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Morphological comparisons of metacarpal I for Australovenator wintonensis and Rapator ornitholestoides: implications for their taxonomic relationships

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Various comparisons of left metacarpal I of the Australovenator wintonensis holotype have been made with Rapator ornitholestoides. These specimenswere identified as being morphologically more similar than either was to that of the neovenatorid Megaraptor namunhuaiquii. Owing to thepoor preservation of A. wintonensis and R. ornitholestoides, distinct morphological separation between the two appeared minimal. The recent discovery of a near perfectly preserved right metacarpal I of A. wintonensis enables a direct and accurate comparison with R. ornitholestoides. Distinctmorphological differences exist between the metacarpals of the two species. A re-evaluation of the age of the A. wintonensis holotype site (AODL85 ‘Matilda Site’) with zircon dating reveals a maximum age of 95 Ma, 10 Ma younger than the Griman Creek Formation at Lightning Ridge,from which R. ornitholestoides was recovered. This age difference detracts from the probability that the specimens belong to the same genus.
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Alcheringa: An Australasian Journal of Palaeontology
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Morphological comparisons of metacarpal I
for Australovenator wintonensis and Rapator
ornitholestoides: implications for their taxonomic
relationships
Matt A. White, Peter L. Falkingham, Alex G. Cook, Scott A. Hocknull & David
A. Elliott
To cite this article: Matt A. White, Peter L. Falkingham, Alex G. Cook, Scott A. Hocknull & David
A. Elliott (2013) Morphological comparisons of metacarpal I for Australovenator wintonensis
and Rapator ornitholestoides: implications for their taxonomic relationships, Alcheringa: An
Australasian Journal of Palaeontology, 37:4, 435-441, DOI: 10.1080/03115518.2013.770221
To link to this article: http://dx.doi.org/10.1080/03115518.2013.770221
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Morphological comparisons of metacarpal I for
Australovenator wintonensis and Rapator ornitholestoides:
implications for their taxonomic relationships
MATT A. WHITE, PETER L. FALKINGHAM, ALEX G. COOK, SCOTT A. HOCKNULL and DAVID A.
ELLIOTT
WHITE, M.A., FALKINGHAM, P.L., COOK, A.G., HOCKNULL, S.A. & ELLIOTT, D.A., 2013. Morphological comparisons of metacarpal I for
Australovenator wintonensis and Rapator ornitholestoides: implications for their taxonomic relationships. Alcheringa 37, 435441. ISSN 0311-
5518.
Various comparisons of left metacarpal I of the Australovenator wintonensis holotype have been made with Rapator ornitholestoides. These speci-
mens were identied as being morphologically more similar than either was to that of the neovenatorid Megaraptor namunhuaiquii. Owing to the
poor preservation of A. wintonensis and R. ornitholestoides, distinct morphological separation between the two appeared minimal. The recent dis-
covery of a near perfectly preserved right metacarpal I of A. wintonensis enables a direct and accurate comparison with R. ornitholestoides. Distinct
morphological differences exist between the metacarpals of the two species. A re-evaluation of the age of the A. wintonensis holotype site (AODL
85 Matilda Site) with zircon dating reveals a maximum age of 95 Ma, 10 Ma younger than the Griman Creek Formation at Lightning Ridge,
from which R. ornitholestoides was recovered. This age difference detracts from the probability that the specimens belong to the same genus.
Matt A. White[fossilised@hotmail.com], School of Engineering, The University of Newcastle, Callaghan, New South Wales 2308, Australia; Peter
L. Falkingham[pfalkingham@rvc.ac.uk], Department of Comparative Biomedical Sciences, Structure and Motion Laboratory, Royal Veterinary
College, London, UK; Alex G. Cook [alex.cook@y7mail.com] and Scott A. Hocknull [scott.hocknull@qm.qld.gov.au], Ancient environments,
Queensland Museum, Hendra, Queensland, 4011, Australia; David A. Elliott [david.elliott@aaod.com.au], Australian Age of Dinosaurs Museum
of Natural History, The Jump Up, Winton, Queensland, 4735, Australia.
