
aspect.
548 P A L A E O N T O L O G Y , V O L U M E 5 3
ridge extends cranially and another caudally on the neurocentral
suture. The rib facets are situated high on the neural arch, and
their neural spines are transversely expanded. From the diapoph-
ysis, a lateral lamina, the acdl and another lamina, the pcdl,
extend downward and join the cranial and the caudal part of the
neurocentral contact. The prezygapophyses are slightly elongated
craniocaudally and extend well beyond the plane of the cranial
centrum facet. The prezygapophyses are joined on the midline at
the dorsal margin of the neural canal by tprl. The prezygapophy-
seal process is supported by the prominent cprl connecting the
prezygapophyses ventrolaterally with the cranial part of the cen-
trum and by the prdl extending laterally from the prezygapophy-
sis to the cranial part of the diapophysis which occurs above the
neurocentral junction. The postzygapophyseal facets are large,
subcircular, moderately tilted and are set well apart. A pair of
laminae originates from the postzygapophysis; the podl connects
the postzygapophysis to the diapophysis, while cpol connects the
postzygapophysis with the caudal part of the neurocentral con-
tact. The cranialmost dorsal vertebra (D1) has a coalesced neural
spine, which is short, transversely expanded, convex on its cranial
surface and concave on its rear surface. The neural arch arises
from the cranial half of the centrum. The sprl flares from the
caudal part of the prezygapophysis towards the cranial surface of
the neural spine; the spol extends from the postzygapophysis to
the caudal surface of the neural spine. On the ventral surface, a
median keel and sharp ridges on the ventrolateral edges are well
defined.
The second cranial dorsal vertebra (D2) (ISIR 81) in the
mounted skeleton is taller, but its centrum length is shorter
than the previous vertebra. An isolated vertebra (ISIR 703)
A
E
B
C
D
100 mm
TEXT -F IG . 11 . Ilium of Barapasaurus tagorei. A, ISIR 51, left ilium as mounted, in articulation with pubis, ischium and femur,
lateral view. B–D, ISIR 52, right ilium in B, lateral and C, medial aspects, and D, oblique view looking into acetabulum, ventral aspect.
E, left ilium as mounted, with proximal femur, proximal ischium, sacral spines, and proximal caudal vertebrae, oblique posterolateral
aspect.
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 549
(Text-fig. 6C–D) is similar to ISIR 81 with respect to its cen-
trum proportions, but is somewhat smaller in overall dimen-
sions. Because it is detached and can be examined from all sides,
this vertebra adds considerable knowledge of this position in the
vertebral column. These two vertebrae represent a position cau-
dal to the first cranial dorsal, perhaps the next vertebra in suc-
cession and are assigned the position of second dorsal (D2). Like
the first cranial dorsal, D2 is opisthocoelous. The centrum is
deeply constricted, and the planes of the centrum facets are
inclined forward with respect to the long axis of the centrum,
the third vertebra in the cervicodorsal region to continue the
arched profile in lateral aspect. This vertebra has shallow depres-
A B
100 mm
C D
cc c
cc
f-ris-ris-l
il-l il-r
is-ris-l
il-l
ilpp-ls
il-r
ilpp-r
p p
l-fr-f
f-l
cc
cc
f-l
p
E F G H K
J
I
TEXT -F IG . 12 . Pelvis of Barapasaurus tagorei. A–D, ISIR 54, right ischium in A, anterior, B, lateral, C, posterior and D, medial
aspects. E–H, ISIR 57, right pubis in E, lateral, F, posterior, G, medial, and H, distal aspects. I, posterior view of pelvis in mounted
skeleton, showing left and right ischia projecting rearward towards viewer with tail deflected to right side as mounted, obscured pubes
projecting forward, caudal vertebrae and left and right femora. J, oblique view of pelvis of mounted skeleton, left side, showing
articulation of left and right ischia, left femur, lateral surface of left ilium, medial surface of right ilium and anterior caudal vertebrae.
K, anterior view of pelvis in mounted skeleton from within ribcage, showing left and right ischia, pubic peduncle of ischia, left and
right pubes, sacral vertebrae and left and right femora.
550 P A L A E O N T O L O G Y , V O L U M E 5 3
sions on the lateral surface of the centrum, but they are not
pleurocoels. Two laminae, the ppdl and the acpl originating
from the parapophysis (which has moved upward compared to
its position on D1), extend to the diapophysis and the cranial
part of the centrum, respectively. The diapophysis is lower than
on the succeeding dorsals. The pcdl extends from the diapophy-
sis to the caudal part of the neurocentral junction.
The prezygapophysis is shortened, and the facets are more or
less subcircular, less tilted and project more outward than for-
ward. Like the first cranial dorsal vertebra, the tprl form wing-
like expansions that buttress the prezygapophyseal process. The
prdl and the cprl extend from the prezygapophysis to the diap-
ophysis occurring above the neurocentral junction and the cra-
nial part of the centrum, respectively. The postzygapophysis is
shortened, and the facets are closer to the neural spine but away
from each other. The podl extends from the postzygapophysis to
the diapophysis. The prominent transverse processes are
supported by the tprl. The neural arch arises on the cranial
A B C
D
f4t 100 mm
E F
TEXT -F IG . 13 . Femora of
Barapasaurus tagorei. A–D, ISIR 741
right femur in A, anterior, B, lateral, C,
posterior, and D, proximal aspects; E–F
ISIR 58, right femur in the mounted
skeleton in E, lateral and F, posterior
aspects.
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 551
two-thirds of the centrum and is more erect than in the first
cranial dorsal vertebra, and the diapophysis is lower than on
succeeding dorsals. The rib facets, like those on the first cranial
dorsal, are situated high on the neural arch. The neural spine is
flat and transversely expanded dorsally. The sprl extends from
the prezygapophysis to the cranial surface of the neural spine;
the spol is not clearly preserved.
The third cranial dorsal vertebra (D3) in the mounted skele-
ton (ISIR 80) with its opisthocoelous centrum is quite similar to
the previous vertebra barring its tall height and lower centrum
length. The width of the centrum is narrow in the middle, less
than half of the cranial face. The large parapophyses face more
outward than downward. The acpl develops as a supporting
ridge extending from the parapophyses to the centrum. The ppdl
extends from the parapophysis to the diapophysis. The prezyga-
pophyseal facets are narrow, less tilted and more transversely
elongate than the previous vertebra and are projected more out-
ward than forward from the neural spine. The tprl meet just
dorsal to the neural canal opening and then descend as a single
lamina. The prdl connects the prezygapophysis to the diapophy-
sis above the neurocentral junction. The postzygapophyseal fac-
ets, however, are closer together. Accordingly, the tpol meet
some way up from the neural canal opening and descend as a
single lamina. The transverse process arises on the neural arch to
A B C D
f
fit
as Mt III
Mt II
Mt I
Mt V
Mt IV
J
F G H IK
L
E
100 mm
TEXT -F IG . 14 . Lower leg of Barapasaurus tagorei. A–E, ISI R62, left tibia in A, anterior, B, lateral, C, posterior, D, medial, and E,
distal aspects. F–K, ISIR 64, left fibula in F, anterior, G, lateral, H, posterior, I, medial, J, proximal and K, distal aspects. L,
photograph of lower limb, left rear leg on mounted skeleton, posterior aspect; astragalus in oblique posterodorsal aspect showing
articular surface for reception of femur and posterior margin. A–K, scale as shown; L, enlarged for details in articulation. The
calcaneum, which was not mounted in this skeleton, would fit between the distal extremity of the femur and metatarsal I–III.
552 P A L A E O N T O L O G Y , V O L U M E 5 3
a height roughly equalling the centrum height and it remains
more or less in the same position in the subsequent dorsal verte-
brae. It is directed outward and very slightly upward and located
closer to the postzygapophyses than the cranial dorsals. The podl
extends from the postzygapophysis to the diapophysis. The neu-
ral canal is tabular, with an oval cross-section in the middle and
the height of the neural canal is relatively larger than its width.
The floor of the canal is slightly sunk into the body of the cen-
trum. The neural spine is flattened craniocaudally and wide
transversely. The transverse width reduces considerably towards
the base, more on the cranial face than on the caudal.
The dorsal vertebrae of Barapasaurus have hollow neural
spines whose chambers communicate directly with the neural
canal (Jain et al. 1979; Britt 1993; Wedel 2003). Wilson (1999)
interpreted the vertebral laminae as primarily pneumatic in ori-
gin but also have a secondary function, and these laminae
evolved initially to partition pneumatic diverticula. Wedel
(2003) concluded that the presacral vertebrae of basal sauropods
were probably pneumatized by diverticula of cervical air sacs.
On the basis of presence of neural spine laminae and suprame-
dullary chambers, he further suggested that the presacral verte-
bral pneumatic fossae of Barapasaurus are osteological correlates
of a system of pneumatic diverticula.
The anatomy of another isolated opisthocoelous dorsal verte-
bra (ISIR 701) (Text-fig. 6A–B) assigned to position D3 differs
from the previously described dorsals in several respects. The
planes of the articular facets of the centrum are orthogonal with
respect to the long axis of the centrum, and the neural arch is
simple and arises from nearly the full length of the centrum.
The depression on the dorsolateral surface of the centrum is
broad and poorly defined. The centrum shape in lateral aspect is
a modified spool, with broad ventrolateral ridges and a weak
ventral keel.
Another cranial dorsal vertebra with an indeterminate position
from the associated skeleton (ISIR 139) is a complete neural
arch and neural spine. The transversely expanded spine is weakly
convex cranially and weakly concave on its caudal surface. The
neural spine is constricted at its base and expands upward to
form a fan shape in cranial and caudal aspects.
Dorsal vertebrae (D4–D14). Jain et al. (1979) described the first
two dorsal vertebrae (D1–D2) as opisthocoelous; the remaining
dorsals (D3–D14) are essentially platycoelous. The mounted skel-
eton corresponds to the description of Jain et al. (1979) with 14
dorsal vertebrae (Text-fig. 5) of which the first two are opistho-
coelous, and the rest are variously platycoelous or slightly bicon-
cave and short. All the centra are real; neural arches and neural
spines of the few cranial and some caudal dorsal vertebrae in the
mounted skeleton are real; those in the middle are recon-
structed. These dorsal vertebrae form an arch that rises gently
rearward, where the top of the arch is established several posi-
tions cranial to the sacral complex. These vertebrae articulate in
closed-pack condition in the mounted skeleton, and this arch
appears to be natural. The centrum length in these vertebrae is
remarkably uniform and shorter than the centra of the cranial
dorsals. All centra have deep dorsolateral depressions, but none
have pleurocoels. The centra are strongly constricted and spool-
shaped.
The parapophyses in the fourth dorsal vertebra D4 (ISIR 122)
are shared by the neural arch although they are mainly on the
centrum. A vertical ridge supports each parapophysis ventrally
A B
C
Mt IIIMt V IV
Mt
a
fi
Mt I
Mt IIMt III
Mt IV
a
t fi fi t
a
Mt III Mt II
f
Mt IMt IV
Mt VMt V
G
100 mm
D
E
F
TEXT -F IG . 15 . Tarsus of
Barapasaurus tagorei. A–C, right tarsus
in the mounted skeleton in A, posterior,
B, anterior, and C, lateral aspects. D–G,
ISI R743, right calcaneum in D, anterior,
E, proximal, F, distal, and G, medial
aspects. Calcaneum was not installed on
the mounted skeleton. The 10-cm scale
applies to the calcaneum only; scale for
A–C modified slightly to provide
perspective.
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 553
on the lateral face of the centrum. In the succeeding dorsals, the
position of the parapophyses gradually migrates upward and
arises almost entirely on the neural arch thereafter to the caudal
dorsals. The parapophyses from the mid-dorsals onwards are
supported below by the acpl, which joins the ventral end of the
cprl. A pcpl joins the parapophysis to the cranial and posterolat-
eral part of the centrum. The ppdl in the mid-dorsals and caudal
dorsals extend from the parapophyses to the diapophyses.
The prezygapophyses in the mid-dorsals are very low on the
neural arch, with the development of a hypantrum in D5 (ISIR
123) and in the succeeding dorsals. The prezygapophyseal facets
curve downward medially and slightly outward, forming addi-
tional articulation surfaces. The prezygapophyses change to a
higher position from the mid-dorsal towards the caudal dorsal
vertebrae. From the mid-dorsals rearward, the prdl connecting
the prezygapophyses to the diapophyses moves further upward.
