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Purported latest bone of a plated dinosaur (Ornithischia: Stegosauria), a

"dermal plate" from the Maastrichtian (Upper Cretaceous) of southern India

Article in Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen · July 2017
DOI: 10.1127/njgpa/2017/0671

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E N. Jb. Geol. Paläont. Abh. 285/1 (2017), 241–000
Stuttgart, July 2017
Article

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,

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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

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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
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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
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as indicated by a pes print of Deltapodus sp., a stegosaurian ichnotaxon.

Key words: Dinosauria, Ornithischia, Stegosauria, Dravidosaurus, Deltapodus, Upper Cretaceous,

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India, bones, footprints.

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1. Introduction mation (uppermost Upper Cretaceous, Maastrichtian).

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The site was in the Cauvery Basin near Kallamedu in

Stegosauria is a clade of quadrupedal, graviportal, the Tiruchirapalli district, Tamil Nadu, southern India.
herbivorous ornithischian dinosaurs with an array of The bones included those interpreted as stegosaurian
dermal plates and spines in two parasagittal rows that and a large dermal plate was illustrated. Subsequently,
extend along the top of the body. The group is best rep- Yadagari & Ayyasami (1979: 529) noted the occurrence
resented by partial to complete skeletons and isolated of theropod, sauropod and stegosaur bones from above
bones from the Middle and Late Jurassic but there are a layer containing a lower Maastrichtian foraminiferal
only a few records from the Early Cretaceous (Galton fauna (Pachydiscus otacodensis Zone, Yadagari &
& Upchurch 2004; Maidment et al. 2008; Maidment Ayyasami 1979, tab. 1, fig. 523, map). The very large
2010; Galton 2012; Pereda-Suberbiola et al. 2012, bones, which came from this excavation site north east
2015; Borinder et al. 2016). The most recent occur- of Kallamadu Village, were described as the theropod
rence for stegosaurs is from the Late Cretaceous of Bruhathkayosaurus matleyi Yadagiri & Ayyasami,
India. 1987. However, this taxon is listed as a nomen dubi-
Anonymous (1978) reported the discovery of bones um, Sauropoda indet., by Upchurch et al. (2004; also
excavated during a geological survey by Ponnala Yad- Hone et al. 2016) and, as one of us (Ayyasami) notes,
agari and K rishnan Ayyasami of the Kallamedu For- only bones of this taxon were found in situ and none

©2017 E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany www.schweizerbart.de

DOI: 10.1127/njgpa/2017/0671 0077-7749/2017/0671 $ 0.00
2 P.M. Galton and K. Ayyasami

of stegosaurs. New vertebrate remains, including dino- open texture (Fig. 1G; Buffrénil et al. 1986; Main et
saur teeth, are reported from the Kallamadu Formation al. 2005; Hayashi et al. 2012), would not be preserved
(Prasad et al. 2013; Goswami et al. 2013) and the verte- without the protective outer cortex. This is also true for
brate fauna of the Maastrichtian of the Cauvery Basin the dermal armor of ankylosaurs (Hayashi et al. 2010).
is discussed by Verma (2015; also Verma et al. 2016). Hulke (1881, pl. 76, fig. 1) figured a large lateral spike
of the nodosaurid ankylosaur Polacanthus foxii (Lower
Previously, the partial skull and postcranial skel-
Cretaceous, England) that is represented by a partial
eton of the stegosaur Dravidosaurus blanfordi Yada-
mostly hollow thin shell of outer cortex with most of
gari & Ayyasami, 1979 was described from the older
the internal cancellous bone eroded away.
marine limestone beds in the Trichinopoly Group (Co-
Apart from being sub-triangular in outline in
niacian, Kosmaticeras theobaldianum Zone), also in
“side” views (Fig. 1A, D), this bone does not resemble
the Cauvery Basin west of Siranattam Village in the
any of the dermal plates of the Morrison (Upper Ju-
Tirachirapalli district of southern India (see Yada-
rassic) stegosaurs Stegosaurus stenops (see Gilmore
gari & Ayyasami 1979, tab. 1, fig. 1). Galton (1981)
1914; Maidment et al. 2015) and Hesperosaurus mjosi
re-identified some of the cranial bones of Dravidos-
(Carpenter et al. 2001; Carpenter 2010; Saitta 2015
aurus blanfordi and mistakenly cited the plate figured
as Stegosaurus mjosi), or those of any other stegosaur
in Anonymous (1978) as belonging to this taxon (see
or of any ankylosaur (see skeletal restorations in Paul
Section 3 for further details on Dravidosaurus).
2016). This is especially true for the dermal armor of
The citations by Galton (1981; also 1990: 435, 2012: titanosaurian sauropods (see Matley 1924, pls. 12, 13;
296; Galton & Upchurch 2004: 343; Pereda-Suberbio- also not equivalent to any of the four types of armor

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la et al. 2012: 75) for the occurrence of stegosaur bones that occur in titanosaurs, see d’Emic et al. 2009).

