Purported latest bone of a plated dinosaur (Ornithischia: Stegosauria), a "dermal plate" from the Maastrichtian (Upper Cretaceous) of southern India

Abstract

A stegosaurian "dermal plate" was reported from the Kallamedu Formation (Upper Cretaceous, Maastrichtian) of southern India. However, histologically the dermal plates of stegosaurs, typified by Stegosaurus (Upper Jurassic, USA), have a thin outer cortex enclosing very cancellous bone having large vascular spaces. The Kallamedu fragment of eroded compact bone has no cortex and is probably from a sauropod dinosaur. Bones found in situ in this formation typically disintegrate very quickly pon exposure to the air, but this bone is well preserved, although worn. It was found as float in a stream bed, indicating that there is a stratum upstream with well-preserved bones, that is still to be discovered in the Maastrichtian of southern India. Stegosaur remains from the underlying Coniacian (Upper Cretaceous) of southern India represents the most recent osteological record of a stegosaur. However, stegosaurs may have continued into the Maastrichtian (Lameta Formation) of western India as indicated by a pes print of Deltapodus sp., a stegosaurian ichnotaxon. © 2017 E. Schweizerbart'sche Verlagsbuchhandlung, Stuttgart, Germany.

Full text

uncorrected proofs
Purported latest bone of a plated dinosaur (Ornithischia: Stegosauria),
a “dermal plate” from the Maastrichtian (Upper Cretaceous) of
southern India
Peter M. Galton and Krishnan Ayyasami
With 1 figure
Abstract: A stegosaurian “dermal plate” was reported from the Kallamedu Formation (Upper Cre-
taceous, Maastrichtian) of southern India. However, histologically the dermal plates of stegosaurs,
typified by Stegosaurus (Upper Jurassic, USA), have a thin outer cortex enclosing very cancellous bone
having large vascular spaces. The Kallamedu fragment of eroded compact bone has no cortex and is
probably from a sauropod dinosaur. Bones found in situ in this formation typically disintegrate very
quickly pon exposure to the air, but this bone is well preserved, although worn. It was found as float in
a stream bed, indicating that there is a stratum upstream with well-preserved bones, that is still to be
discovered in the Maastrichtian of southern India. Stegosaur remains from the underlying Coniacian
(Upper Cretaceous) of southern India represents the most recent osteological record of a stegosaur.
However, stegosaurs may have continued into the Maastrichtian (Lameta Formation) of western India
as indicated by a pes print of Deltapodus sp., a stegosaurian ichnotaxon.
Key words: Dinosauria, Ornithischia, Stegosauria, Dravidosaurus, Deltapodus, Upper Cretaceous,
India, bones, footprints.
1. Introduction
Stegosauria is a clade of quadrupedal, graviportal,
herbivorous ornithischian dinosaurs with an array of
dermal plates and spines in two parasagittal rows that
extend along the top of the body. The group is best rep-
resented by partial to complete skeletons and isolated
bones from the Middle and Late Jurassic but there are
only a few records from the Early Cretaceous (Galton
& UpchUrch 2004; MaidMent et al. 2008; MaidMent
2010; Galton 2012; pereda-SUberbiola et al. 2012,
2015; borinder et al. 2016). The most recent occur-
rence for stegosaurs is from the Late Cretaceous of
India.
anonyMoUS (1978) reported the discovery of bones
excavated during a geological survey by Ponnala yad-
aGari and KriShnan ayyaSaMi of the Kallamedu For-
mation (uppermost Upper Cretaceous, Maastrichtian).
The site was in the Cauvery Basin near Kallamedu in
the Tiruchirapalli district, Tamil Nadu, southern India.