The cprl extend from the prezygapophyses to the cranial part of
the centra. The transverse processes in the dorsal vertebrae are
constituted essentially by the combination of prdl, podl and the
posteroventrally directed pcdl. The transverse processes are
mostly directed laterally and are situated high in the caudal dor-
sals. The postzygapophyses become almost horizontal from mid-
dorsals rearward, and the tilt of the facets becomes low and in
the caudal dorsals, the facets are small. The tprl is present in the
mid-dorsals and in the caudal dorsals. The sprl extends from the
prezygapophysis to the cranial surface of the neural spine in
the mid-dorsals. The tpol are also present in the dorsal series.
The cranial face of the neural arch in the cranial dorsals is
concave on either side, but this face on the mid-dorsal vertebrae
is flat, nearly vertical, and bends forward from the parapophyses.
The neural canal in the cranial dorsals is tabular with an oval
cross-section in the middle. The floor of the canal is slightly
depressed on the centrum. The canal is larger, and there is a rel-
ative increase in the height of the openings compared to their
A
H
E F G
I J K
L
B100 mm
C
D
III
I
Mt
Mt II Mt I IV II
aa
a
I
IVMtII
MtI
TEXT -F IG . 16 . Rear foot of Barapasaurus tagorei. A–C, mounted skeleton, left foot in A, anterior, B, medial, and C, posterior
aspects. I, II, IV, metatarsals, original bone; unlabelled elements are restored. D, mounted skeleton, right foot in anterior aspect. E–H,
ISIR 83, isolated right ungual I in E, lateral, F, dorsal, G, medial, and H, proximal aspects. I–L, ISIR 84, isolated right ungual III in I,
lateral, J, dorsal, K, medial, and L, proximal aspects. 10-cm scale identical in A and B slightly larger in C.
554 P A L A E O N T O L O G Y , V O L U M E 5 3
width; caudally, the canal remains more or less the same except
that the cranial opening is more subcircular in D4 (ISIR 122) but
tall and oval in D7 (ISIR 125). The neural canal in the caudal
dorsals is roughly triangular and taller than wide. In the mid-
dorsals, the canal opens dorsally through a narrow slit-like open-
ing into a large cavity. Below this slit-like opening, the canal may
be narrow and deeply sunk on the centrum. The base of the neu-
ral spine forms the roof of the cavity, and the floor of the cavity
is depressed on either side of the slit-like opening. The cranial
opening of the neural canal is surrounded by a deep fossa
between the cprl, the dorsal surface of the centrum and the tprl.
Neural spines are simple and subrectangular in cross-section.
None of the dorsals in the collections has a divided neural spine.
Hyposphene-hypantrum articulations in the middle and caudal
dorsal vertebrae are well developed. The prsl and the posl cover-
ing the cranial aspect of the neural spine from the base to the top
are present in the dorsals. The posl are prominent ridges that
diverge upwards and fade away near the top of the spine. The
neural spines in the cranial dorsals are flattened craniocaudally
but wide transversely. This configuration is even more prominent
in the cervicodorsal vertebrae of the Colbert collection. The cra-
nio-caudal thickness of the neural spine increases caudally. The
cranio-caudal and transverse widths of the neural spines in the
mid-dorsals are roughly equal at the tip. The width of the cranial
face of the neural spine becomes narrow towards the base of the
spine. The caudal face of the spine is, however, wider than the
cranial face. The lateral faces of the neural spines are gently con-
cave in mid-dorsals, but the concavity disappears near the top of
the spine. The spdl extending from the diapophysis to the lateral
part of the neural spine is prominent from the mid-dorsals to the
caudal dorsals; this lamina is joined by the spol at mid-spine
forming a composite lateral lamina (Wilson and Sereno 1998;
Wilson 1999).
Jain et al. (1979) described and figured a caudal dorsal verte-
bra D9 (ISIR 127) with unusual anatomy in the region of the
neural canal and neural arch. This vertebra (Text-fig. 5B) is
complete except for the neural spine. The neural canal is steeply
triangular. The neural arch coalesces fore and aft in weak bridges
of bone that enclose a rectangular depression, which deepens
rearward, then communicates with the neural canal in an elon-
gate slot about 2 cm wide.
One mid- to caudal dorsal vertebra (ISIR 700) (Text-fig. 6E–
F) in the associated skeleton of Colbert collection is platycoelous
and has a rectangular centrum with flattened ventral surface.
The dorsolateral depression is broad and shallow, with no indi-
cation of a pleurocoel. The neural arch arises from the full
length of the centrum, a feature difficult to establish on the
mounted dorsals. This vertebra, like several others in the
mounted skeleton, has an unusual anatomy in the neural canal.
Jain et al. (1979) described this feature as a depression in the
neural arch dorsal to the neural canal, and aptly called it the
dorsal cavity. This intravertebral depression is apparently related
to another modification of the neural canal, as described below.
Another mid to caudal dorsal vertebra (ISIR 727) (Text-
fig. 6I–L) has similar anatomy, including the intravertebral dor-
sal cavity, and a second unusual feature. The expanded cranial
face of the neural arch forms a distinctive, nearly hemispherical
depression. This depression forms the caudal limit of an uniden-
tified intervertebral expansion of the region of the neural canal.
Several of the mounted vertebrae have this feature as well. This
depression communicates horizontally with the neural canal and
the intravertebral dorsal cavity. Jain et al. (1979) described these
depressions as excavations on the cranial and caudal faces of the
caudal dorsals, now in the mounted skeleton. A consequence of
these excavations is confinement of the neural canal cavity
within the neural arch. Thus, the intravertebral cavity for the
neural canal is short and communicates dorsally through a small
opening into the intravertebral dorsal cavity. The confined neu-
ral canal in these vertebrae extends horizontally through its cra-
nial and caudal openings into the much larger intervertebral
depressions. Jain et al. (1979) considered several explanations for
the neural anatomy of the caudal dorsals. According to their
description, sacral vertebrae (now mounted) possess these fea-
tures as well.
Sacral vertebrae
Jain et al. (1975) described the anatomy of the sacral vertebrae
of B. tagorei. The mounted sacrum has four co-ossified sacral
vertebrae with amphiplatyan centra. Neural spines are high, and
the sacricostal yokes are set close together. For completeness,
their measurements of this sacrum are repeated here: greatest
length of the four conjoined centra, 705 mm; greatest width at
first sacral, 460 mm. The sacral ribs are greatly expanded later-
ally where they coalesce to form the sacricostal yoke, for contact
with the ilium; the yoke is long and curved. The fourth sacral
vertebra has enormous sacral ribs that are taller than the cen-
trum. The caudal facet of the centrum of the fourth sacral is
slightly concave. The neural spines of the second and third sacral
are co-ossified. The transversely expanded neural spines are tall
and fan-shaped in cranial aspect. These features are illustrated
with the ilium and pelvis, below.
A large, isolated sacral vertebra (ISIR 737), probably from the
position of sacral 1, lacks the neural arch, which had not fused
to the centrum. This vertebra is as large as those are on the
mounted skeleton. Its lack of ossification indicates that individu-
als could have achieved considerably greater size than indicated
by the mounted skeleton. The floor of the neural canal is repre-
sented by a narrow, continuous groove on the dorsal surface of
the centrum.
Caudal vertebrae
In the mounted skeleton, the caudal series (Text-fig. 7A–B) is
arranged by decreasing diameter of the centra. All caudal centra
are biconcave, and the articular ends have subcircular outline.
Caudal vertebrae lack pleurocoels, but have broad dorsolateral
depressions on the centra. The proximal caudals are axially
shorter than their diameter. Passing distally the centra becomes
elongate or spool-shaped with the decrease in the diameter and
with an increasing length of the centra. The distal caudals are
more rod-like, and a couple of them are fused. Ventrally the
centra are concave, smooth without any keel. Most of the
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 555
caudals have weakly developed chevron facets ventrally, produc-
ing a somewhat squared profile in cranial and caudal aspect.
The first caudal vertebra in the mounted skeleton (ISIR 745)
(Text-fig. 7A) has a complete, triangular neural arch and an
erect neural spine. The transverse process expands laterally and
forms a sturdy wing-like process. Because of this expansion, this
vertebra might be considered a sacro-caudal, but there is no
indication that it contributed to the support of the ilia in this
stage of growth, or that it would have supported the ilia later in
ontogeny. Its centrum is taller (dorsoventral axis) than long
(cranio-caudal axis), whereas mid- and caudal centra are longer
than tall. The caudal face of the centrum of the first caudal ver-
tebra slopes a little caudoventrally at a level below the cranial
face, indicating that the tail is slightly downturned in this region.
However, this feature is not prominent in other cranial vertebrae
of the caudal series. It seems that the tail droops a little behind
the pelvis and then remains straight (Text-fig. 3).
In the first few caudals, the neural arches cover almost the
total length of the centrum, whereas it is confined to the cranial
half of the centrum in middle caudals, while the distal caudals
lack neural arches. The transverse processes in the second caudal
vertebra in the mounted series are expanded laterally, their rela-
tive size reduced rearward but quite prominent to the middle of
the caudal series. The prezygapophyses, preserved in proximal
caudals and Ca14 (ISIR 747) (in the mounted skeleton) project
anterodorsally and bend forward. The postzygapophyses are not
preserved. The neural canal in the first caudal is triangular but
becomes rounded caudally. Caudal vertebrae in positions Ca13-14
(ISIR 746–747) and Ca32 (ISIR 734) include neural spines which
are simple, laterally compressed and project dorsally. In Ca13-14
(ISIR 746–747), the spines are comparatively tall, while in Ca32
(ISIR 734) the spine is short and is inclined backward.
The vertebra in the position of Ca33 (ISIR 748) (Text-fig. 7A)
in the mounted skeleton appears to be nearly complete. Its neu-
ral spine is inclined rearward and broadly expanded. This verte-
bra lacks zygapophyses. The last mounted caudal is in the
position of Ca43 (ISIR 134); it possesses an elongate centrum.
An isolated distal caudal (ISIR 739 found near the pubis, ISIR
740) with a complete neural arch is approximately the same size
as the last several caudals in the mounted skeleton. Its enor-
mous, rod-like neural spine is broadly oval in cross-section and
elongate, extending caudally well beyond the caudal articular
facet. Five isolated caudals (ISIR 704, ISIR 144, ISIR 145, ISIR
146 and ISIR 147) from the Colbert excavation have amphicoe-
lous to amphyplatyan centra. They lack neural arches, which
were separated at their growth lines on the centra, indicating
subadult anatomy. Eleven distal caudals (ISIR 706–716) are simi-
larly elongated, and considering their size, have unexpectedly
robust neural arches. The centra are fused on the two smallest
caudal vertebrae, perhaps representing the terminus of the tail.
The centra are amphicoelous to amphiplatyan, with circular to
ovoid cross-sections.
Chevrons
Three chevrons in the collection (ISIR 723, ISIR 724–725)
permit limited characterization of the haemal arch anatomy.
One (ISIR 725) (fused by preservation to the right femur, ISIR
58, on the mounted skeleton) is V-shaped with a closed proxi-
mal cross-bar (Text-fig. 7C); the position in the tail for this
chevron is more caudal than the smaller two. The other two
chevrons, ISIR 723 (Text-fig. 7D) and ISIR 724, are Y-shaped
(forked), with fused cranial and caudal projections. In ISIR 723,
the dorsal surface of the haemal canal is open, lacking a proxi-
mal cross-bar.
Pectoral girdle
There are three scapula-coracoids in the collection: left scapula
and coracoid (ISIR 68 and ISIR 69, mounted skeleton) and right
scapulacoracoid (ISIR 92) (Text-fig. 8). The left scapula (ISIR
68) probably represents a smaller individual, where the scapula
and coracoid are disarticulated, but it is fairly complete except
for the cranial and caudal margins. The left coracoid (ISIR 69)
and the right scapulacoracoid (ISIR 92) are more or less in close
size range. In the coracoid, the cranio-ventral part is missing.
The scapular blade is damaged in cranial and caudal margins in
both the specimens. A major part of the scapular blade and the
lower portion of the coracoid of ISIR 92 are missing. Although
the scapula and coracoid of ISIR 69 and ISIR 92 are coalesced,
the general curvature of the bones is well preserved. Jain et al.
(1975, pl. 93A) briefly described the scapula and coracoid.
