OSTEOLOGY OF BARAPASAURUS TAGOREI (DINOSAURIA: SAUROPODA) FROM THE EARLY JURASSIC OF INDIA

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

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4)


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


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).


(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


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.


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.


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.


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.


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.


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.


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.


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.


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.


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.


(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.


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.


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.


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.


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.


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


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


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


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


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


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


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).


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.


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


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


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.


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

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OSTEOLOGY OF BARAPASAURUS TAGOREI (DINOSAURIA: SAUROPODA) FROM THE EARLY JURASSIC OF INDIA

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