The bones included those interpreted as stegosaurian
and a large dermal plate was illustrated. Subsequently,
yadaGari & ayyaSaMi (1979: 529) noted the occurrence
of theropod, sauropod and stegosaur bones from above
a layer containing a lower Maastrichtian foraminiferal
fauna (Pachydiscus otacodensis Zone, yadaGari &
a
yyaSaMi
1979, tab. 1, fig. 523, map). The very large
bones, which came from this excavation site north east
of Kallamadu Village, were described as the theropod
Bruhathkayosaurus matleyi yadaGiri & ayyaSaMi,
1987. However, this taxon is listed as a nomen dubi-
um, Sauropoda indet., by U
pchUrch
et al. (2004; also
H
one
et al. 2016) and, as one of us (a
yyaSaMi
) notes,
only bones of this taxon were found in situ and none
©2017 E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany www.schweizerbart.de
DO I : 10.1127/ njgpa/20 17/0671 007 7-7 749/2 017/0 671 $ 0.0 0
N. Jb. Geol. Paläont. Abh. 285/1 (2017), 241–000 Article
Stuttgart, July 2017
E

uncorrected proofs
2 P.M. Galton and K. Ayyasami
of stegosaurs. New vertebrate remains, including dino-
saur teeth, are reported from the Kallamadu Formation
(praSad et al. 2013; GoSwaMi et al. 2013) and the verte-
brate fauna of the Maastrichtian of the Cauvery Basin
is discussed by VerMa (2015; also VerMa et al. 2016).
Previously, the partial skull and postcranial skel-
eton of the stegosaur Dravidosaurus blanfordi y
ada
-
Gari & ayyaSaMi, 1979 was described from the older
marine limestone beds in the Trichinopoly Group (Co-
niacian, Kosmaticeras theobaldianum Zone), also in
the Cauvery Basin west of Siranattam Village in the
Tirachirapalli district of southern India (see yada-
Gari
& a
yyaSaMi
1979, tab. 1, fig. 1). G
alton
(1981)
re-identified some of the cranial bones of Dravidos-
aurus blanfordi and mistakenly cited the plate figured
in anonyMoUS (1978) as belonging to this taxon (see
Section 3 for further details on Dravidosaurus).
The citations by Galton (1981; also 1990: 435, 2012:
296; G
alton
& U
pchUrch
2004: 343; p
ereda
-S
Uberbio
-
la et al. 2012: 75) for the occurrence of stegosaur bones
in the Maastrichtian of India, the latest record of the
group, is based on the claim by yadaGari & ayyaSaMi
(1979: 529) given above. However, the only informa-
tion provided to date on this material is a photograph
of the dermal plate in anonyMoUS (1978). Owing to
the potential importance of such a geologically young
stegosaur, a more thorough description is long overdue.
Institutional abbreviations: GSIG, Geological Survey of
India, Western Region Gandhinagar, Gujarat, India; GSIH,
Geological Survey of India, Southern Region, Hyderabad,
India; and YPM VP, Division of Vertebrate Paleontology,
Peabody Museum of Natural History, Yale University, New
Haven, Connecticut, USA.
2. Description and comparisons
Dermal armor occurs in three groups of dinosaurs that
are possibly represented in the Upper Cretaceous of
India: stegosaurs (yadaGiri & ayyaSaMi 1979; Galton
1981), ankylosaurs (c
hatterjee
& r
Udra
1996: 518;
undescribed vertebrae, scapulocoracoid, humerus, fe
-
mur, osteoderms), and titanosaurian sauropods (the
armor of “Lametosaurus indicus” of Matley 192 4,
pls. 12, 13 that is not ankylosaurian, see c
arrano
&
S
aMpSon
2008: 208). Using the dermal plates of Stego-
saurus as a comparative standard, the plates consist
of a thin layer of compact bone enclosing cancellous
bone, with large vascular spaces that, because of its
open texture (Fig. 1G; B
Uffrénil
et al. 1986; M
ain
et
al. 2005; HayaShi et al. 2012), would not be preserved
without the protective outer cortex. This is also true for
the dermal armor of ankylosaurs (h
ayaShi
et al. 2010).
HUlKe (1881, pl. 76, fig. 1) figured a large lateral spike
of the nodosaurid ankylosaur Polacanthus foxii ( Lower
Cretaceous, England) that is represented by a partial
mostly hollow thin shell of outer cortex with most of
the internal cancellous bone eroded away.