The scapula (ISIR 68) (Text-figs. 8A–E) on the mounted skel-
eton and isolated element (ISIR 92) is slender, with a modestly
expanded blade showing a gentle dorsoventral convexity on its
lateral surface. The medial surface is slightly concave. An obli-
que, faint ridge originates above the glenoid cavity and fades
away laterally in the middle of the bone. Medially a similar but
prominent ridge is present. The scapular blade is much thick-
ened in the region of these two ridges. From its distal extremity,
the cranial border curves gradually downward and expands to
form a relatively weak acromium, producing a concave profile in
lateral aspect. The caudal border is nearly straight in lateral pro-
file, expanding only slightly where it contributes to the glenoid
facet. The scapula thickens cranially towards the articulation
with the coracoid. The scapula-coracoid articulation is weakly
sigmoid in dorsal aspect. The caudal end is thick and rugose
indicating the presence of suprascapular cartilage in life. Passing
forward the blade gradually narrows in the mid-length and then
flares again cranially. The cranial edge of the blade is very thin,
forming a shallow curvature to meet ventrally with the thickened
acromial outgrowth. The ventral edge is equally thin, narrows
anteriorly and expands in the thickened glenoid region. Laterally,
the fossa for supracoracoscapular muscle is shallow and forms
an arc between the acromium and the glenoid rim.
The glenoid is semi-elliptical in outline; the articular surface is
rugose facing craniodorsally with a central dip. In ISIR 68, the
articular surface for coracoid is rough, pitted and considerably
thickened. In ISIR 69 and ISIR 92, a faint ridge marks the con-
tact between the two elements, where these are coalesced.
The coracoid is subcircular in outline, with a convex lateral
surface and a fairly concave medial surface. The caudal border is
nearly circular. The small coracoid foramen is situated cranial to
the glenoid cavity near the scapulacoracoid articulation. Below
556 P A L A E O N T O L O G Y , V O L U M E 5 3
the foramen, an antero-caudal ridge divides the bone into two
sloping surfaces. The glenoid surface on the coracoid is subtrian-
gular, rugose, facing somewhat laterally and caudally. Its inner
half is slightly hollowed out. The ventro-medial edge of coracoid
shows a smooth and longitudinal groove for probable contact
with the sternal plate. The coracoid thickens caudally to contrib-
ute to the deeply concave glenoid cavity, which it shares with
the scapula, for reception of the humerus.
Forelimb
The forelimb in B. tagorei is shorter than the hind limb; in the
partial association of the small individual, the humerus is 78Æ5per cent of the length of the femur, and the combined length of
humerus + radius is 83Æ1 per cent of that of the femur + tibia.
The proximal segment is distinctly longer than the distal; the
length of the radius is 70 per cent of that of the humerus, and
the length of the tibia is 60Æ5 per cent of that of the femur. A
definitive quadrupedal pose for Barapasaurus has been inferred
from its robust and elongate forelimb (Wilson 2005b).
Humerus. Humeri in the Barapasaurus collection are remarkably
uniform despite pronounced differences in size. There are three
left humeri (ISIR 70; Text-figs. 9A–F), (ISIR 86 and ISIR 87)
and three right (ISIR 85, ISIR 88 and ISIR 93). The humeri ISIR
70 and ISIR 85 represent the smallest groups. These are about
the same size and proportions and possibly belong to the same
individual. ISIR 70 is complete, but the shaft is distorted,
whereas the latter is deficient in the shaft region. ISIR 86 is a
medium sized, least distorted humerus and is the basis for resto-
ration along with ISIR 70. However, its proximal end is partly
damaged. ISIR 87 is the proximal half of a large humerus, the
shaft showing a central cavity. ISIR 88 is fairly complete except
at the two ends. It is much less distorted and is intermediate in
size between ISIR 86 and ISIR 87. ISIR 93 comprises four pieces
which cannot be fitted properly. Its estimated length is compara-
ble with ISIR 87. It is somewhat crushed antero-caudally, and
the two ends bear well-marked rugosities.
The humerus is long, slender and expanded transversely at
either end. The proximal expansion, with the prominent delto-
pectoral crest, is larger than the distal extremity. The two expan-
sions, however, make a slight angle (15 degree) with each other.
The bone is flattened cranio-caudally throughout its length. Its
proximal articular surface is convexly rounded, triangular, with
apex in caudal position; this end is marked by rugosities in the
large humerus. Opposite the apex, on the cranial border of the
articular facet, is a distinctive sulcus. The shaft is narrowest at
its middle (approximately one-third of the proximal expansion)
and expands distally to form slightly separated, convex articular
surfaces for the radius and ulna; it is ovoid in cross-section and
shows a central cavity in ISIR 87.
In lateral aspect, the humerus is sigmoid. Cranially the medial
margin of the humerus is more deeply curved than the lateral
margin. The deltopectoral crest is confined approximately to the
proximal third of the humerus and is situated on the craniolat-
eral margin of the surface. The crest does not form a sharp acute
apex; it is produced into a thick and flat ridge and makes a shal-
low concavity with the adjacent cranial surface, which is other-
wise almost flat. Apparently, this concavity was for the insertion
of the supracoracoscapularis and coracobrchialis muscles. Later-
ally, a prominent ridge runs throughout the length of the bone.
The distal end is slightly inclined cranially and is highly rugose
and flat. The transverse diameter is twice the sagittal diameter.
The radial and ulnar condyles are feebly developed and are sepa-
rated by a longitudinal groove cranially. Caudally the anconeal
fossa is moderate in depth.
Radius and ulna. Among the epipodials are one left radius
(ISIR 71), one right radius (ISIR 89), two left ulnae (ISIR 72
and ISIR 91) and one right ulna (ISIR 90). From the field evi-
dence, it seems that the radius ISIR 71, ulna ISIR 72 and
humerus ISIR 70 belong to the left side of a small individual;
the radius and the ulna were found on either side of the
humerus in association. Similarly, the right epipodials of a lar-
ger individual are represented by the radius (ISIR 89) and ulna
(ISIR 90), which were found side by side in the excavation. A
left ulna (ISIR 91) was found two metres away from these two
bones and is comparable in size (Text-fig. 2, Association G).
However, no compatible humerus was recovered alongside,
although a left humerus (ISIR 87) which seems to be the right
side of the second association was obtained far away from this
spot. ISIR 71 and ISIR 72 are fairly complete, well preserved
and almost undistorted. The restoration of the radius and ulna
are entirely based upon this material. In ISIR 89 and ISIR 90,
the ends are worn and the shafts are deficient; ISIR 91 is better
preserved and is almost complete. Its shaft and distal end show
minor erosion.
The radius (Text-fig. 9M–R) is cylindrical, shorter and slim-
mer than the ulna and its length is 70 per cent of that of the
humerus. The shaft is narrow, long, ovoid in cross-section,
weakly bowed in cranial and caudal aspects, and straight in
medial and lateral aspects. The shaft expands smoothly into an
inflated distal extremity with rugose texture even more pro-
nounced than on the proximal end. Proximally, the radius
expands cranio-laterally to caudo-medially and shows rugosities.
The proximal articular facet is convex and ovoid in profile. Lat-
erally, a sinuous ridge runs lengthwise and terminates distally as
a projection for the ligamentous connection to the ulna. Below
this projection is a flat, triangular facet for contact with the
ulna. The distal facet is also convex, pronouncedly rugose but is
distinctly circular in profile.
The ulna (Text-fig. 9G–L) is much heavier and slightly longer
than the radius. It has an enlarged and robust proximal end, a
narrow shaft, and a slightly expanded distal end. The proximal
end is triradiate with prominent craniolateral and craniomedial
processes and less prominent caudal process. Its rugose triangu-
lar proximal facet has a moderately deep cranial groove formed
by craniomedial and craniolateral processes to receive the proxi-
mal end of the radius. The cross-section of the bone is triangular
for the proximal two-thirds of its length, beyond which the shaft
becomes ovoid in cross-section. The radial groove on the medial
surface is well marked and deeply set. The cranial apex of the
proximal end is bluntly rounded, whereas the caudal one pro-
jects sharply as a thin flange. From it, a sharp ridge runs obli-
quely to the distal end. A corresponding tuberosity is present on
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 557
this ridge, as in the radius, near the distal end, presumably for
the ligament attachment to bind the radius. Below this tuberos-
ity is a facet for reception of the radius. Thus, the two bones
were locked essentially parallel to the each other in life without
any supination. The shaft is straight, and distally the bone is
expanded a little transversely; the distal facet is elongated, rugose
and concave for reception of the carpus.
Manus
Metacarpals. There are three metacarpal bones (Text-fig. 10) in
the collection. These are the third (ISIR 94), fourth (ISIR 95)
and the fifth (ISIR 96) of the left side. Except the long third
metacarpal (ISIR 94), the metacarpals are subequal in length.
They were collected as surface finds and may belong to a single
individual. The third and fifth metacarpals are complete, whereas
the fourth is represented by the proximal half only. In addition,
an isolated element (ISIR 108) is probably the first metacarpal
of left side.
The third metacarpal (ISIR 94) (Text-fig. 10E–H) has
enlarged ends and a narrow shaft, where the greatest diameter
of the proximal end makes a small angle with the transverse
diameter of the distal one. The proximal end is subtriangular
with lateral extension. The medial margin forms the base of
the triangular shape, the lateral margin tapers into an apex, to
overlap the medio-proximal corner of the fourth metacarpal.
From the caudal apex of the triangular end, a blunt but pro-
nounced ridge continues up to the mid-length of the bone.
The summit of the ridge shows rough end surface. Beyond the
ridge, the shaft is narrow and triangular in cross-section. The
distal end is convex, with an asymmetrical trochlear facet, and
is roughly rectangular in outline. On the lateral surface near
the distal extremity, a longitudinal groove indicates the passage
of a flexor tendon.
The proximal end of the fourth metacarpal (ISIR 95) is trian-
gular, convex, and elongated transversely. The ventral margin
forms the broad base of the proximal triangle. The cranial apex
of the triangle is continued as a ridge downward, demarcating
the two separate contact surfaces for Mc III and Mc V on either
side. The caudal surface in the proximal half has a shallow con-
cavity. The proximal end narrows abruptly to narrow shaft. The
distal half is lacking.
The fifth metacarpal (ISIR 96) (Text-fig. 10I–M) is a short
and squat bone, with expanded proximal extremity and a
broad shaft. The flat proximal extremity is subtriangular and
slopes laterally. The shaft is flattened cranio-caudally. Posteri-
orly the shaft is concave. The distal end is ovate and elongated
transversely.
An isolated metacarpal (ISIR 108) (Text-fig. 10A–D), proba-
bly the first metacarpal of the left side, is long and stout with
little expansion at the two ends. The proximal end is roughly
subtriangular and is more expanded than the distal end. Later-
ally a blunt, hook-like projection occurs, below which the shaft
is semicircular in cross-section, robust and broad and main-
tains an almost uniform width. The laterodistal corner is bro-
ken. The distal end is semi-rectangular with a median broad
groove.
Phalanges. There are four ungual phalanges in the Barapasaurus
collection, two from the manus and two from the pes (described
below), ranging in size from large to small. Unguals ISIR 749
(Text-fig. 10N–Q) and ISIR 110 (Text-fig. 10R) are the ungual
phalanges, digit I of the manus of different individuals. The sur-
face texture of the larger of the two manual unguals (ISIR 749)
is extremely rugose and pitted proximally, and smoother distally
where the claw sheath covered the bone in life. The articular sur-
face is irregular rather than smooth, indicating limited move-
ment and weight bearing by this digit. The shaft is tightly
curved, and the bone terminates in a blunt knob-like protuber-
ance. The smaller ungual (ISIR 110) belonged to a young indi-
vidual. It is similar to the larger ungual, but less heavily rugose
and pitted. The proximal end is oval, recurved, blunt and con-
cave with a median ridge and two symmetrical depressions
beneath the proximal margin of the sheath. The articular surface
is concave, smooth and bears a prominent rim of constriction
just below this surface. The cranial surface is pitted, indicating a
horny sheath of cover during life. The lateral and medial sur-
faces have a groove extending from the middle to the tip of the
claw. The distal end is bluntly rounded. Barapasaurus might
have a tubular manus as suggested by Upchurch et al. (2004).
Although the collected bones of Barapasaurus lack a complete
preserved manus, Wilson (2005b) inferred reduction of manual
phalanges in this taxon.
Pelvic girdle
The collection includes five ilia, five ischia and five pubes, of
which two distinct size groups and associations can be sepa-
rated. A right ilium (ISIR 52), one left (ISIR 54) and one right
(ISIR 115) ischia, one left (ISIR 117) and one right (ISIR 57)
pubes belong to a small individual (Text-fig. 2, Association C).
One left (ISIR 111) and one right (ISIR 51) ilia, one left (ISIR
114) and one right (ISIR 53) ischia and one left (ISIR 56) and
one right (ISIR 55) pubes belong to a large individual (Text-
fig. 2, Association A). The sacrum (ISIR 50) was found along
with the pelvis of the associated large individual; however, two
ischia were slightly away from the locality. In addition to these
partial associations, a few isolated pelvic bones were also col-
lected: these include one right (ISIR 112) and one left ilia
(ISIR 113), one left ischium (ISIR 116) and one left pubis
(ISIR 118) (Text-fig. 2, Association F). All the pelvic bones
occurred as disarticulated elements without any co-ossification
between them.