Also afliated with Australian Age of Dinosaurs Museum of Natural His-
tory, The Jump Up, Winton, Queensland 4735, Australia. Also afliated with Department of Ecology and Evolutionary Biology, Division of Biol-
ogy and Medicine, Brown University, USA. Received 22.9.2012; revised 13.1.2013; accepted 17.1.2013.
Key words: Australovenator wintonensis,Rapator ornitholestoides, metacarpal, theropods.
AUSTRALIAN THEROPOD DINOSAUR fossils are
rare with only ve taxa so far formally recognized, and
most based on isolated fragmentary remains. These
include: Walgettosuchus woodwardi Huene, 1932;
Kakuru kujani Molnar & Pledge, 1980; Rapator orni-
tholestoides Huene, 1932; Timimus hermani Rich &
Vickers-Rich & Vickers-Rich, 1994; and Australoventa-
tor wintonensis Hocknull et al. 2009 (Agnolin et al.
2010). Isolated theropod postcranial bones have also
been recovered from the Early Cretaceous, Flat Rocks
locality in southern Victoria. These were identied as
pertaining to allosauroids, tyrannosauroids, ceratosauri-
ans, ornithomimosaurians, a basal coelurosaur and at
least three indeterminate maniraptoran taxa (Benson
et al. 2012).
Australias most complete theropod dinosaur, A.
wintonensis, was discovered interspersed with remains
of the sauropod Diamantinasaurus matildae Hocknull
et al. 2009 at the type locality AODL 85 Matilda
Sitenear Winton in central-western Queensland, Aus-
tralia (Hocknull et al. 2009). Australovenator winton-
ensis was later placed within a new family-level
clade, Neovenatoridae, which also included Neovena-
tor salerii Hutt et al., 1996, Aerosteon riocoloradensis
Sereno et al., 2008, Chilantaisaurus tashuikouensis
Hu, 1964, Fukuiraptor kitadaniensis Azuma & Curry,
2000, Megaraptor namumhuaiquii Novas, 1998 and
Orkoraptor burkei Novas et al., 2008 (Benson et al.
2010).
The initial description of A. wintonensis incorporated
a comparison of its poorly preserved left metacarpal I
with the same element of R. ornitholestoides (Hocknull
et al. 2009). This revealed that metacarpal I of A.
wintonensis lacks a prominent caudomedial process pro-
jecting proximo-dorsally, but possesses subequal distal
condyles and a at proximal articular surface, which
differ from the concave surface form evident in R.
ornitholestoides; the straight distal lateral condyle also
contrasts with the disto-laterally projecting distal
lateral condyle in R. ornitholestoides (Hocknull et al.
2009).
Reanalysis of the R. ornitholestoides type specimen
(NHMUK R3718) by Holtz et al. (2004), resulted in a
revised placement with alvarezsaurids. Agnolin et al.
(2010) alternatively identied two critical tetanuran
characteristics: a lateral articular facet that extends well
Ó2013 Association of Australasian Palaeontologists
http://dx.doi.org/10.1080/03115518.2013.770221
Alcheringa: An Australasian Journal of Palaeontology 2013.37:435-441.
proximo-distally; and a deep proximal articular surface
that articulates with the distal carpal. Rapator ornithol-
estoides was, thus, specically excluded from Coeluros-
auria based on the rounded medial margin on its
metacarpal I; this differs from the typical sharp-edged
condition in coelurosaurians (Rauhut & Xu 2005,
Agnolin et al. 2010). Agnolin et al. (2010) further
concluded that R. ornitholestoides was virtually
identical to both A. wintonensis and the South American
theropod Megaraptor namumhuaiquii and was, there-
fore, probably an indeterminate megaraptoran, rendering
R. ornitholestoides anomen dubium. The most recent
assessment by Carrano et al. (2012) agreed with this
proposal but could not rene the placement of R. orni-
tholestoides beyond Neovenatoridae indet., because
metacarpal I is absent in N. salerii.