Apart from being sub-triangular in outline in
“side” views (Fig. 1A, D), this bone does not resemble
any of the dermal plates of the Morrison (Upper Ju-
rassic) stegosaurs Stegosaurus stenops (see GilMore
1914; MaidMent et al. 2015) and Hesperosaurus mjosi
(Carpenter et al. 2001; Carpenter 2010; Saitta 2015
as Stegosaurus mjosi), or those of any other stegosaur
or of any ankylosaur (see skeletal restorations in PaUl
2016). This is especially true for the dermal armor of
titanosaurian sauropods (see Matley 1924, pls. 12, 13;
also not equivalent to any of the four types of armor
that occur in titanosaurs, see d’eMic et al. 2009).
Unlike thyreophoran dermal plates, this bone
(ca. 175 mm long close to base, ca. 190 mm high, ca.
65 mm at thickest) does not consist of cancellous bone
with large vascular spaces Fig. 1A-E cf. Fig. 1F) or
taper from base to apex in “edge-on” views (Fig. 1B, C
cf. Fig. 1A). There are no vertical vascular grooves on
the outer surface of the cortex, as occur in Morrison
stegosaurs (see GilMore 1914; oStroM & McintoSh
1999; G
alton
2010), because this layer is completely
missing. The only smooth surfaces are those of unpre-
pared matrix (m, Fig. 1B-E). The bone surfaces are all
very rough and irregular with no trace of an externally
smooth thin outer layer of compact bone. This is not
an artefact of mechanical preparation because the bone
surfaces at cross-sections of the matrix-bone intersec-
tion are similar (Fig. 1B, C, E). This “dermal plate” is
an eroded piece of compact bone that, based on its size,
was probably derived from a long bone of a sauropod
dinosaur. In this context it is interesting that h
one
et
al. (2016: 7) cite Bruhathkayosaurus (from Kallamedu
Formation near Kallamedu) as an anomalously large
sauropod dinosaur (yadaGiri & ayyaSaMi 1987: ilium
length 1200 mm, femur distal condylar width 750 mm,
tibia length 2000 mm).
3. Discussion
With reference to the Kallamedu Formation (Upper
Cretaceous, Maastrichtian) near Kallamedu, Tiruchi-

uncorrected proofs
Purported latest bone of a plated dinosaur (Ornithischia: Stegosauria) 3
Fig. 1. A-E – Alleged stegosaur dermal plate originally figured by A
nonyMoUS
(1978), GSIH (unnumbered) from Upper
Cretaceous (Maastrichtian) of southern India. It is here identified as a piece of eroded compact bone, possibly part of the
long bone of a sauropod dinosaur: A, side view matching original figure; B-C, edge on views of A: B, from left and C,
from right; D, opposite side to A and E, basal view with surface A below. F – Stegosaurian ichnotaxon Deltapodus sp.,
GSIG (unnumbered) from Upper Cretaceous (Maastrichtian) of western India, plaster cast taken from imprint of right pes.
G – Stegosaurus stenops, YPM VP 1856 [1512Y, Box 3], Upper Jurassic of Como Bluff, Wyoming, USA, vertical transverse
section of large dermal plate in the basal region (from bUffrénil et al. 1986). Abbreviations: m, matrix; II, digit II; scale
bars = 50 mm (A-E), 100 mm (F) and 10 mm (G).