Ilium. The ilium was described and illustrated by Jain et al.
(1975, figs 3–4; Jain et al. (1979), pls 93–94) on the basis of ISIR
51 and ISIR 52. In the mounted skeleton, the ilia are ISIR 51
and ISIR 111. These ilia are generally deficient in the blade in
the caudo-dorsal region, whereas the cranial process of the blade
is well developed. Except ISIR 112, the ilia are not very crushed
and are in a good state of preservation. The iliac profile (Text-
fig. 11A–B) in lateral aspect is rounded dorsally with a gentle
convexity. The curved and subtriangular iliac crest extends crani-
ally to form the prominent preacetabular process. The pubic
peduncle is likewise prominent and extends downward and
558 P A L A E O N T O L O G Y , V O L U M E 5 3
slightly cranially. The ischiadic peduncle is short and subrectan-
gular in lateral view. The acetabulum is deeply concave, and the
bone is thickened medially. The sacricostal yoke does not con-
tribute to the deep medial wall of the acetabulum. As mounted,
the dorsal crests of the ilia are narrow and converge cranially.
The ilia are firmly ankylosed to the sacral ribs (Text-fig. 5E).
Pubis. ISIR 57 (Text-fig. 12E–H) is the best-preserved pubis,
which retains the twist below the proximal and distal regions. In
other specimens, there is a variation of the amount of the twist
because of different degree of crushing. All the specimens are
deficient in the caudal margin, in the region between the two
symphyses. Jain et al. (1975, 1979) described and discussed the
implications of the unusual anatomy of the pubis based on ISIR
55, ISIR 56 and ISIR 57; to this list ISIR 117 may be added. They
suggested that the pubis is somewhat shorter than this ischium
and illustrated these two bones in profile with considerable dif-
ference in length. However, the pubis (ISIR 117) and ischium
(ISIR 54) in the associated skeleton ‘Colbert excavation’ are
almost exactly the same length. The pubic apron is developed
but without substantial thickening and without obscuring the
basic anatomy of the shaft. The proximal articulations with the
ilium and ischium are borne by the stout iliac and ischiadic
peduncles, respectively. The proximal part of the pubis is pierced
by large obturator foramen, which is situated well within the
pubic apron. The pubic apron articulated with its partner along
nearly the full length of the midline. The resulting pelvic basin,
which is the opening between the pubes beneath the sacrum, was
consequently narrow, as recognized by Jain et al. (1979). The
shaft of the pubis is straight in anterolateral profile and expands
distally to form a greatly enlarged distal extremity to meet its
partner at the midline. The pubic shaft in posterolateral aspect is
steeply curved. The distal extremities of the pubes articulate
along the midline, forming a cranioventrally sloping trough.
Ischium. Jain et al. (1975) described and illustrated the ischium
of B. tagorei on the basis of two specimens, ISIR 53 and ISIR 54
(Text-fig. 12A–D). The curvature of the ischiadic shaft is pre-
served only in ISIR 54, which is fairly complete. The relatively
delicate cranial flange was invariably damaged in all specimens.
The shaft of the ischium is slender and straight, transversely
compressed and longer than the pubis shaft. Its distal extremity
is moderately expanded, subrounded, and the symphyseal contact
is narrow. In articulation with the other bones of the pelvis, it is
oriented downward and rearward, at roughly 45 degree angle
with respect to the horizontal axis of the vertebral column. It is
almost exactly the same length as the pubis (see above), rather
than longer as originally published. Its articulation with the ilium
is broad and robust where it contacts the iliac peduncle. Its pubic
peduncle is deep and equally robust. The ischiadic contribution
to the acetabulum is roughly equal to that of the pubis.
Hind limb
The Barapasaurus mounted skeleton includes the femora, tibiae,
right fibula, right astragalus, three left metatarsals and two
ungual phalanges. In addition, rear limb bones in the associated
skeleton include both femora, left tibia and both fibulae.
Femur. There are seven well-preserved femora in the Barapasau-
rus collection; these include three right femora ISIR 58, ISIR 99
and ISIR 100; and four left femora, ISIR 59, ISIR 60, ISIR 97
and ISIR 98. The femora are fairly complete except ISIR 100 in
which only the proximal half is retained. The lengths of the
femora range from 875 to 1365 mm. The right femur on the
mounted skeleton (ISIR 58) (Text-fig. 13E–F) has a length of
1167 mm. It appears that ISIR 58 and ISIR 59 represent a pair
of femora of a juvenile individual. Most of the femora are com-
pressed craniocaudally from burial; consequently, the shafts are
flattened, and the condyles are also partly damaged.
The femur is long and slender with expanded ends and a
straight shaft. The prominent, hemispherical head is set at a
right angle to the straight, slender shaft. The proximal end is
rugose and ovate, with two distinct levels. The higher one is
more rugose with deep furrows forming the proper head, while
lower part is less rugose and grades laterally into the greater tro-
chanter. Laterally, a shallow depression occurs just below the
greater trochanter. The shaft has a fairly uniform width, ovoid
in cross-section, with a central cavity (ISIR 100). There is no les-
ser trochanter, but the fourth trochanter is prominent and forms
a distinctive ridge (Text-fig. 13B). The fourth trochanter is situ-
ated on the caudo-medial margin, and its apex is slightly higher
than the midlength. It projects caudally as a ridge-like process
with an acuminate and declined tip. The medial surface of the
fourth trochanter shows a rough surface.
Distally the robust tibial and fibular condyles are well devel-
oped and pronounced. The medial condyle is larger than the
lateral condyle. The two condyles are separated by the inter-
condylar groove. The lateral condyle is further subdivided into
an internal condyle and a lateral epicondyle by a lateral longi-
tudinal grove (Janensch 1961) which might have provided the
passage of a strong tendon. The medial and the internal condyle
together receive the proximal end of the tibia while the lateral
condyle articulates with the fibula.
Some morphological variations as a result of growth have been
noted on the femora of B. tagorei. On the basis of the greatest
length of the femora, three distinct size groups can be recognized;
these are small (ISIR 59 = 875 mm and ISIR 60 = 885 mm),
medium (ISIR 99 = 101 mm) and large (ISIR 58 = 122 mm;
ISIR 97 = 1365 mm; ISIR 98 = 132 mm and ISIR 100 =
134 mm (estimated). The ends of the small femora do not have
any rugosities, whereas the larger femora are marked by pro-
nounced rugosity, and the medium sized femora show only some
pitting. The head of the large femora is distinct, whereas in the
medium and small femora the head cannot be demarcated from
the rest of the proximal end.
The anatomy of the fourth trochanter resembles that
described by Yadagiri (2001) for Kotasaurus, but is not suffi-
ciently preserved to permit confident comparison.
Tibia and fibula. The epipodials include four left tibiae (ISIR 61,
ISIR 62, ISIR 63 and ISIR 102), three right tibiae (ISIR 77, ISIR
101 and ISIR 103), two left fibulae (ISIR 64 and ISIR 105) and
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 559
two right fibulae (ISIR 104 and ISIR 106). A partial association
of a hind limb was recognized when the tibia, ISIR 62 was found
along the side of the fibula, ISIR 64; these two were recovered
near the femur, ISIR 60 (Text-fig. 2, Association C). The right
tibia, ISIR 101 occurred nearby the right femur, ISIR 58 (Text-
fig. 2, Association H). The size of ISIR 101 and ISIR 102 indi-
cate a probable pair and their corresponding fibulae may be ISIR
105 and ISIR 106. The tibia, ISIR 103 and the fibula ISIR 104
may belong to the same side of a large individual. The tibia ISIR
62 is least damaged and retains the twist of the two ends.
Although the tibia, ISIR 63 is represented by only the distal end,
it possesses the notch for the astragalus. The fibulae ISIR 64 and
ISIR 104 are fairly complete.
Jain et al. (1975) described the tibia of B. tagorei. The tibia
length (ISIR 62 = 505 mm) is short relative to the length of the
femora (865 mm and 867 mm) in the associated skeleton. The
lengths of the right tibia (ISIR 103) and right femur (ISIR 58) in
the mount are 714 and 1167 mm, respectively, approximately
the same ratio (61 per cent) as in the associated skeleton (58 per
cent). The tibia (Text-fig. 14A–E, L) is robust, with well-devel-
oped cnemial crest. The proximal articular surface is rugose,
ovoid and interrupted by a notch. The cnemial crest is a thin
vertically elongated flange protruding craniolaterally for the
attachment of triceps femoris muscle. It extends up to one-
fourth of the tibia length and is separated from the rest of the
proximal end by a prominent notch in the caudolateral aspect.
The remainder of the proximal end is differentiated into two
subequal medial and lateral expansions caudally for the articula-
tion of the femur. The lateral expansion and the cnemial crest
converge downward forming a triangular surface for the contact
with the fibula but with a notch between them. In cranial pro-
file, the shaft is broad and straight. The shaft is semi-circular in
cross-section and expands slightly towards the distal extremity.
The distal articular facet is distinctly concave, triangular in out-
line and lacks separation between the caudoventral process and
the articular surface for the ascending process. Distally the tibia
articulates with the astragalus by a notch and a descending
flange. The notch is located on the caudolateral corner for the
reception of the ascending process of the astragalus. The
descending flange on the caudomedial surface of the tibia pro-
jects ventrally to fit into the astragalus. A faint groove between
the notch and the descending flange marks the passage of a ten-
don for the flexor muscle of the foot. Rugose marks occur along
the periphery of the distal end.
The fibula (ISIR 64, Text-fig. 14F–K) is slender and trans-
versely compressed, expanded proximally and distally, and nar-
rowest at the centre of the shaft. The weakly crescentic proximal
end adjoins the crista lateralis of the tibia, and together these
surfaces articulate with the lateral condyle of the femur. In lat-
eral view, the fibular shaft is narrow and straight from its proxi-
mal extremity to near the distal extremity, where it expands
somewhat to form the carpal facet and becomes sigmoid in pro-
file. The lateral trochanter is weakly developed depicting the ori-
gin of the flexor digitorium longus muscle (Borsuk-Bialynicka
1977; Wilson and Sereno 1998), and the tibial articular surface
is a narrow ridge parallel to the shaft proximally, but rather
oblique near its distal extremity. A raised elliptical rugosity
occurs on the lateral side of the mid-shaft of the fibula for the
articulation of the tibia (Wilson and Sereno 1998); the muscle
scars on the fibula are considered as a derived condition of
Barapasaurus (Wilson and Sereno 1998). The articular facets are
convex. The distal end is elongated and subrounded, highly
rugose and fits to the lateral surface of the astragalus.
Tarsus. The tarsus is represented by a right astragalus (ISIR 107)
which was found in association with tibia ISIR 103 and a right
calcaneum (ISIR 743). There are three metatarsals of the right
side in the collection; these include the metatarsal I (ISIR 65),
metatarsal II (ISIR 66) and metatarsal IV (ISIR 67). These three
were found associated with the astragalus (ISIR 107), tibia (ISIR
103) and fibula (ISIR 104). Metatarsal I and IV are complete
and undistorted; metatarsal II is damaged in the shaft and proxi-
mal region. The ends of the metatarsals are rugose.
The right astragalus (Text-figs 14L, 15A–C) is flat and subtri-
angular. The proximal surface bears a prominent ascending pro-
cess and a medial depression; the ascending process is convex
and articulates with the distal extremity of the tibia. Medially,
the base of the ascending process slopes downward gradually
into the depression. Its convex distal surface is uniformly con-
toured, broad and rugose. The cranial margin is straight and
rugose, with a gently sloping surface for the articulation of the
metatarsals. Separate facets for individual metatarsals are not
discernible. The calcaneal articular surface is indistinct. The
caudal surface is less rugose and exhibits an upturned lip which
superficially covers the tibia. The medial margin is rugose and
tapers cranially. The lateral surface is quite deep; its upper half
is quite smooth, probably for articulation with the calcaneum,
and the lower half is rugose. Other anatomical features of this
element cannot be discerned owing to its position in the
mounted skeleton.
The calcaneum (ISIR 743) (Text-fig. 15D–G) is roughly qua-
drangular. Cranially, it has a convex outline. The proximal artic-
ular surface is semi-trapezoidal, slightly concave and smooth for
reception of the fibula. The medial surface, which articulates
with the astragalus, is gently convex and smooth. Laterally, the
calcaneum is a little thickened but smooth and gently rounded.