New skeletal elements of the holotype specimen of
A. wintonensis (AODF 604) have been found at the ori-
ginal AODL 85 locality near Winton, central western
Queensland (White et al. 2012). Of particular interest is
a complete right metacarpal I whose exceptional preser-
vation has revealed novel features not observable on the
only other known specimen (see Hocknull et al. 2009,
Fig. 27AF; White et al., 2012, Fig. 11). This discovery
has enabled more accurate comparisons with R. orni-
tholestoides and critical determination of whether
A. wintonensis and R. ornitholestoides are indeed
morphologically compatible and/or belong to the same
genus.
Stratigraphy, palaeoenvironment and
age
Vertebrate fossils from the opal elds of Lightning
Ridge, New South Wales, including the only known
specimen of R. ornitholestoides, all derive from the
Wallangulla Sandstone Member of the Griman Creek
Formation (Byrnes 1977, Molnar 1980, Molnar &
Galton 1986, Dettmann et al. 1992). Sedimentology and
regional stratigraphy indicates that this unit was most
likely a uviatile freshwater to estuarine deposit, with
rivers owing westward into an epicontinental sea
(Byrnes 1977, Molnar 1980, Molnar & Galton 1986,
Dettmann et al. 1992). Palynomorphs from the Griman
Creek Formation are attributable to the Australian paly-
nological [AP] unit APK5 (Burger 1980, 1992, Price
1997), which indicates a middle Albian age (ca 105
million years).
The holotype of A. wintonensis (AODF 604) was
excavated from the lowermost part of the Winton
Formation, which is the youngest subdivision of the
mid-Cretaceous Rolling Downs Group. The lowermost
beds of the Winton Formation are interpreted to
constitute a proximal deltaic to coastal plain uvial suc-
cession, developed following regression of an
epicontinental seaway during the latest Albian. Abun-
dant macrooral and microoral (McLoughlin et al.
1995, Price 1997, Pole & Douglas 1999, Dettmann &
Clifford 2000, Pole 2000, Clifford & Dettmann 2005,
Dettmann et al. 2009, McLoughlin et al. 2010)
evidence suggests a latest Albian to Cenomanian age,
Fig. 1. Stratigraphy of the Surat and northern Eromanga basins correlated with the palynozonation of Price (1997). The Surat Basin includes the
Griman Creek Sandstone, which preserved the Rapator specimen. The northern Eromanga Basin includes the Winton Formation, which preserved
Australovenator wintonensis.
436 MATT A. WHITE ET AL. ALCHERINGA
Alcheringa: An Australasian Journal of Palaeontology 2013.37:435-441.
conforming to the AP units APK6 (uppermost) and
APK7. The A. wintonensis type stratum was interpreted
to be latest Albian [(Dettmann & Clifford 2000) corre-
sponding to the uppermost APK6 (White et al. 2012)].
Recent zircon dating of the A. wintonensis type site
AODL 85 Matilda Site(Fig. 1) inferred a Cenomanian
age ca 95 Ma (Greentree 2011), thus making this partic-
ular stratum around 10 million years younger than the
Lightning Ridge deposits.
Materials and Methods
Computed tomography
The A. wintonensis metacarpal I was scanned for
computed tomography (CT) at Queensland Xray, Mac-
kay Mater Hospital, central eastern Queensland using
a Philips Brilliance CT 64-slice machine. Mimics
version 10.01 was used to view all CT scans, take
Fig. 2. Rapator ornitholestoides NHMUK R3718. The holotype left metacarpal I specimen of Rapator ornitholestoides in photographed and
computer-rendered views: A, Dorsal photograph; B, Dorsal render; C, Medial photograph; D, Medial render; E Ventral photograph; F, Ventral
render; G, Lateral photograph; H, Lateral render; I, Proximal photograph; J, Proximal render; K, Distal photograph; L, Distal render. Scale bar =
5 cm.