uncorrected proofs
4 P.M. Galton and K. Ayyasami
rapalli district in the Cauvery Basin of southern India,
blanford (1864: 139) noted that the beds consisted of a
mass of white sands and grey sandy clays and “imbed-
ded in the deposits large bones are numerous, but so
saturated with water and so very friable, that it is im-
possible, even with the greatest care, to extract them in
anything like a recognizable condition.” Matley (1929)
noted that this was the situation during the monsoon
season whereas during the dry season desiccation, with
expansion by day and contraction at night, results in
the bones being split into fragments. Consequently,
the bones are poorly preserved and the photographs
used for Bruhathkayosaurus by yadaGari & ayyaSaMi
(1987) were taken while the bones were still exposed
in situ. The bones started to disintegrate in the field
jackets even before reaching the GSIH and no longer
exist. This is in marked contrast to the “dermal plate”
(Fig. 1A-E) that, even though lacking a resistant outer
cortex, is still very well preserved. However, one of
us (K.A.) was present when Panalla YadaGiri located
this isolated piece of bone near the excavation site as
float in the bed of a stream, a tributary of the Maru-
daiyar River (yadaGari & ayyaSaMi 1979, fig. 1, map).
Consequently, it was not found in situ with the other
bones that, as a
yyaSaMi
notes, did not include any of
stegosaurs. This is extremely important because some-
where upstream to this site there should be a horizon
with very well preserved large bones, something still
to be discovered in the Maastrichtian of southern India.
As regards to Dravidosaurus blanfordi, c
hatterjee
& rUdra (1996: 518) noted that “we visited the site
and found only fragmentary remains of plesiosaurs. We
also examined the holotype and could not see anything
related to the stegosaurian plates and skull claimed by
these authors. Instead, the bones are highly weathered
limb and girdle elements and may belong to plesio-
saurs.” This was cited by MaidMent et al. (2008; also
M
aidMent
2010) who, until the bones are independently
re-described, regard this taxon as a nomen dubium and
the specimen as Stegosauria indet. (but as ?Stegosauria
indet., p
ereda
-S
Uberbiola
et al. 2015; not demonstrably
ornithischian, WilSon et al. 2011: 982). VerMa (2 015:
57) lists this taxon as a marine reptile and it is given
as a taxon of plesiosaur by V
erMa
et al. (2016: 320).
However, there is no correspondence between the pho-
tographs of the bones of Dravidosaurus in y
adaGari
& ayyaSaMi (1979, pls. 1-4; also figs. 2-7) with figures
of the limbs and girdles of a plesiosaur (cf. a
ndrewS
1910, 1913; WilliSton 1903; O’GorMan et al. 2015).
This is especially true for a small tooth (crown width
and height ca 1 mm with three crenulations, total length
with long root ca 2.5 mm; yadaGari & ayyaSaMi 1979,
fig. 4, pl. 1, fig. 3a, b). However, one of us (K.a.) is cur-
rently working on undescribed bones that support the
presence of a stegosaur in the type quarry in the Conia-
cian (Upper Cretaceous) of southern India.
An isolated impression of a small right pes (225
mm long), the basis for a plaster cast (Fig. 1F), was
found in the uppermost limestone unit of the Lameta
Formation (given as Infratrappean Sequence; Maas-
trichtian) of the Jetholi area near Balasinor in Kheda
District in Gujarat, western India (see Mohabey 1986
for photos and map). This pes print was referred to the
stegosaurian ichnotaxon Deltapodus sp. by MateUS et
al. (2011). Deltapodus brodricki whyte & roMano,
1995, which was based on a supposed sauropod track
from the Middle Jurassic of England, was re-identified
as representing the prints of a three toed stegosaur like
Stegosaurus by w
hyte
& r
oMano
(2001, tab. 1 for de-
tails; for photographs in situ as preserved, see whyte &
r
oMano
1993, fig. 3 for holotype and L
oMax
& t
aMUra
2014, figs. 130A, B, 131A, B for other examples). The
identification of Deltapodus brodricki as a stegosaurian
footprint is supported by subsequent studies as cited in
Galton (2017). The Indian pes print of Deltapodus sp.
(Fig. 1F) agrees with those of Deltapodus brodricki
(see w
hyte
& r
oMano
2001, fig. 3A, tab. 1; G
alton
2017, fig. 2N) in several characters, viz., it is triangular
with a well-developed heel, digitigrade, three very wide
blunt digits, toes radiating and not separated by well-
developed hypicies (angles), and weakly mesaxonic
(digit III only slightly longer than II and IV) (w
hyte
& r
oMano
20 01; L
i
et al. 2012). Deltapodus also oc-
curs in the Lower Cretaceous of Spain (coboS et al.