The caudal surface is flat. The distal articular surface for the
metatarsals is convex, rough and has a fine mediolateral ridge
on the caudodistal margin, which might have attached the fleshy
pad (Bonnan 2000). Wilson (2005b) concurred that the pes of
Barapasaurus was supported by heel pad. Bonnan mistakenly
mentioned the presence of calcaneum in Barapasaurus (2000,
2005; Bonnan, pers. comm. 2007). It had not been reported ear-
lier. The calcaneum in Text-fig. 15D–G would fit into the space
between the distal extremity of the fibula and lateral metatarsals
in the mounted skeleton (Text-fig. 16A–C).
Pes
Metatarsals. Right metatarsals I, II, and IV (Text-figs. 15A–C,
16A–D) are elongate, with proximodistal lengths of 178, 259
and 250 mm. Metatarsal I (ISIR 65) is the shortest, but heavi-
est of the three, and its shaft is twisted. The twist of the proxi-
mal expansion is 55 degrees with respect to the distal one. Its
proximal articular surface is more greatly expanded than the
560 P A L A E O N T O L O G Y , V O L U M E 5 3
distal end. The proximal end is elongated and ovoid, with its
apex pointing laterally for overlapping the adjacent metatarsal
II. The shaft is broad and compressed cranio-caudally; its lat-
eral margin forms a sharp ridge terminating distally into a
notch. The medial margin is flat and broad. The distal extrem-
ity forms an asymmetric ginglymus where the lateral condyle is
larger than the medial one.
Metatarsal II (ISIR 66) is longer and slimmer than the first,
but about the same length as metatarsal IV. The proximal end
has a biconcave outline and is more expanded than the distal
end. Medially, the proximal surface is produced into a concave
flange for the reception of metatarsal I. The shaft is considerably
more robust, flattened, oval in cross-section and shows a similar
ridge and a notch on the lateral margin as in the metatarsal I.
The distal end has a symmetric ginglymus surface with a deep
median groove.
Metatarsal IV (ISIR 67) is long and slender. The proximal end
is narrow, elongated and subtriangular with a sharp apex point-
ing medially. The shaft has ridges both laterally and medially
and similar notches on the distolateral margin as in metatarsal I
and II. The distal end is squared and anterocaudally thickened;
no distinct grove is visible on the ginglymus surface.
In life, the pes was semiplantigrade as indicated by the pres-
ence of weakly developed lateral trochanter on the fibula and the
long metatarsus. The twisted metatarsal I and robust condition
of metatarsals I and II indicate that much of the weight borne
by the pes was on the inner side of the toes. Wilson (2005b)
suggested that the pedal unguals of Barapasaurus were deflected
laterally relative to the long axis of each digit and of the foot
itself. He considered this feature as a derived character of Bara-
pasaurus.
Phalanges. As mentioned in the description of the manus, there
are four dissociated ungual phalanges, two of which are pedal.
ISIR 83 and ISIR 84 (Text-fig. 16D–L) belong to digit I and digit
III of the right pes. These two ungual phalanges are similar in
general build, recurved, somewhat blunt and pitted on the sur-
face. The proximal end is oval, concave with a median ridge
with two symmetric depressions on its side. The articular surface
is extensive up to the cranial tip, allowing considerable flexion
and extension. The cranial margin is longer that the caudal one.
PHYLOGENETIC PERSPECTIVES
Upchurch et al. (2007a) in their phylogenetic analysis
diagnosed 292 characters for 34 taxa of basal saur-
opodomorphs and on the basis of published information
(e.g. Jain et al. 1975, 1979; Wilson and Sereno 1998)
compared Barapasaurus which scores 99 characters in
their data matrix (Upchurch et al. 2007a). It may be
mentioned that because of lack of skull, braincase and
mandible, the first 85 characters could not compared.
Hence, 33Æ9 per cent character scoring out of 292 charac-
ters was actually used. Yates (2007a) carried out another
phylogenetic analysis of the basal sauropodomorphs and
identified 351 characters from which Barapasaurus shares
96 characters and the first 106 characters from his data
matrix belonged to skull, braincase and mandible. Hence,
he used 27Æ53 per cent of the characters for Barapasaurus.
Later Yates (2007b) added more characters in his previous
phylogenetic analysis. Subsequently, Smith and Pol (2007)
described a basal sauropodomorph from Antarctica and
carried out cladistic analysis and added eight more char-
acters to the character scoring of Yates’s (2007a, b) phylo-
genetic analyses; but according to them, Barapasaurus
does not share any of those characters.
The present description of Barapasaurus improves the
character scoring of Yates (2007a, b), Smith and Pol
(2007) and Upchurch et al. (2007a). In the present work,
the data matrix comprising 34 taxa and 292 morphologi-
cal characters that was initially used by Upchurch et al.
(2007a) is modified (Table 4), especially those character
states that are related to Barapasaurus. However, several
of their characters, especially those with ratios could not
be used because of lack of associated ⁄ articulated bones.
The present description improves the scoring of Barapa-
saurus with a total of 143 characters out of 292 and 48Æ97
per cent of character scoring can be used. It may be
mentioned here that most of the character states for
TABLE 4 . Revised data matrix of Barapasaurus tagorei Upchurch et al. (2007a) that was used for the current phylogenetic analysis.
81 91 101 111
? ? ? ? ? 1 ? 1 1 1 1 1 ? ? ? 1 ? 1 1 ? ? ? ? ? 0 1 0 ? 0 ? ? 1 0 1 ? 1 1 0 1 1
121 131 141 151
0 0 0 1 0 0 1 0 0 1 1 1 1 1 0 ? ? 1 1 0 0 1 0 1 1 1 0 0 1 0 1 1 ? ? 0 0 1 0 ? 1
161 171 181 191
? ? 0 0 1 0 1 1 1 0 1 0 1 1 ? 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? 1 0 ? ? ? ? ? ? ? ?
201 211 221 231
? ? 0 1 1 1 0 1 1 0 0 1 1 1 0 1 0 0 0 1 1 0 0 1 1 1 0 1 1 1 0 0 1 0 ? ? 1 10 1
241 251 261 271
1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 ? ? ? ? 0 1 1 1 ? 1 0 ? 1 ? 1 0
281 291
? ? ? ? ? ? 0 ? ? ? ? 1
For the character details, see Upchurch et al. (2007a).
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 561
Vulcanodon are either missing ⁄ unknown (characters rang-
ing from 1–138 and 179–208). The data matrices of Yates
(2007a) and Smith and Pol (2007) deal mostly with pros-
auropods and were not used here.
The modified data matrix is subjected to a maximum
parsimony analysis using PAUP 4.0b10 (Swofford 2000).
Multistate characters were treated as unordered, and all
characters were weighted equally. Unknown ⁄ missing char-
acters were coded as ‘?’. The heuristic search algorithm
was used with default settings. The analysis recovered 47
most parsimonious trees (MPTs), each with tree lengths
(TL) of 738 steps, a consistency index (CI) of 0Æ40
rescaled to 0Æ26, a retention index (RI) of 0Æ65, and a
homoplasy index of 0Æ6. The topology of the strict con-
sensus tree is shown in Text-figure 17, which is generally
compatible with that of Upchurch et al. (2007a).
Although Vulcanodon and Kotasaurus were considered as
basal in comparison with Barapasaurus (Upchurch et al.
2007a), in the current work Barapasaurus was found to
be more basal than Vulcanodon and is further removed
from the Eusauropoda (sensu Upchurch et al. 2007a). A
bootstrap analysis (1000 replicates) shows moderate to
strong support (>60 per cent) for the fully resolved clades
within the strict consensus tree. In addition, topological
constraints were created to explore two separate hypothe-
ses. These comprise: (1) Kotasaurus and Barapasaurus are
sister taxa; and (2) Barapasaurus falls within Eusauro-
poda. In the current analysis, it was found that 21 MPTs
(TL = 748) were generated when the first constraint was
enforced showing that another 10 steps are required for
Kotasaurus and Barapasaurus to be sister taxa. On the
other hand, the second constraint resulted in the genera-
tion of 675 MPTs (TL = 741), showing that three more
steps are necessary to incorporate Barapasaurus within
Eusauropoda. In both the cases when the constraints were
applied, the resulting topologies show that the Sauropoda
constitutes a monophyletic clade, the Eusauropoda with
Camarasaurus and Omeisaurus remaining as sister taxa,
similar to that seen in the unconstrained MPTs.
DISCUSSION
Previous work on classification of Barapasaurus
In 1975, Jain et al. described B. tagorei under the Infraor-
der Sauropoda but did not assign it to a family. They
drew attention to the similarity of some aspects of the
anatomy of B. tagorei to that of the Prosauropoda, espe-
cially with respect to the sacrum. Gauthier (1986) gave an
outline of the sauropod relationship where he considered
Barapasaurus as one of the most primitive sauropods
along with Vulcanodon and listed at least 20 synapomor-
phies uniting them with other sauropod taxa. Subsequent
workers (Bonaparte 1986; Mcintosh 1990; Wilson and
Sereno 1998; Upchurch et al. 2004, 2007a) also kept
Barapasaurus within Sauropoda. Wilson and Sereno
(1998) defined Sauropoda as all sauropodomorphs more
closely related to Saltasaurus than to Plateosaurus and
proposed 17 synapomorphies for Sauropoda among which
Barapasaurus shares (1) quadrupedal posture with colum-
nar limbs and short metapodial; (2) four sacral vertebrae;
(3) low deltopectoral crest of humerus; (4) absence of
TEXT -F IG . 17 . Strict consensus tree of the 47 MPTs found by
the heuristic search in PAUP 4Æ0b10 (Swofford 2000). Tree
statistics are as follows: TL = 738 steps; CI = 0.40; RCI = 0.26;
HI = 0.60; RI = 0.65. A, Sauropoda. B, Eusauropda. C,
synapomorphies are as follows: elliptical ⁄ subrectangular
transverse section of the femoral shaft, angle between femoral
head and transverse axis of the distal end is close to 0 degree,
tibia smaller than femur, and the presence of extensor
depression on the femoral distal end.
562 P A L A E O N T O L O G Y , V O L U M E 5 3
olecranon process; (5) triradiating proximal end of ulna
with deep radial fossa; (6) ilium with low and subrectangu-
lar ischial peduncle; and (7) ischial shaft equal to pubic
shaft. The other proposed synapomorphies are difficult to
ascertain because of non-availability of associated bones.
Upchurch et al. (2004) defined Sauropoda as a stem-based
taxon with 15 characteristic features some of which were
earlier suggested by Wilson and Sereno (1998); Barapa-
saurus shares (1) triradiate proximal end of ulna; (2)
middle and distal portions of the femoral shaft straight in
cranial view. Upchurch et al. (2007a) produced a detailed
phylogenetic analysis on basal sauropodomorph relation-
ships and maintained Sauropoda as a stem-based taxon
which includes Eusauropoda, Barapasaurus, Vulcanodon,
Kotasaurus, Chinshakiangosaurus, Antetonitrus, Blikana-
saurus, ‘melanorosaurids’ and Jingshanosaurus.
As has been mentioned earlier, evolutionary relation-
ships of sauropods by using cladistic analysis have been
considered by several workers, namely Russell and Zheng
(1994), Calvo and Salgado (1995), Salgado et al. (1997),
Upchurch (1995, 1998), Sereno (1997, 1998), Wilson and
Sereno (1998), Wilson (2002, 2005a), Yates (2003, 2007a,
b), Upchurch et al. (2004) and Smith and Pol (2007).
These analyses showed that Barapasaurus was a basal
sauropod belonging to Eusauropoda. Upchurch (1995)
first defined Eusauropoda as a new taxonomic group con-
taining all sauropods except the vulcanodontids and diag-
nosed 14 characters out of which Barapasaurus shares (1)
slightly procumbent tooth crowns; (2) 12 cervical verte-
brae; (3) shorter and robust pubis; (4) less pronounced
torsion between the distal and proximal ends of the
pubis; and (5) very short and robust Mt I. Upchurch
(1995) separated another clade Euhelopidae and diag-
nosed five characters from which Barapasaurs possesses
only two: (1) forked chevrons which may belong to mid-
dle caudals; and (2) the first caudal rib co-ossified with
the vertebrae and is quite expanded to give a fan-shape in
anterior view.
Wilson and Sereno (1998) defined Eusauropoda as a
stem-based taxon and included all sauropods more closely
related to Saltasaurus than to Vulcanodon and diagnosed
40 characters. Because of lack of skull ⁄ skull elements, out
of 40 characters the first 13 cranial characters are not
known in Barapasaurus, and from the rest, Barapasaurus
shares (1) spatulate crown; (2) tooth enamel with wrin-
kled texture; (3) opisthocoelous cervical centra; (4) dorsal
neural spines broader transversely than craniocaudally;
(5) fork-shaped distal chevron; (6) fibular lateral trochan-
ter present; (7) minimum shaft width of metatarsal I
greater than that of metatarsal II–IV; and (8) pedal pha-
langes broader. Wilson and Sereno (1998) proposed
another clade ‘Barapasaurus + Omeisaurus + Neosauro-
poda (sensu Upchurch 1995)’ and diagnosed seven char-
acters of which Barapasaurus has 6 characters, including
(1) neural arches of caudal cervical vertebrae and cranial
dorsal vertebrae with interprezygapophyseal lamina; (2)
opisthocoelous cranial dorsal centra; (3) neural arches of
the middle and caudal dorsal vertebrae with composite
lateral lamina; (4) presence of sacricostal yoke; (5) fibula
with broad triangular scar for tibia; and (6) astragalar
posterior fossa divided by crest.