ALCHERINGA THEROPOD METACARPAL 437
Alcheringa: An Australasian Journal of Palaeontology 2013.37:435-441.
measurements and create a three-dimensional mesh.
This was imported into the graphic design package
Rhinoceros 4.0, which rendered a high-denition sur-
face model.
Photogrammetry
The R. ornitholestoides metacarpal (NHMUK R3718)
was digitized using the freely available open source
photogrammetry software bundler (Snavely et al. 2006,
2007) and CMVS/PMVS (Furukawa et al. 2010, Furuk-
awa & Ponce 2010) as outlined by Falkingham (2012).
One hundred and twenty-eight photographs were taken
using a Canon G11 digital camera (10 mp) to compile a
360° digital reconstruction of the specimen. Separate
models were produced of the dorsal and ventral surfaces
(using 78 and 50 photographs respectively), maintaining
considerable overlap at the sides. These were generated
using an 8-core workstation with 32GB RAM, then
scaled and aligned in the free software Meshlab. The
resultant complete point cloud (consisting of the merged
dorsal and ventral models) comprised 941 025 points,
and was then surfaced using the Poisson Surface
Reconstruction lter in Meshlab.
Institutional abbreviations
AODF, Australian Age of Dinosaurs Fossils; AODL,
Australian Age of Dinosaurs Locality; NHMUK, Natu-
ral History Museum of United Kingdom
Morphological comparisons
To assist with our analysis, the newly discovered right
metacarpal I of A. wintonensis (AODF 604) was mir-
rored onto the corresponding left metacarpal I of R.
ornitholestoides (NHMUK R3718: see Figs 2, 3). The
following observational perspectives follow the order of
gured specimens (Fig. 4).
Dorsal view
The dorso-lateral process is not as proximally posi-
tioned in A. wintonensis as in R. ornitholestoides, creat-
ing a shallower dorso-proximal articular facet. The
proximo-medial process in R. ornitholestoides appears
slightly broader. Medially, the A. wintonensis metacar-
pal is bowed more laterally than in R. ornitholestoides.
The lateral side is near straight from directly proximal
of the distal condyle to the proximal end of the dorso-
lateral process, whereas R. ornitholestoides has a
slightly bowed shaft medially portraying the A. winton-
ensis specimen to be slightly broader. The medial con-
dyle in A. wintonensis is splayed more medially than in
R. ornitholestoides. Proximo-distally, the medial con-
dyle of A. wintonensis is orientated at around 60°,
whereas in R. ornitholestoides it is more proximo-dis-
tally orientated with a 40° medial orientation. The inter-
condyle region separating the medial and lateral
condyles is shallow in A. wintonensis, whereas it is dee-
ply concave in R. ornitholestoides. A pronounced
rounded ridge occurs directly proximal to the medial
condyle of R. ornitholestoides but is absent in A. win-
tonensis (Fig. 4AD).
Ventral view
The ventral lateral process is sharply bowed medially at
the proximal end of A. wintonensis, whereas it is more
shallowly bowed in R. ornitholestoides. However, this
might be an artefact of poor preservation. The medial
condyle in A. wintonensis is orientated medially in com-
parison with the more proximo-distal orientation it
occupies in R. ornitholestoides (Fig. 4EH).
Medial view
Both specimens are quite similar in medial view, the
only distinct difference being that the proximo-central
process appears rounded in A. wintonensis, whereas it
appears atter on the ventral side in R. ornitholestoides.
This is probably due to a portion of the specimen not
being preserved. Additionally, the dorso-lateral process
is positioned more proximally in R. ornitholestoides
(Fig. 4IL).
Lateral view
In lateral view, A. wintonensis has a proximo-medial
ridge, which tapers into the proximal articular facet.