2010; paScUal et al. 2012), western USA (Milân et al.
2015) and China (x
inG
et al. 2013). As M
ateUS
et al.
(2011: 656) note, the occurrence of skeletal remains
of a stegosaur from the Upper Cretaceous (Coniacian)
of southern India would be in accord with the record
of the stegosaurian Deltapodus-like pes print from the
Maastrichtian of western India.
Acknowledgements
PMG thanks D. M
ohabey
(Nagpur University, Nagpur, India)
for a copy of the original photograph (Fig. 1F) and for try-
ing to locate the cast of the pes in the GSIG collection. We
thank the reviewers P.M. barrett (Natural History Museum,
London, UK), K. Carpenter (University of Utah – Eastern,
Price, Utah, USA, and J.I. KirKland (Utah Geological Sur-
vey, Salt Lake City, Utah, USA) for their constructive com-
ments that improved this paper.

uncorrected proofs
Purported latest bone of a plated dinosaur (Ornithischia: Stegosauria) 5
References
andrewS, c.w. (1910): A Descriptive Catalogue of the Ma-
rine Reptiles of the Oxford Clay. Based on the l
eedS
Col-
lection in the British Museum (Natural History), London.
Part I. – xvii + 205 pp.; London (British Museum).
andrewS, c.w. (1913): A Descriptive Catalogue of the Ma-
rine Reptiles of the Oxford Clay. Based on the l
eedS
Col-
lection in the British Museum (Natural History), London.
Part II. – xix + 206 pp.; London (British Museum).
anonyMoUS (1978): New dinosaurian remains. – Geological
Survey of India, News, 9 (5): 4.
b
lanford
, H.F. (1864): On the Cretaceous and other rocks of
the South Arcot and Trichinopoly districts, south India. –
Memoirs of the Geological Survey of India, 4 (1): 1-127.
borinder, n.h., poropat, S.f. & Kear, b.p. (2016): Reas-
sessment of the earliest documented stegosaurian fossils
from Asia. – Cretaceous Research, 68: 61-69.
bUffr énil, V. de, farlow, j.o. & ricqléS, a. de (1986):
Growth and function of Stegosaurus plates: evidence
from bone histology. – Paleobiology, 12 (4): 459- 473.
c
arpent er
, K. (2010): Species concept in North American
stegosaurs. – Swiss Journal of Geosciences, 103 (2): 155-
162.
carpent er, K., MileS, c.a. & cloward, K. (20 01): Ne w
primitive stegosaur from the Morrison Formation, Wyo-
ming. – In: carpent er, K. (Ed.): The Armored Dino-
saurs: 55-75; Bloomington (Indiana University Press).
c
arrano
, M.t. & S
aMpSon
, S.d. (2008): The phylogeny of
Ceratosauria (Dinosauria: Theropoda). – Journal of Sys-
tematic Palaeontology, 6 (2): 183-236.
c
hatterjee
, S. & r
Udra
, d.K. (1996): KT events in India:
impact, rifting, volcanism and dinosaur extinction. –
Memoirs of the Queensland Museum, 39: 489-532.
coboS, a., royo-terreS, r., lUqUe, l., alcalá, l. &
MaMpel, L. (2010): An Iberian stegosaurs paradise: The
Villard et Arzobispo Formation (Tithonian-Berriasian)
in Teruel (Spain). – Palaeogeography, Palaeoclimatology,
Palaeoecology, 293: 223-236.
D’eMic, M.d., wilSon, j.a. & chatterjee, S. (2009): The
titanosaur (Dinosauria: Sauropoda) osteoderm record:
Review and first definitive specimen from India. – Jour-
nal of Vertebrate Paleontology, 29 (1) : 165-177
Galton, P.M. (1981): Craterosaurus pottonensis Seeley, a
stegosaurian dinosaur from the Lower Cretaceous of
England, and a review of Cretaceous stegosaurs. – Neues
Jahrbuch für Geologie und Paläontologie, Abhandlungen,
161: 28-46.