According to Upchurch et al. (2004), Eusauropoda is a
node-based taxon which defines the most recent common
ancestor of Shunosaurus and Saltasaurus and all the
descendants of that ancestor. They diagnosed 38 characters
including some of Wilson and Sereno’s suggested character
states. The additional common characters of Barapasaurus
includes (1) caudal margins of caudal cervical neural
spines sloping strongly forward in lateral view; (2) dorsal
surfaces of sacral plates level with the dorsal margin of the
ilium; (3) dorsal margin of the ilium gently convex in lat-
eral view; (4) reduced ischial peduncle of ilium so that the
long axis of the iliac blade slopes craniodorsally in lateral
view; and (5) femoral cranial trochanter absent. Wilson
(2005a) considered the distinctive spatulate shape and
crowns of individual tooth with wrinkled enamel texture
as characteristic features for Eusauropoda. Barapasaurus
teeth were mildly spatulate and had weakly crenulated
texture on the enamel, indicating its affinity towards
Eusauropoda. Wedel et al. (2000) and Wedel (2003) also
considered Barapasaurus as ‘Eusauropoda’ on the basis of
presacral vertebrae which bear lateral pneumatic fossae on
the centra, some of which are deeper than others but do
not meet each other closely to produce a medial septum.
Upchurch et al. (2007a) carried out another phyloge-
netic analysis of basal Sauropoda and maintained their
earlier definition of Eusauropoda (sensu Upchurch et al.
2004); their analysis removed Barapasaurus from Eusauro-
poda. According to their phylogenetic analysis, Sauropoda
includes the two Indian sauropods Barapasaurus and
Kotasaurus as well as Vulcanodon, Chinshakiangosaurus,
Antetonitrus, Blikanasaurus, ‘melanorosaurids’ and Jings-
hanosaurus and Eusauropoda. Since Eusauropoda (sensu
Upchurch et al. 2007a) is mainly defined by cranial char-
acters, whether Barapasaurus may be included within Eus-
auropoda or not cannot be assessed because of missing
cranial characters. On the other hand, Barapasaurus along
with Eusauropoda shares the synapomorphies such as
opisthocoelus cervical centra, greater height of the mid-
cervical neural arches in comparison with the centrum
height, presence of spinodiapophyseal lamina on middle
and caudal dorsal vertebrae, presence of postzygapophyse-
al lamina on all dorsal vertebrae, forked middle and distal
chevrons, strongly convex dorsal iliac margin, middle and
distal portions of the pubis lying in the same plane as the
proximal end, absence of lesser trochanter and later-
ally directed cnemial crest (Node U of Upchurch et al.
2007a). The suggestion by Upchurch et al. (2007a) that
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 563
Barapasaurus was a basal sauropod lying outside the Eus-
auropoda is consistent with other phylogenetic analyses of
Benton et al. (2000), Yates and Kitching (2003), and Gal-
ton and Upchurch (2004). However, Allain and Aquesbi
(2008) included Barapasaurus within the Eusauropoda.
Present work
The current phylogenetic analysis based on the revised
data matrix of Upchurch et al. (2007a) shows that the
Sauropoda including the Indian forms Barapasaurus and
Kotasaurus along with Antetonitrus, Camarasaurus, Chin-
shakiangosaurus, Omeisaurus, Jingshanosaurus, Shunosaurus
and Vulcanodon is fully resolved (Text-fig. 17) with the
nodes achieving values of more than 60 per cent. These
taxa share the characters such as the elliptical or subrect-
angular transverse section of the femoral shaft, angle
between the femoral head and transverse axis of the distal
end is close to 0 degree, tibia smaller than femur (charac-
ters 249–250 of Upchurch et al. 2007a, subsequently
quoted here as ‘C249’ etc.) and presence of extensor
depression on the distal end of the femur (C252). Barapa-
saurus is further removed from Eusauropoda and is found
to be more basal in comparison with Vulcanodon (Text-
fig 17) based on the different parameters of the mid-
caudal centra (C145, length of the mid caudal vertebrae
compared to the height of cranial articulation is less than
2 in Kotasaurus and Vulcanodon but >2 in Barapasaurus),
presence of caudal hyposphenal ridge, length of the base of
the caudal neural spines (C147–148), cranial disappearance
of caudal ribs (C150) and subtriangular outline of the
distal end of the ischium (C234). Barapasaurus shows
advanced traits in comparison with Kotasaurus in several
characters. These include opisthocoelus cervical centra
(C106), presence of spinodiapophyseal lamina on middle
and caudal dorsal vertebrae (C124) and composite lateral
laminae on dorsal neural spines (C130), shape of the scap-
ular blade (C157), extent of the deltopectoral crest (C165),
different parameters of the ilium (C207–C209), position
and profile of the fourth trochanter (C241–C242), ratio
between the tibial and femoral length (C251), laterally
directed cnemial crest (C253) and presence of muscle scar
on the lateral surface of the fibula. In contrast, Kotasaurus
shows advanced traits based on mid-caudal centra (C145),
caudal neural spines (C148), ratio between the radial and
humeral length (C176) and metatarsal length: width ratio
(C269).
Some comments on Barapasaurus and Kotasaurus
Until recently, only two sauropods had been known from
the lower part of the Kota Formation of the Pranhita-
Godavari basin of India. Apart from B. tagorei another
species of basal sauropod, K. yamanpalliensis was col-
lected by Yadagiri (1988, 2001) from the same strati-
graphic horizon but from a different locality. The fossil
locality of Kotasaurus is situated 5 km west of the village
Yamanpalli which is 40 km north of Pochampalli village,
the type fossil locality of Barapasaurus (Text-Fig. 1). Mul-
tiple taxa of sauropods in the same level of a stratigraphic
horizon are known to occur in different parts of the
world. Kotasaurus and Barapasaurus occur in the same
stratigraphic level of the Early Jurassic Lower Kota For-
mation of Pranhita-Godavari basin. Although only two
sauropods are now known from this horizon, more
sauropod material will likely to be described in future
from the Kota Formation.
Recently, questions have been raised on the validity of
the genus Kotasaurus yamanpalliensis. Yates (2007a) men-
tioned in his article that Kotasaurus is a chimera (a view
held by Oliver Rauhut, p. 30 in Yates 2007a) and he did
not use its characters in his phylogenetic analysis. Allain
and Aquesbi (2008) also refrained from using the charac-
ter states of Kotasaurus as the assigned taxon might
belong to more than one species (a view held by Jeff Wil-
son, p. 401 in Allain and Aquesbi 2008). Rauhut recently
opined (Rauhut and Lopez-Arbarello 2008, p. 561) that at
least one more taxon of sauropod is included in Kotasau-
rus. Three authors (SB, SR and DPS) of the present article
examined the mounted skeleton and other isolated mate-
rial of Kotasaurus. It must be mentioned here that there
are a few bones in both the mounted skeleton and the
isolated ones, which bear some similarities with Barapa-
saurus and may belong to Barapasaurus. However, there
are also major osteological dissimilarities between Kota-
saurus and Barapasaurus which separate these two taxa.
Some of the differences were mentioned by Yadagiri
(2001). The vertebral morphology of the presacral series
of the two taxa is quite different. Barapasaurus is charac-
terized by the presence of acamerate vertebrae morphol-
ogy, 16 vertebral laminae and hollow neural spines in the
presacral series, whereas vertebrae of Kotasauras presacral
series have ‘lateral depressions, which may be either deep
and small, or shallow and large’ and 7 vertebral laminae
(4 in cervical and 3 in dorsal vertebrae) (Yadagiri 2001,
p. 246). Gigantism and elongation of neck are the major
biomechanical problems of sauropods which were mostly
resolved by complex vertebral pneumaticity, origination
of vertebral laminae, high and hollow dorsal neural spines
which lighten as well as increase the strength of the verte-
bral column. Although Barapasaurus does not have exten-
sively subdivided internal structures in the centra of
advanced sauropods such as Mamenchisaurus, diplodocids
and brachiosaurids, the presence of pneumatic fossae, 16
vertebral laminae and hollow neural spines in Barapasau-
rus indicates the beginning of increasing body size and
564 P A L A E O N T O L O G Y , V O L U M E 5 3
length of neck in sauropod evolutionary history, and it
may be said that Barapasaurus marks the beginning of
gigantic sauropods. The neural canal of the caudal mid-
dorsals in Barapasaurus is a specialized feature; it becomes
narrow and deeply sunk on the centrum ventrally but
opens dorsally into a large cavity through a narrow slit-
like opening. The base of the neural spine forms the roof
of the cavity and the floor of the cavity is depressed on
either side of the slit-like opening. In contrast the neural
canal in Kotasaurus is a normal tubular form running
almost for the whole length of the arch. Moreover, the
transverse processes of Barapasaurus are mostly directed
laterally while in Kotasaurus these are directed upward.
The second major difference between the two taxa is
the sacrum. Four co-ossified sacral vertebrae have been
recognized in Barapasaurus since its description (Jain
et al. 1975). However, the presence of four co-ossified
sacral vertebrae is questionable in Kotasaurus. Yadagiri
(1988, p. 103) mentioned the presence of three co-ossified
sacral vertebrae but in the same paper he wrote that there
are two conjoined sacral vertebrae, while the third one is
broken (p. 110). Later, Yadagiri (2001, p. 242) mentioned
the presence of ‘19 sacrals (one with three fused centra,
two with fused centra)’ but none with four fused centra.
In the illustration of sacral vertebrae, there are four con-
joined sacral vertebrae (Yadagiri 2001, p. 245). He further
wrote, ‘The full restoration of the sacrum was also based
on features of the incomplete specimens. The co-ossified
sacrum consists of four centra’ (Yadagiri 2001, p. 246). In
this publication, the length of the sacrum is also mea-
sured on the basis of the fourth sacral vertebra. Three of
the present authors (SB, SR and DPS) checked the
sacrum in the mounted skeleton of Kotasaurus and noted
that there are actually three co-ossified sacral vertebrae,
while a fourth loose sacral vertebra is attached to it.
Besides the mounted skeleton, there is no other sacrum
with four co-ossified centra in the collection. It appears
that Kotasaurus probably had three co-ossified sacral ver-
tebrae and a loose dorsosacral for strengthening of the
sacrum, whereas a dorsal vertebra was added to the
sacrum of Barapasaurus.
Among other differences, the Kotasaurus scapula is tall
but significantly narrower than Barapasaurus whose scapu-
lar blade shows gentle dorsoventral convexity, larger prox-
imal expansion and a relatively weak acromium. The ilia
of the two taxa differ considerably. The iliac blade of
Barapasaurus is comparatively high, and the dorsal margin
is rounded and gently convex; the preacetabular process is
quite prominent, curved and subtriangular (Text-fig. 11A–
B), whereas the iliac blade of Kotasaurus is quite low, and
as a result, the tall neural spine is visible in the mounted
skeleton; the dorsal margin of the ilium is almost straight,
and the preacetabular process extends comparatively
farther and rises above the level of pubic peduncle. The
caudal part of the ilium of Kotasaurus extends well beyond
the ischiadic peduncle in the mounted skeleton, but it is
broken caudally; an isolated specimen of the ilium in the
collection, though small, reveals a blunt shape of the cau-
dalmost part. The caudal part of the ilium of Kotasaurus is
larger than that of Barapasaurus. The acetabular part of
the ilium of Kotasaurus is wider than Barapasaurus. The
obturator foramen of Kotasaurus is significantly larger
than Barapasaurus. Another important difference is the
lesser trochanter of the femur which is present in Kotasau-
rus but completely absent in Barapasaurus. The fibula of
Barapasaurus has a broad triangular scar for the tibia
which is not noted in Kotasaurus. The presence of promi-
nent astragalar peg situated anteroventrally is characteristic
of Kotasaurus but is not found in Barapasaurus. From the
above comparative discussion, it is clear that Barapasaurus
has its own unique characteristic features that are different
and is more highly derived than Kotasaurus.