Fig. 3. Metacarpal I specimens of Australovenator wintonensis AODF
604 and Rapator ornitholestoides NHMUK R3718. A, Mirror image
of the right metacarpal I of A. wintonensis;B,Rapator
ornitholestoides left metacarpal I. Scale bar = 5 cm.
438 MATT A. WHITE ET AL. ALCHERINGA
Alcheringa: An Australasian Journal of Palaeontology 2013.37:435-441.
Fig. 4. Morphological comparisons between metacarpal I specimens of Australovenator wintonensis and Rapator ornitholestoides. Computer
generated renders, morphological traces and described characteristics of a right metacarpal I of Australovenator mirror imaged to a left metacarpal
I and re-scaled to compare with the left Rapator metacarpal I. Computer render of Australovenator in: A, dorsal; E, ventral; I, medial; M, lateral;
Q, distal; U, proximal views. Computer render of Rapator in: B, dorsal; F, ventral; J, medial; N, lateral; R, distal; V, proximal views.
Morphological trace of Australovenator in: C dorsal; G, ventral; K medial; O, lateral; S, distal; W, proximal views. Morphological trace of Rapator
in: D, dorsal; H, ventral; L, medial; P, lateral; T, distal; X, proximal views. Morphological characteristics; 1, dorso-lateral process; 2, dorso-
proximal articular facet; 3, proximo-medial process; 4, dorsal medio-lateral bow; 5, lateral face; 6, dorsal view of medial condyle; 7, dorsal view of
intercondyle region; 8, region proximal to medial condyle; 9, ventro-lateral process; 10, ventral medio-lateral bow; 11, ventral view of lateral
condyle; 12, ventral view of medial condyle; 13, proximo-medial process; 14, proximo-central ridge; 15, articular facet for metacarpal II; 16,
medial condyle; 17, lateral condyle; 18, distal view of intercondyle region; 19, proximal articular facet; 20, ventro-medial margin; 21, dorsal
surface.
ALCHERINGA THEROPOD METACARPAL 439
Alcheringa: An Australasian Journal of Palaeontology 2013.37:435-441.
The ridge separates two condyles of distal carpal-1.
Rapator ornitholestoides lacks this proximo-medial
ridge. Both specimens have distinctly different lateral
articular facets. The articular facet of A. wintonensis is
broader and extends more proximo-distally implying its
articulation with metacarpal II extends to a greater por-
tion of metacarpal I than in R. ornitholestoides where
the articulation is predominantly constrained to the
proximal half (Fig. 4MP).
Distal view
It is difcult to accurately compare the distal condyles
owing to the poor preservation of R. ornitholestoides,
however the basic outline demonstrates some distinct
differences. The medial condyle is taller than the lateral
condyle in A. wintonensis but in R. ornitholestoides
they are equal in height. Ventrally, the condyles are
more distinctly separated in R. ornitholestoides than in
A. wintonensis. The medial condyle in A. wintonensis is
angled around 20°, whereas the Rapator condyle is
more dorso-ventrally orientated. The lateral condyle in
A. wintonensis has a distinct medial bow, whereas it is
straighter in R. ornitholestoides (Fig. 4QT).
Proximal view
The articular facets of the proximal ends of both speci-
mens are quite different. Australovenator wintonensis
has a prominent medial ridge on the articular surface
that separates the condyles on distal carpal-1 and a ven-
tro-medial ridge, both of which are absent in R. ornithol-
estoides. The dorsal surface is more distinctly bowed in
R. ornitholestoides emphasizing a more elongate dorso-
lateral process compared with A. wintonensis. These fea-
tures indicate distinct differences between the distal car-
pal-I bones of both specimens (Fig. 4UX).
Conclusions
The discovery of a near perfectly preserved right meta-
carpal I belonging to the holotype (AODF604) of Aus-
tralovenator wintonensis, has enabled a detailed
comparative reappraisal of the structurally similar taxon
Rapator ornitholestoides (NHMUK R3718). This pro-
vides an advantage over previous evaluations (e.g.,
Agnolin et al. 2010), which were based on the incom-
plete left metacarpal I described by Hocknull et al.