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in the Maastrichtian of India, the latest record of the Unlike thyreophoran dermal plates, this bone
group, is based on the claim by Yadagari & Ayyasami (ca. 175 mm long close to base, ca. 190 mm high, ca.
(1979: 529) given above. However, the only informa- 65 mm at thickest) does not consist of cancellous bone
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tion provided to date on this material is a photograph with large vascular spaces Fig. 1A-E cf. Fig. 1F) or
of the dermal plate in Anonymous (1978). Owing to taper from base to apex in “edge-on” views (Fig. 1B, C
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the potential importance of such a geologically young cf. Fig. 1A). There are no vertical vascular grooves on
stegosaur, a more thorough description is long overdue. the outer surface of the cortex, as occur in Morrison
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stegosaurs (see Gilmore 1914; Ostrom & McIntosh

1999; Galton 2010), because this layer is completely
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Institutional abbreviations: GSIG, Geological Survey of missing. The only smooth surfaces are those of unpre-
India, Western Region Gandhinagar, Gujarat, India; GSIH,
pared matrix (m, Fig. 1B-E). The bone surfaces are all
Geological Survey of India, Southern Region, Hyderabad,
co

India; and YPM VP, Division of Vertebrate Paleontology, very rough and irregular with no trace of an externally
Peabody Museum of Natural History, Yale University, New smooth thin outer layer of compact bone. This is not
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Haven, Connecticut, USA. 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
2. Description and comparisons an eroded piece of compact bone that, based on its size,
was probably derived from a long bone of a sauropod
Dermal armor occurs in three groups of dinosaurs that dinosaur. In this context it is interesting that Hone et
are possibly represented in the Upper Cretaceous of al. (2016: 7) cite Bruhathkayosaurus (from Kallamedu
India: stegosaurs (Yadagiri & Ayyasami 1979; Galton Formation near Kallamedu) as an anomalously large
1981), ankylosaurs (Chatterjee & Rudra 1996: 518; sauropod dinosaur (Yadagiri & Ayyasami 1987: ilium
undescribed vertebrae, scapulocoracoid, humerus, fe- length 1200 mm, femur distal condylar width 750 mm,
mur, osteoderms), and titanosaurian sauropods (the tibia length 2000 mm).
armor of “Lametosaurus indicus” of Matley 1924,
pls. 12, 13 that is not ankylosaurian, see Carrano &
Sampson 2008: 208). Using the dermal plates of Stego-
saurus as a comparative standard, the plates consist 3. Discussion
of a thin layer of compact bone enclosing cancellous With reference to the Kallamedu Formation (Upper
bone, with large vascular spaces that, because of its Cretaceous, Maastrichtian) near Kallamedu, Tiruchi-
 Purported latest bone of a plated dinosaur (Ornithischia: Stegosauria) 3

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Fig. 1. A-E – Alleged stegosaur dermal plate originally figured by Anonymous (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).
4 P.M. Galton and K. Ayyasami

rapalli district in the Cauvery Basin of southern India, with long root ca 2.5 mm; Yadagari & Ayyasami 1979,
Blanford (1864: 139) noted that the beds consisted of a fig. 4, pl. 1, fig. 3a, b). However, one of us (K.A.) is cur-
mass of white sands and grey sandy clays and “imbed- rently working on undescribed bones that support the
ded in the deposits large bones are numerous, but so presence of a stegosaur in the type quarry in the Conia-
saturated with water and so very friable, that it is im- cian (Upper Cretaceous) of southern India.
possible, even with the greatest care, to extract them in An isolated impression of a small right pes (225
anything like a recognizable condition.” Matley (1929) mm long), the basis for a plaster cast (Fig. 1F), was
noted that this was the situation during the monsoon found in the uppermost limestone unit of the Lameta
season whereas during the dry season desiccation, with Formation (given as Infratrappean Sequence; Maas-
expansion by day and contraction at night, results in trichtian) of the Jetholi area near Balasinor in Kheda
the bones being split into fragments. Consequently, District in Gujarat, western India (see Mohabey 1986
the bones are poorly preserved and the photographs for photos and map). This pes print was referred to the
used for Bruhathkayosaurus by Yadagari & Ayyasami stegosaurian ichnotaxon Deltapodus sp. by Mateus et
(1987) were taken while the bones were still exposed al. (2011). Deltapodus brodricki Whyte & Romano,
in situ. The bones started to disintegrate in the field 1995, which was based on a supposed sauropod track
jackets even before reaching the GSIH and no longer from the Middle Jurassic of England, was re-identified
exist. This is in marked contrast to the “dermal plate” as representing the prints of a three toed stegosaur like
(Fig. 1A-E) that, even though lacking a resistant outer Stegosaurus by Whyte & Romano (2001, tab. 1 for de-
cortex, is still very well preserved. However, one of tails; for photographs in situ as preserved, see Whyte &
us (K.A.) was present when Panalla Yadagiri located Romano 1993, fig. 3 for holotype and Lomax & Tamura