Concluding remarks
To conclude, the position of Barapasaurus within the
phylogenetic tree of the sauropods is redefined. More
scores on the different character states used by previous
workers to build up the sauropod phylogeny can be now
used for Barapasaurus through a detailed study of the
described as well many undescribed material belonging to
that taxon. The new phylogenetic tree suggests Barapasau-
rus is more advanced than Kotasaurus but is more basal
in comparison with Vulcanodon. However, Barapasaurus
is removed from Eusauropoda. Kotasaurus has been
accepted here as a valid taxon, though there may be irreg-
ularities in the mounted skeleton. All the material
assigned to Kotasaurus may not be of same taxon, but
there is undoubtedly at least a second distinct sauropod
present in the Kota Formation. Finally, it may be men-
tioned that the overall morphological features of B. tagor-
ei such as quadrupedal posture, spatulate teeth with
wrinkled enamel texture, shortening of the trunk, com-
plex vertebral laminae and pneumatic fossae in the presa-
cral series, articulation of hyposphene-hypantrum in the
dorsal vertebrae, strengthening of sacrum by the addition
of vertebra making four co-ossified sacral vertebrae, nar-
rowness of the pubic apron, and slender, columnar limbs
set the trend for future sauropod evolution.
Acknowledgements. The discovery of B. tagorei was made in 1958
by a team consisting of the late Pamela Lamplugh Robinson of
University College, London, the late S. L. Jain and T. K.
RoyChowdhury of the Indian Statistical Institute. T. S. Kutty
joined the team during the excavation in 1961. Subsequent exca-
vation (known as the ‘Colbert excavation’) in 1964 was carried
out by the late Edwin. H. Colbert, late S. L. Jain and late P. L.
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 565
Robinson. These scientists along with S. Chatterjee (presently at
Texas Tech University) did the basic descriptions of B. tagorei.
The expeditions were part of an integrated programme of the
study of the Gondwana rocks of the Pranhita-Godavari basin
which was sponsored by the Indian Statistical Institute and
partly financed by the Royal Society, London. SB and DPS are
thankful to T. K. RoyChowdhury for discussions and clarifica-
tion of certain aspects of the anatomy of B. tagorei. The authors
are grateful to T. K. RoyChowdhury for allowing them to use
his unpublished geological map of the Pranhita-Godavari basin.
Constructive criticism and suggestions by Jeff Wilson, Oliver
Rauhut and another anonymous reviewer have improved the
paper. We thank Victor Leshyk for the drawing in Text-figure 3.
Thanks are due to Dr. K. Ayyasami of the Geological Survey of
India, Hyderabad and Dr. B. G. Sidharth of the Birla Science
Museum, Hyderabad for access to Kotasaurus material. The late
Edwin H. Colbert encouraged this project and provided sage
advice in its early stages. The Colbert Endowment of The
Museum of Northern Arizona and the Indian Statistical Institute
provided travel and research fund to DDG.
Editor. Oliver Rauhut
REFERENCES
A L L A I N , R. and A Q U E S B I , N. 2008. Anatomy and phyloge-
netic relationships of Tazoudasaurus naimi (Dinosauria,
Sauropoda) from the Early Jurassic of Morocco. Geodiversitas,
30, 345–424.
—— D E J A X , J., M E Y E R , C., M O N BA R O N , M., M ON T E -
N A T , C., R I C H I R , P., R O CH D Y , M., R US S E L L , D. and
T A QU E T , P. 2004. A basal sauropod dinosaur from the early
Jurassic of Morocco. Comptes Rendus de l’Academie des Scien-
cies, 3, 199–208.
A V E R I A N O V , A. O. 2002. Early Cretaceous ‘symmetrodont’
mammal Gobitheriodon from Mongolia and the classification
of ‘Symmetrodonta’. Acta Palaeontologia Polonica, 47, 705–716.
B A N D Y O P A D H Y A Y , S. and R O Y C H O W D H U R Y , T. K.
1996. Beginning of the continental Jurassic in India: a pala-
eontological approach. Museum of Northern Arizona Bulletin,
60, 371–378.
—— and R UD R A , D. K. 1985. Upper Gondwana Stratigraphy,
north of the Pranhita-Godavari confluence, Southern India.
Journal Geological Society of India, 26, 261–266.
—— and S E N G UP TA , D. P. 2006. Vertebrate faunal turnover
during the Triassic–Jurassic transition: an Indian scenario. 77–
85. In H A R R I S , J. D., L U CA S , S. G., K I R K L A N D , J. D.
and M I L N E R , A. R. C. (eds). Terrestrial Triassic–Jurassic
transition. New Mexico Museum of Natural History and
Science Bulletin, 37, New Mexico, 607 pp.
—— R OY CH O W DH U R Y , T. K. and S E N GU PT A , D. P.
2002. Taphonomy of some Gondwana Vertebrate Assemblages
of India. Sedimentary Geology, 147, 219–245.
B A R R E T T , P. M. 1999. A sauropod dinosaur from the Lower
Lufeng Formation (Lower Jurassic) of Yunnan Province, Peo-
ple’s Republic of China. Journal of Vertebrate Paleontology, 19,
785–787.
—— and UP C HU R CH , P. 2007. The evolution of feeding
mechanisms in early sauropodomorph dinosaurs. 91–112. In
BA R R E T T , P. M. and B A T T E N , D. J. (eds). Evolution and
palaeobiology of early sauropodomorph dinosaurs. Special Papers
in Palaeontology, 77, 289 pp.
B E N T ON , M. J., J U UL , L., S T O R RS , G. W. and G A L -
T ON , P. M. 2000. Anatomy and systematics of the prosauro-
pod dinosaur Thecodontosaurus antiquus from the Upper
Triassic of southwest England. Journal of Vertebrate Paleontol-
ogy, 20, 77–108.
B ON A PA R T E , J. F. 1986. The early radiation and phyloge-
netic relationships of the Jurassic sauropod dinosaurs, based
on vertebral anatomy. In P A DI A N , K. (ed.) The Beginning of
the age of the dinosaurs. Cambridge University Press, Cam-
bridge, 376 pp.
B ON N A N , M. F. 2000. The presence of a calcaneum in a dipl-
odocid sauropod. Journal of Vertebrate Paleontology, 20, 317–
323.
—— 2003. The evolution of manus shape in sauropod dinosaurs:
implications for functional morphology, forelimb orientation
and sauropod phylogeny. Journal of Vertebrate Paleontology,
23, 595–613.
—— 2005. Pes Anatomy in sauropod dinosaurs: implications for
functional morphology, evolution, and phylogeny. 346–380. In
T I DW E L L , V. and CA R P E N T E R , K. (eds). Thunder-
Lizards: The Sauropodomorph Dinosaurs. Indiana University
Press, Bloomington, Indiana, 495 pp.
B OR S UK - BI A L Y N I CK A , M. 1977. A new camarasaurid
sauropod Opisthocoelicaudia skarzynskii, gen. n., sp. n. from
the Upper Cretaceous of Mongolia. Palaeontologia Polonica,
37, 341–349.
B R I T T , B. B. 1993. Pneumatic postcranial bones in dinosaurs
and other archosaurs. Ph. D. dissertation, University of Cal-
gary, Calgary, 383 pp.
B UF F E T A UT , E., S U T E E TH O R N , V., CU N Y , G., T ON G ,
H., L E L O E U FF , J., K H A N S UB H A , S. and J ON -
G A UT C HA R I Y A K UL , S. 2000. The earliest known sauro-
pod dinosaur. Nature, 407, 72–74.
C A L V O , J. O. 1994. Jaw mechanics in sauropod dinosaurs.
Gaia, 10, 183–193.
—— and S A L G A DO , L. 1995. Rebbachisaurus tessonei, sp.
nov. A new Sauropoda from the Albian-Cenomanian of
Argentina; new evidence on the origin of the Diplodocidae.
Gaia, 11, 13–33.
C HA RI G , A. J., A T T R I D G E , J. and CR O M P T ON , A. W.
1965. On the origin of the sauropods and the classification of
Saurischia. Proceedings Linnean Society of London, 176, 197–
221.
C OL B E R T , E. H. 1980. A Fossil Hunter’s Notebook, My Life
with Dinosaurs and Other Friends. E. P. Dutton, New York,
242 pp.
—— 1989. Digging Into the Past, An Autobiography. Red Demb-
ner Enterprises Corporation, New York, 456 pp.
C OO P E R , M. R. 1984. A reassessment of Vulcanodon karibaen-
sis Raath (Dinosauria: Saurischia) and the origin of the Sauro-
poda. Palaeontologia Africana, 25, 203–231.
D A T TA , P. M. 1981. The first Jurassic mammal from India.
Zoological Journal of Linnean Society of London, 73, 307–312.
566 P A L A E O N T O L O G Y , V O L U M E 5 3
—— and DA S , D. P. 2001. Indozostrodon simpsoni, gen. et. sp.
nov., an Early Jurassic megazostrodontid mammal from India.
Journal of Vertebrate Paleontology, 21, 528–534.
—— M A N N A , P., GH O S H , S. C. and DA S , D. P. 2000. The
first Jurassic turtle from India. Palaeontology, 43, 99–109.
D ON G Z H I - M I N G 1992. The dinosaurian faunas of China.
Springer–Verlag, Berlin, 188 pp.
E V A N S , S. E., PR A S A D, G. V. R. and M A N H A S , B. K.
2001. Rhynchocephalians (Diapsida : Lepidosauria) from the
Jurassic Kota Formation. Zoological Journal of the Linnean
Society, 133, 309–334.
—— ————2002. Fossil lizards from the Jurassic Kota Forma-
tion of India. Journal of Vertebrate Paleontology, 22, 299–312.
G A L T O N , P. M. and U PC H UR C H , P. 2004. Prosauropoda.
232–258. In W E I S H A M P E L , D. B., D O DS O N , P. and
O S M O L S KA , H. (eds). Dinosauria II. University of Califor-
nia Press, Berkeley, 880 pp.
—— and V A N HE E RD E N , J. 1985. Partial hindlimb of Bli-
kanasaurus cromptoni n. gen. and n. sp., representing a new
family of prosauropod dinosaurs from the Upper Triassic of
South Africa. Geobios, 18, 509–516.
—— UP CH U R CH , P. and V A N H E E R DE N , J. 1998. Anat-
omy of the prosauropod dinosaur Blikanasaurus cromptoni
(Upper Triassic, South Africa), with notes on the other tetra-
pods from the lower Elliot Formation. Palaontologische Zeit-
schrift, 72, 163–177.
—— V A N H E E R D E N and Y A T E S , A. M. 2005. The postcra-
nial anatomy of referred specimens of the sauropodomorph
dinosaur Melanorosaurus from the Upper Triassic of South
Africa. 1–37. In T I D W E L L , V. and C A R P E N T E R , K.
(eds). Thunder-lizards: The sauropodomorph dinosaurs. Indiana
University Press, Bloomington, Indiana, 495 pp.
G A U FF R E , F.–. X. 1993. The most recent Melanorosauridae
(Saurischia, Prosauropoda). Lower Jurassic of Lesotho with
remarks on the prosauropod phylogeny. Neues Jahrbuch fur
Geologie und Paleontlologie, Monatschefte, 1993, 648–654.
G A U T HI E R , J. 1986. Saurischian monophyly and the origin of
birds. 1–55. In PA D I A N , K. (ed.). The origin of birds and
the evolution of flight. Memoir California Academy of Science,
8, 98 pp.
G O V I N D A N , A. 1975. Jurassic fresh water ostracods from the
Kota limestone of India. Palaeontology, 18, 207–216.
G H OS H, J. G. 1994. Geological map of Gondwana basin of the
Pranhita-Godavari valley, Andhra Pradesh, India. Geological
Survey of India Publication, Sheet No. 1.
H A U GH T O N , S. H. 1924. The fauna and stratigraphy of the
Stormberg Series. Annals of South African Museum, 12, 323–
497.
H E X I N - L U , W A N G CH A N G - S N A H G , L I U S HA N G -
Z HO N G , Z H OU F E N G - Y U N , L I U TU - Q I A N G , C A I
KA I - J I and DA I B I N G 1998. A new species of sauropod
from the Early Jurassic of Gongxian County, Sichuan. Acta
Geologica Sichuan, 18, 1–6. [In Chinese, English Abstract].
H U E N E , F. von 1932. Die fossile Reptil-Ordnung Saurischia,
ihre Entwicklung und Geschichte. Monograph Geologie und
Palaeontologie, 4, 1–361.
J A I N , S. L. 1959. Fossil fishes from the Kota Formation of
India. Proceedings Geological society of London, 1565, 26–27.
—— 1973. New specimens of Lower Jurassic holostean fishes
from India. Palaeontology, 16, 149–177.
—— 1974a. Indocoelacanthus robustus n. gen., n. sp., (Coelacan-
thidae, Lower Jurassic) the first fossil coelacanth from India.