(2009). Key features that Agnolin et al. (2010) consid-
ered to be diagnostic are here shown to be products of
post-mortem damagee.g., the at proximal articular
surface and sub-equal distal condyles. Furthermore,
Agnolin et al. (2010) concluded that there was minimal
morphological variation between R. ornitholestoides, A.
wintonensis and the South American taxon Megaraptor
namumhuaiquii; although R. ornitholestoides and A.
wintonensis were interpreted to be structurally more
compatible than either was to M. namumhuaiquii. How-
ever, our assessment of these taxa identied sufcient
differences to warrant assignment to separate genera.
Moreover, the forms of the articular facets of distal car-
pal 1, metacarpal II and manual phalanx I-1 are clearly
different. Finally, clear temporal separation of R. orni-
tholestoides and A. wintonensis by at least 10 million
years demonstrates stratigraphical distinction, and sug-
gests that other theropod material from Lightning Ridge
should reveal contrasting taxonomic elements.
Acknowledgements
We gratefully thank Sarah Woolridge, Ian Pengelly and
Queensland Xray staff in Mackay for their assistance in
CT scanning specimens. We thank P. Hurst from Image
Resources at the Natural History Museum, London for
the Rapator specimen photographs. We thank Sandra
Chapman (NHMUK) for access to the Rapator
specimen. We thank Maxine Macmillan for the
excellent preparation of right metacarpal I of
Australovenator. Special thanks to Stephen
McLoughlin, Roger Benson, Fernando Novas and Ben
Kear for their useful suggestions and criticisms to
improve this paper. We thank Colin McHenry for
assistance with Rhinoceros 4.0 software. We thank the
following people for their help, encouragement and
support: Judy Elliott, Brant Bassam, Tricky Sloan,
George Sinapius. We thank the following organizations
for their help and support; Queensland Xray Mackay,
Australian Age of Dinosaurs Natural History Museum,
Queensland Museum, Natural History Museum of
United Kingdom and the University of Newcastle.
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Alcheringa: An Australasian Journal of Palaeontology 2013.37:435-441.
... Cessation of sedimentation in and the onset of uplifting of the Surat and Eromanga Basins in the late Early Cretaceous is currently hypothesised to have led to the formation of opal beds in many areas of the GAB through erosion and oxidation of volcaniclastic sediments deposited between 130-95 Mya into in a cold, oxygen-deprived fluvial-deltaic environment (Rey, 2013). The Griman Creek Formation at Lightning Ridge arguably contains the most abundant fossil record of Cretaceous terrestrial fauna in Australia (Dettmann et al., 1992), with crocodylomorphs (Etheridge, 1917;Molnar, 1980;Molnar & Willis, 2000), australosphenidian mammals (Archer et al., 1985;Rich, Flannery & Archer, 1989;Flannery et al., 1995), ornithopod dinosaurs (Molnar & Galton, 1986), megaraptoran theropods (White et al., 2013;Bell et al., 2015), enantiornithine birds (Molnar, 1999), plesiosaurs (Kear, 2006a), turtles (Smith, 2010;Smith & Kear, 2013), dipnoan lungfish (Kemp & Molnar, 1981;Kemp, 1993;Kemp, 1997) and a possible synapsid (Clemens, Wilson & Molnar, 2003)in addition to numerous species of non-marine macro-invertebrates (Byrnes, 1977;Hocknull, 2000;Kear & Godthelp, 2008;Hamilton-Bruce & Kear, 2010) and plants. Preservation of fossils at Lightning Ridge—including those specimens described here—is commonly in the form of natural casts, or pseudomorphs, in non-precious opal (e.g.,Molnar & Willis, 2000;Clemens, Wilson & Molnar, 2003;Bell et al., 2015). ...
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