G
alton
, p.M. (1990): Stegosauria. – In: w
eiShaMpel
, d.b.,
dodSon, p. & oSMólSKa, H. (Eds.): The Dinosauria (1
st
edition): 343-362; Berkeley (University of California
Pre ss).
Galton, P.M. (2010): Species of plated dinosaur Stegosaurus
(Morrison Formation, Late Jurassic) of western USA:
new type species designation needed. – Swiss Journal of
Geosciences, 103 (2): 187-198.
Galton, P.M. (2012): Stegosaurs. – In: brett-SUrMan, M.K.,
holtz, t.r. Jr. & farlow, J.O. (Eds.): The Complete
Dinosaur (2nd edition): 482-504; Bloomington (Indiana
Un iversit y P ress).
G
alton
, P.M. (2017): Purported earliest bones of a plated di-
nosaur (Ornithischia: Stegosauria): a “dermal spine” and
a centrum from the Aalenian-Bajocian (Middle Jurassic)
of England, with comments on other early thyreopho-
rans. – Neues Jahrbuch für Geologie und Paläontologie,
Abhandlungen, 285 (1): 1-10.
Galton, p.M. & UpchU rch, P. (2004): Stegosauria. – In:
w
eiShaMpel
, d.b., d
odSon
, p. & o
SMólSKa
, H. (Eds.):
The Dinosauria (2
nd
edition): 343-362; Berkeley (Univer-
sity of California Press).
G
ilMore
, C.W. (1914): Osteology of the armoured Dinosauria
in the United States National Museum, with special refer-
ence to the genus Stegosaurus. – Bulletins of the United
States National Museum, 89: 1-143.
G
oSwaMi
, a., p
raSad
, G.V.r., V
erMa
, o., f
lynn
, j.j. & b
en
-
Son, r.b.j. (2013): A troodontid dinosaur from the latest
Cretaceous of India. – Nature Communications, DOI:
10.1038/ncom ms2716.
h
ayaShi
, S., c
arpent er
, K., S
cheyer
, t.M., w
atabe
, M. &
SUzUKi, D. (2010): Function and evolution of ankylosau-
rian dermal armor. – Acta Palaeontologica Polonica, 55
(2): 213-228.
h
ayaShi
, S., c
arpenter
, K., w
atabe
, M. & M
c
w
hinney
, l.a.
(2012): Ontogenetic histology of Stegosaurus plates and
spikes. – Palaeontology, 55 (1): 145 -161.
hone, d.w.e., farKe, a.a. & wedel, M.j. (2016): Ontog-
eny and the fossil record: what, if anything, is an adult
dinosaur. – Biological Letters, 12: 1-8, 20150947 http://
dx.doi.org/10.1098/rsbl.2015.0947
hUlKe, j.w. (1881): Polacanthus foxii, a large undescribed
dinosaur from the Wealden Formation in the Isle of
Wight. – Philosophical Transactions of the Royal Society
of London, 172: 653-662.
li, j., locKley, M.G., zha nG, Y., hU, S., MatSUKawa, M.
& b
ai
, Z. (2012): An important ornithischian tracksite
in the Early Jurassic of the Shenmu Region, Shaanxi,
China. – Acta Geologica Sinica, 86 (1): 1-10.
loMax, d.r. & taMUra, N. (2014): Dinosaurs of the British
Isles. – 414 pp.; Manchester (Siri Scientific Press).
M
aidMent
, S.C.R. (2010): Stegosauria: a historical review of
the body fossil record and phylogenetic relationships. –
Swiss Journal of Geosciences, 103 (2): 199-210.