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this isolated piece of bone near the excavation site as 2014, figs. 130A, B, 131A, B for other examples). The

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float in the bed of a stream, a tributary of the Maru- identification of Deltapodus brodricki as a stegosaurian
daiyar River (Yadagari & Ayyasami 1979, fig. 1, map). footprint is supported by subsequent studies as cited in
Consequently, it was not found in situ with the other Galton (2017). The Indian pes print of Deltapodus sp.
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bones that, as Ayyasami notes, did not include any of (Fig. 1F) agrees with those of Deltapodus brodricki
stegosaurs. This is extremely important because some- (see Whyte & Romano 2001, fig. 3A, tab. 1; Galton
ed

where upstream to this site there should be a horizon 2017, fig. 2N) in several characters, viz., it is triangular
with very well preserved large bones, something still with a well-developed heel, digitigrade, three very wide
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to be discovered in the Maastrichtian of southern India. blunt digits, toes radiating and not separated by well-
As regards to Dravidosaurus blanfordi, Chatterjee developed hypicies (angles), and weakly mesaxonic
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& Rudra (1996: 518) noted that “we visited the site (digit III only slightly longer than II and IV) (Whyte
and found only fragmentary remains of plesiosaurs. We & Romano 2001; Li et al. 2012). Deltapodus also oc-
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also examined the holotype and could not see anything curs in the Lower Cretaceous of Spain (Cobos et al.
related to the stegosaurian plates and skull claimed by 2010; Pascual et al. 2012), western USA (Milân et al.
these authors. Instead, the bones are highly weathered
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2015) and China (Xing et al. 2013). As Mateus et al.

limb and girdle elements and may belong to plesio- (2011: 656) note, the occurrence of skeletal remains
saurs.” This was cited by Maidment et al. (2008; also of a stegosaur from the Upper Cretaceous (Coniacian)
Maidment 2010) who, until the bones are independently of southern India would be in accord with the record
re-described, regard this taxon as a nomen dubium and of the stegosaurian Deltapodus-like pes print from the
the specimen as Stegosauria indet. (but as ?Stegosauria Maastrichtian of western India.
indet., Pereda-Suberbiola et al. 2015; not demonstrably
ornithischian, Wilson et al. 2011: 982). Verma (2015:
57) lists this taxon as a marine reptile and it is given
as a taxon of plesiosaur by Verma et al. (2016: 320). Acknowledgements
However, there is no correspondence between the pho- PMG thanks D. Mohabey (Nagpur University, Nagpur, India)
tographs of the bones of Dravidosaurus in Yadagari for a copy of the original photograph (Fig. 1F) and for try-
& Ayyasami (1979, pls. 1-4; also figs. 2-7) with figures ing to locate the cast of the pes in the GSIG collection. We
thank the reviewers P.M. Barrett (Natural History Museum,
of the limbs and girdles of a plesiosaur (cf. Andrews London, UK), K. Carpenter (University of Utah – Eastern,
1910, 1913; Williston 1903; O’Gorman et al. 2015). Price, Utah, USA, and J.I. Kirkland (Utah Geological Sur-
This is especially true for a small tooth (crown width vey, Salt Lake City, Utah, USA) for their constructive com-
and height ca 1 mm with three crenulations, total length ments that improved this paper.
 Purported latest bone of a plated dinosaur (Ornithischia: Stegosauria) 5

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very Basin, South India: implications for Gondwanan
paleobiogeography. – Journal of Vertebrate Paleontol- P eter M. Galton, University of Bridgeport, Bridgeport,
ogy, 33 (6): 1260-1268. Connecticut, USA; Curatorial Affiliate, Peabody Museum
Saitta, E.T. (2015): Evidence for sexual dimorphism in of Natural History, Yale University, New Haven, Connecti-
the plated dinosaur Stegosaurus mjosi (Ornithischia, cut, USA. Present address: 1065 Vintage Drive, Rio Vista,
Stegosauria) from the Morrison Formation (Upper Ju- California 94571, USA;
rassic) of western USA. – PloSONE, 10 (4): e0123503. e-mail: pgalton@bridgeport.edu
DOI:10.1371/journal.pone.0123503 Krishnan Ayyasami, formerly of the Palaeontology Division,
Upchurch, P., Barrett, P.M. & Dodson, P. (2004): Sauropo- Geological Survey of India, Hyderabad 500068, India;
da. – In: Weishampel, D.B., Dodson, P. & Osmólska, H. e-mail: ayyasami@gmail.com
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