Journal of Paleontology, 48, 49–62.
—— 1974b. Jurassic pterosaur from India. Geological Society of
India, 15, 330–335.
—— 1980. The continental Lower Jurassic fauna from the Kota
Formation. 99–123. In J A CO B S , L. L. (ed.). Aspects of Verte-
brate History. Museum of Northern Arizona Press, Flagstaff,
407 pp.
—— 1983. A review of the genus Lepidotes (Actinopterygii:
Semionotoformes) with special reference to the species from
Kota Formation (Lower Jurassic), India, Journal Palaeontologi-
cal Society of India, 28, 7–42.
—— R O B I N S O N , P. L. and R O Y C H O W D H U R Y , T. K.
1962. A new vertebrate fauna from the Early Jurassic of Dec-
can, India. Nature, 194, 755–757.
—— KU T TY , T. S., RO Y C H OW DH UR Y , T. K. and
CH A T T E R J E E , S. 1975. The sauropod dinosaur from the
Lower Jurassic Kota Formation of India. Proceedings of the
Royal Society of London, 188A, 221–228.
—— —— —— —— 1979. Some characteristics of Barapasaurus
tagorei, a sauropod dinosaur from the Lower Jurassic of Dec-
can, India. 204–216. In L A S K A R , B. and R A J A R A O , C. S.
(eds). Fourth International Gondwana Symposium 1, Hindus-
than Publishing Corporation, New Delhi, 384 pp.
J A N E N S C H, W. 1961. Die Gliedmassen und Gliedmassengur-
tel der Sauropoden der Tendaguru-Schichten. Palaeontograph-
ica, 3, 177–235.
K I N G , W. 1881. The geology of the Pranhita–Godavari valley.
Memoir Geological Survey of India, 18, 151–311.
K UT T Y , T. S., C HA T T E R J E E , S., G A L TO N , P. M. and
UP C HU R CH , P. 2007. Basal sauropodomorphs (Dinosauria:
Saurischia) from the Lower Jurassic of India: their anatomy
and relationships. Journal of Paleontology, 81, 1552–1574.
M A R S H , O. C. 1878. Principal characters of American Jurassic
dinosaurs Part I. American Journal of Science, Series, 3, 16.
M CI N T OS H , J. S. 1990. Sauropoda. 345–401. In W E I S H A M -
PE L , D. B., DO DS O N , P. and O S M OL S K A , H. (eds). The
Dinosauria. University of California Press, Berkeley, 733 pp.
O W E N , R. 1842. Report on British fossil reptiles Part II. British
Association of Advancement of Science, 11, 60–204.
P A N DE , K. 2002. Age and duration of the Deccan Trap, India:
Constraints from Radiometric and Paleomagnetic data.
Proceedings Indian Academy of Sciences, 111, 115–123.
P A T T E R S ON , C. and O W E N , H. G. 1991. Indian isolation or
contact? A response to Briggs. Systematic Zoology, 40, 96–100.
P R A S A D , G. V. R. 2003. Stratigraphic distribution and diver-
sity of Mesozoic mammals of India. 124. In XIX Indian Collo-
quium on Micropalaeontology and Stratigraphy & Symposium
on Recent Developments in Indian Ocean Palaeontology and
Palaeoclimate: Varanasi, Benaras Hindu University, 277 pp.
—— and A R R A T I A , G. 2004. Elasmobranch and actinoptery-
gian remains from the Jurassic and Cretaceous of India. 625–
634. In A R R A T I A , G. and T I N TO R I , A. (eds). Mesozoic
Fishes 3 – Systematics, Palaeoenvironments and Biodiversity,
Verlag Dr. Friedrich Pfeil, Munchen, Germany, 649 pp.
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 567
—— and M A N H A S , B. K. 1997. A new symmetrodont mam-
mal from the Lower Jurassic Kota Formation, Pranhita-
Godavari valley, India. Geobios, 30, 563–572.
—— —— 2002. Triconodont mammals from the Jurassic Kota
Formation of India. Geodiversitas, 25, 445–464.
—— —— 2007. A new docodont mammal from the Jurassic
Kota Formation of India. Palaeontologia Electronica, 10, 1–11.
R A A T H , M. A. 1972. Fossil vertebrate studies in Rhodesia: a
new dinosaur (Reptilia: Saurischia) from near the Triassic–
Jurassic boundary. Arnoldia, 5, 1–37.
R A U HU T , O. W. M. and L O PE Z - A R B A R E L L O, A. 2008.
Archosaur evolution during the Jurassic: a southern perspec-
tive. Revista de la Associacion Geologica Argentina, 63, 557–
585.
R O BI N S O N , P. L. 1970. The Indian Gondwana formations a
review. First International Symposium on Gondwana Strati-
graphy, I. U. G. S., South America, 201–268.
—— and RO Y C H OW DH UR Y , T. K. 1962. A new vertebrate
fauna from the Early Jurassic of Deccan, India. Nature, 194,
755–757.
R U DR A , D. K. 1982. Upper Gondwana stratigraphy and sedi-
mentation in the Pranhita-Godavari valley, India. Quarterly
Journal of the Geological, Mining and Metallurgical Society of
India, 54, 56–79.
—— and M A U L I K , P. K. 1994. Lower Jurassic Kota Limestone
of India. 185–191. In G I E R L O W S K I - K O R D E S C H , E.
and KE L T S , K. (eds). Global Geological Record of Lake Basins,
Cambridge University Press, Cambridge, 461 pp.
R U S S E L L , D. A. and Z HE N G Z H O N G 1994. A large
mamenchisaurid from the Junggar Basin, Xinjiang, People’s
Republic of China. Canadian Journal of Earth Sciences, 30,
2082–2995.
S A L G A D O, L., C OR I A , R. A. and C A L V O, J. O. 1997. Evo-
lution of titanosaurid sauropods I: Phylogenetic analysis based
on postcranial evidence. Ameghinia, 34, 3–32.
S C H A E F FE R , B. and P A T TE R S ON , C. 1984. Jurassic fishes
from the Western United States with comments on Jurassic
fish distribution. American Museum Novitates, 2796, 1–86.
S E E L E Y , H. G. 1888. The classification of the Dinosauria.
Report British Association of Advancement of Science, 1887,
698–699.
S E N G U PT A , S. 1970. Gondwana sedimentation around
Bheemaram (Bhimaram), Pranhita – Godavari valley, India.
Journal of Sedimentary Petrology, 40, 140–170.
S E R E N O , P. C. 1997. The origin and evolution of dinosaurs.
Annual Reviews of Earth and Planetary Sciences, 25, 435–489.
—— 1998. A rationale for phylogenetic definitions, with applica-
tion to the higher level of taxonomy of Dinosauria. Neues
Jahrbuch fur Geologie und Palaontologie, Abhandlungen, 210,
41–83.
S I M M ON S , D. J. 1965. The non-therapsid reptiles of the Luf-
eng Basin, Yunnan, China. Fieldiana Geology, 15, 1–93.
S M I T H , N. D. and P OL , D. 2007. Anatomy of a basal saur-
opodomorph from the Early Jurassic Hanson Formation of
Antarctica. Acta Palaeontologia Polonica, 52, 657–674.
S W OF F OR D , D. L. 2000. PAUP*. Phylogenetic Analysis Using
Parsimony (*and other methods), Version 4.0b4a. Sinauer
Associates Sunderland, Massachusetts.
U PC H UR C H, P. 1995. The evolutionary history of sauropod
dinosaurs. Philosophical Transactions of the Royal Society of
London, Series B, 349, 365–390.
—— 1998. The phylogenetic relationships of sauropod dinosaurs.
Zoological Journal of the Linnean Society, 124, 43–103.
—— B A R R E T T , P. M. and D OD S O N , P. 2004. Sauropoda.
259–322. In W E I S HA M P E L , D. B., D OD S O N , P. and
O S M O LS KA , H. (eds). Dinosauria II. University of Califor-
nia Press, Berkeley, 880 pp.
—— —— and G A L T O N , P. M. 2007a. A phylogenetic analysis
of basal sauropodomorph relationships: implications for the
origin of sauropod dinosaurs. 57–90. In B A R R E T T , P. M.
and B A T T E N , D. J. (eds). Evolution and palaeobiology of
early sauropodomorph dinosaurs. Special Papers in Palaeonto-
logy, 77, 289 pp.
—— —— Z H A O X I J I N and X U X I N G 2007b. A re-evalua-
tion of Chinshakiangosaurus chunghoensis Ye vide Dong 1992
(Dinosauria, Sauropodomorpha): implications for cranial evo-
lution in basal sauropod dinosaurs. Geological Magazine, 144,
247–262.
W E D E L , M. J. 2003. The evolution of vertebral pneumaticity
in sauropod dinosaurs. Journal of Vertebrate Paleontology, 23,
344–357.
—— C I F E L L I , R. L. and S A N DE R S , R. 2000. Osteology,
paleobiology, and relationshios of the sauropod dinosaur
Sauropseidon. Acta Palaeontologica Polonica, 45, 343–388.
W I L S ON , J. A. 1999. A nomenclature for vertebral laminae in
sauropods and other saurischian dinosaurs. Journal of Verte-
brate Paleontology, 19, 639–653.
—— 2002. Sauropod dinosaur phylogeny: critique and cladistic
analysis. Zoological Journal of the Linnean Society, 136, 217–
276.
—— 2005a. Overview of sauropod phylogeny and evolution.
15–49. In C U RR Y - R O G E R S , K. A. and W I L S O N , J. A.
(eds). The Sauropods: Evolution and Paleobiology. University of
California Press, Berkeley, 349 pp.
—— 2005b. Integrating ichnofossil and body fossil records to
estimate locomotor posture and spatiotemporal distribution of
early sauropod dinosaurs: a stratocladistic approach. Paleo-
biology, 31, 400–423.
—— and S E R E N O , P. C. 1998. Early evolution and higher-
level phylogeny of sauropod dinosaurs. Journal of Vertebrate
Paleontology (Memoir 5), 18, 1–68.
Y A D A G I R I , P. 1984. New symmetrodonts from Kota Forma-
tion (Early Jurassic), India. Journal Geological Society of India,
25, 514–621.
—— 1985. An amphidontid symmetrodont from the Early Juras-
sic Kota Formation, India. Zoological Journal of the Linnean
Society of London, 85, 411–417.
—— 1986. Lower Jurassic lower vertebrates from Kota Forma-
tion, Pranhita-Godavari valley, India, Journal of the Palaeonto-
logical Society of India, 31, 89–96.
—— 1988. A new sauropod Kotasaurus yamanpalliensis from
Lower Jurassic Kota Formation of India. Records of the Geolog-
ical Survey of India, 11, 102–127.
—— 2001. The osteology of Kotasaurus yamanpalliensis, a sauro-
pod dinosaur from the Early Jurassic Kota Formation. Journal
of Vertebrate Paleontology, 21, 242–252.
568 P A L A E O N T O L O G Y , V O L U M E 5 3
—— and PR A S A D, K. N. 1977. On the discovery of new Pholid-
ophorus fishes from the Kota Formation, Adilabad District,
Andhra Pradesh. Journal Geological Society of India, 18, 436–444.
Y A TE S , A. M. 2003. A new species of the primitive dinosaur
Thecodontosaurus (Saurischia: Sauropodomorpha) and its
implications for the systematics of early dinosaurs. Journal of
Systematic Biology, 1, 1–42.
—— 2007a. The first complete skull of the Triassic dinosaur
Melanorosaurus Haughton (Sauropodomorpha: Anchisauria).
9–55. In B A R R E T T, P. M. and B A T T E N , D. J. (eds). Evo-
lution and palaeobiology of early sauropodomorph dinosaurs.
Special Papers in Palaeontology, 77, 289 pp.
—— 2007b. Solving a dinosaurian puzzle: the identity of Aliwalia
rex Galton. Historical Biology, 19, 93–123.
—— and K I T CH I N G, J. W. 2003. The earliest known sauro-
pod dinosaur and the first steps toward sauropod locomotion.
Proceedings of the Royal Society of London, 270, 1753–1758.
Y O U N G , C. C. 1951. The Lufeng saurischian fauna in China.
Palaeontologia Sinica Series C, 13, 1–96.
Z H A N G Y I - H O N G and Y A N G Z H A O - L O N G 1994. A
complete osteology of Prosauropoda in the Lufeng Series, Yun-
nan, China. Jingshanosaurus. Yunnan Science and Technology
Publishing House, Kunming, 100 pp. [In Chinese, English
summary].
B A N D Y O P A D H Y A Y E T A L . : O S T E O L O G Y O F B A R A P A S A U R U S T A G O R E I 569
Top Related