MaidMent, S.c.r., braSSey, c. & bar rett, p.M. (2015):
The postcranial skeleton of an exceptionally complete
individual of the plated dinosaur Stegosaurus stenops
(Dinosauria: Thyreophora) from the Upper Jurassic Mor-
rison Formation of Wyoming, U.S.A. – PloSONE 10 (10):
e0138352. Doi:10.1271/jour na l.p one.0138352
MaidMent, S.c.r., norMan, d.b., barrett, p.M. & Up-
chUrch, P. (2008): Systematics and phylogeny of Stego-
sauria (Dinosauria: Ornithischia). – Journal of System-
atic Palaeontology, 6: 3 6 7 - 4 07.
Main, r.p., ricqlèS, A. de, horner, j.r. & padian, K. (20 05):
The evolution and function of thyreophoran dinosaur
scutes: implications for plate function in stegosaurs. –
Paleobiology, 31 (2): 291-314.
MateUS, o., Milàn, j., roMano, M. & whyte, M.A. (2011):
New finds of stegosaur tracks from the Upper Jurassic
Lourinhã Formation, Portugal. – Acta Palaeontologica
Polonica, 56 (3): 651-658.
Matley, C.A. (1924): Notes on an armoured dinosaur from
the Lameta beds of Jubbulpore. – Records of the Geo-

uncorrected proofs
6 P.M. Galton and K. Ayyasami
logical Survey of India, 55 (2): 105-109.
Matley, C.A. (1929): The Cretaceous dinosaurs of the
Trichinopoly District, and the rocks associated with
them. – Records of the Geological Survey of India, 61
(4): 337-349.
Milan, j., chiappe, l.M., loope, d.b., Ki rKla nd, j.l. &
locKley, M.G. (2015): First report on dinosaur tracks
from the Burro Canyon Formation, San Juan County,
Utah, USA – evidence of a diverse, hitherto unknown
Lower Cretaceous dinosaur fauna. – Annales Societatis
Geologorum Poloniae, 85: 515 -525.
Mohabey, D.M. (1986): Note on dinosaur footprint from
Kheda District, Gujarat. – Journal of the Geological So-
ciety of India, 27: 456-459.
o’GorMan, j.p., SalGado, l., oliVero, e. & MarenSSi, S.A.
(2015): Vegasaurus molyi, gen. et sp. nov. (Plesiosauria,
Elasmosauridae), from the Cape Lamb Member (Lower
Maastrichtian) of the Snow Hill Island Formation, Vega
Island, Antarctica, and remarks on Wedellian Elasmo-
sauridae. – Journal of Vertebrate Paleontology, 35 (3):
e931285 (21 pp). DOI: 10.1080/02724634.2014.931285.
oStroM, j.a. & McintoSh, J.S. (1999): Marshʼs Dinosaurs.
The Collections from Como Bluff. With a New Foreword
by P
eter
d
odSon
and a Historical Update by C
lifford
a. MileS and daVid w. haMblin. – xxiv + 388 pp.; New
Haven (Yale University Press).
paScUal, c., canUdo, j.i., hernandez, n., barco, j.l. &
c
aStanera
, d. (2012): First record of stegosaur dinosaur
tracks in the Lower Cretaceous (Berriasian) of Europe
(Oncala Group, Soria, Spain). – Geodiversitas, 34 (2):
29 7-312.
PaUl, G.S. (2016): The Princeton Field Guide to Dinosaurs
(2nd edition). – 360 pp.; Princeton (Princeton University
Pre ss).
pereda SUber biola, x., día z-Martínez, i., SalGado, l. &
ValaiS, S. de (2015): Síntesis del registro fósil de dino-
saurios tireóforos en Gondwana. – Argentinian Paleon-
tological Association Publication Electronica, 15 (1):
9 0 -1 0 7.
pereda SUberbiola, x., Galton, p.M., MalliSon, h. & no-
VaS
, F. (2012): A plated dinosaur (Ornithischia, Stego-
sauria) from the Early Cretaceous of Argentina, South
America: an evaluation. – Alcheringa, 37: 65-78.
p
raSad
, G.V.r., V
erMa
, o., f
lynn
, j.j. & G
oSwaMi
, a. (2013):
A new Late Cretaceous vertebrate fauna from the Cau-
very Basin, South India: implications for Gondwanan
paleobiogeography. – Journal of Vertebrate Paleontol-
og y, 33 (6): 1260-1268.
Saitta, e.T. (2015): Evidence for sexual dimorphism in
the plated dinosaur Stegosaurus mjosi (Ornithischia,
Stegosauria) from the Morrison Formation (Upper Ju-
rassic) of western USA. – PloSONE, 10 (4): e0123503.
DOI:10.1371/journal.pone.0123503
UpchUrch, p., barrett, p.M. & dodSon, p. (200 4): Sauropo-
da. – In: weiShaMpel, d.b., dodSon, p. & oSMólSKa, H.
(Eds.): The Dinosauria (2
nd
edition): 259-322; Berkeley
(University of California Press).
VerMa, o. (2015): Cretaceous vertebrate fauna of the Cau-
very Basin, southern India: Palaeodiversity and palaeo-
biogeographic implications. – Palaeogeography, Palaeo-
climatology, Palaeoecology, 431: 5 3 - 67.
V
erMa
, o., K
hoSla
, a., G
oin
, f.j. & K
aUr
, J. (2016): Histori-
cal biogeography of the Late Cretaceous vertebrates of
India: Comparisons of geophysical and paleontological
data. – Bulletin of the New Mexico Museum of Natural
History and Science, 71: 317-33 0.
whyte, M.a. & roMano, M. (1993): Footprints of a sau-
ropod dinosaur from the Middle Jurassic of Yorkshire.
– Proceedings of the Geologists’ Association, 104: 195 -
199.
w
hyte
, M.a. & r
oMano
, M. (1995): Probable sauropod foot
-
prints from the Middle Jurassic of Yorkshire, England.
– Gaia, 10: 15-26.
whyte, M.A. & RoMano, M. (2001): Probable stegosaurian
dinosaur tracks from the Saltwick Formation (Middle
Jurassic) of Yorkshire, England. – Proceedings of the
Geologists’ Association, 112: 45-5 4.
W
illiSton
, S.w. (1903): North American plesiosaurs. Part
1. – Geological Series of the Field Columbian Museum,
2 (1) : 1-7 7.
W
ilSon
, j.a., b
arrett
, p.M. & c
arra no
, M.t. (2 011): A n
associated partial skeleton of Jainosaurus cf. septen-
trionalis (Dinosauria: Sauropoda) from the Late Creta-
ceous of Chhota Simla, central India. – Palaeontology,
54: 981-998.
x
inG
, l., l
ocKley
, M.G., M
c
c
rea
, r.t., G
ierlinSKi
, G.d.,
bUcKley, l.G., zhanG, j., qi, l. & jia, C. (2013): First
record of Deltapodus tracks from the Early Cretaceous
of China. – Cretaceous Research, 42: 55-65.
yadaGiri, p. & ayyaSaMi, K. (1979): A new stegosaurian di-
nosaur from Upper Cretaceous sediments of south India.
– Journal of the Geological Society of India, 20: 521-530.
yadaGiri, p. & ayyaSa Mi, K. (1987): A carnosaurian dinosaur
from the Kallamedu Formation (Maastrichtian horizon),
Tamil Nadu. – Geological Survey of India, Special Pub-
lications, 11: 523-528.
Manuscript received: December 18th, 2016.
Revised version accepted by Stuttgart editor: May 14th, 2017.
Addresses of the authors:
peter M. Galton, University of Bridgeport, Bridgeport,
Connecticut, USA; Curatorial Affiliate, Peabody Museum
of Natural History, Yale University, New Haven, Connecti-
cut, USA. Present address: 1065 Vintage Drive, Rio Vista,
California 94571, USA;
e-mail: pgalton@bridgeport.edu
K
riShnan
a
yyaSaMi
, formerly of the Palaeontology Division,
Geological Survey of India, Hyderabad 500068, India;
e-mail: ayyasami@gmail.com