Journal of Vertebrate Paleontology

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New megaraptorid (Dinosauria: Theropoda)

remains from the Lower Cretaceous Eumeralla
Formation of Cape Otway, Victoria, Australia

Stephen F. Poropat, Matt A. White, Patricia Vickers-Rich & Thomas H. Rich

To cite this article: Stephen F. Poropat, Matt A. White, Patricia Vickers-Rich & Thomas H. Rich
(2019) New megaraptorid (Dinosauria: Theropoda) remains from the Lower Cretaceous Eumeralla
Formation of Cape Otway, Victoria, Australia, Journal of Vertebrate Paleontology, 39:4, e1666273,
DOI: 10.1080/02724634.2019.1666273

To link to this article: https://doi.org/10.1080/02724634.2019.1666273

View supplementary material Published online: 10 Oct 2019.

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Journal of Vertebrate Paleontology e1666273 (19 pages)
© by the Society of Vertebrate Paleontology
DOI: 10.1080/02724634.2019.1666273

ARTICLE

NEW MEGARAPTORID (DINOSAURIA: THEROPODA) REMAINS FROM THE LOWER

CRETACEOUS EUMERALLA FORMATION OF CAPE OTWAY, VICTORIA, AUSTRALIA

STEPHEN F. POROPAT, *,1,2 MATT A. WHITE,2,3 PATRICIA VICKERS-RICH, 1,4,5 and THOMAS H. RICH5
1
Faculty of Science, Engineering and Technology, Swinburne University of Technology, John Street, Hawthorn, Victoria 3122,
Australia, sporopat@swin.edu.au; prich@swin.edu.au;
2
Australian Age of Dinosaurs Natural History Museum, The Jump-Up, Winton, Queensland 4735, Australia,
stephenfporopat@gmail.com; fossilised@hotmail.com;
3
School of Environmental and Rural Science, University of New England, Armidale, New South Wales 2351, Australia,
mwhite62@une.edu.au;
4
School of Earth, Atmosphere and Environment, Monash University, Melbourne, Victoria 3800, Australia, pat.rich@monash.edu;
5
Museum Victoria, P.O. Box 666, Melbourne, Victoria 3001, Australia, trich@museum.vic.gov.au

ABSTRACT—Megaraptorid theropods thrived in South America and Australia during the mid-Cretaceous. Their Australian
record is currently limited to the upper Barremian–lower Aptian upper Strzelecki Group and the upper Aptian–lower Albian
Eumeralla Formation of Victoria, the Cenomanian Griman Creek Formation of New South Wales, and the Cenomanian–
lowermost Turonian Winton Formation of Queensland. The latter has produced Australovenator wintonensis, the
stratigraphically youngest and most complete Australian megaraptorid. The Eric the Red West (ETRW) site on Cape
Otway, Victoria (Eumeralla Formation; lower Albian), has yielded two teeth, two manual unguals, and a right astragalus
that are almost identical to the corresponding elements in Australovenator. Herein, we classify these as Megaraptoridae cf.
Australovenator wintonensis. We also reappraise the ‘spinosaurid’ cervical vertebra from ETRW and suggest that it pertains
to Megaraptoridae. Three other theropod elements from ETRW—a cervical rib (preserving a bite mark), a caudal vertebra,
and a non-ungual manual phalanx—are also described, although it is not possible to determine their phylogenetic position
more precisely than Tetanurae (non-Maniraptoriformes). All elements were found in a fluvial deposit, associated with
isolated bones of other theropods, ornithopods, and turtles, amongst others; consequently, no two can be unequivocally
assigned to the same theropod individual. The new specimens from ETRW demonstrate that a megaraptorid theropod
morphologically similar to Australovenator lived during the late Early Cretaceous in Victoria, at a higher paleolatitude than
its northern counterpart. Moreover, they attest to the success of megaraptorids in late Barremian–early Turonian faunas
throughout eastern Australia.

SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

Citation for this article: Poropat, S. F., M. A. White, P. Vickers-Rich, and T. H. Rich. 2019. New megaraptorid (Dinosauria:
Theropoda) remains from the Lower Cretaceous Eumeralla Formation of Cape Otway, Victoria, Australia. Journal of
Vertebrate Paleontology. DOI: 10.1080/02724634.2019.1666273.

INTRODUCTION Barremian–Aptian Birdrong Sandstone (Long and Cruickshank,

1996) is an incorrectly restored elasmosaur cervical vertebra (and
The Australian Mesozoic theropod fossil record extends from
is in fact from the Aptian Windalia Sandstone Member of the
the Late Triassic to the Late Cretaceous (Table S1 in Supplemen-
Muderong Shale; M. Siversson, pers. comm., May 13, 2019),
tal Data). Although ichnological evidence indicates that thero-
whereas the ‘humerus’ from the Maastrichtian Miria Marl
pods were present in Queensland during the Late Triassic
(Long, 1992) appears to be a testudinatan scapula (M. Siversson,
(Thulborn, 2000) and Middle Jurassic (Thulborn, 1990; Molnar,
pers. comm., May 13, 2019). The South Australian record com-
1991), the only pre-Cretaceous body fossil evidence of theropods
prises only the partial holotype right tibia of Kakuru kujani
in Australia is the holotype tibia of Ozraptor subotaii from the
from the Aptian Bulldog Shale (Molnar and Pledge, 1980), now
Bajocian Colalura Sandstone of Western Australia (Long and
regarded as an indeterminate averostran (Agnolin et al., 2010)
Molnar, 1998). By contrast, body fossils and trace fossils of Cre-
or tetanuran (Barrett et al., 2010); the phalanx assigned by
taceous theropods are known from several sites throughout Aus-
Molnar and Pledge (1980) to this taxon was reappraised and
tralia (Table S1). The Western Australian record is restricted to
regarded as Archosauria indet. (Barrett et al., 2010). The
footprints from the Valanginian–Barremian Broome Sandstone
Queensland record comprises enantiornithine remains from the
(Colbert and Merrilees, 1967; Salisbury et al., 2017) and a pedal
upper Albian Toolebuc Formation (Molnar, 1986; Kurochkin
phalanx from the Cenomanian–Coniacian Molecap Greensand
and Molnar, 1997) and theropod footprints, megaraptorid teeth,
(Long, 1995). Two other reports of Western Australian Cretac-
and the megaraptorid Australovenator wintonensis (Table S2)
eous theropods are erroneous: the ‘caudal vertebra’ from the
from the Cenomanian–lower Turonian Winton Formation (Thul-
born and Wade, 1979, 1984; Hocknull et al., 2009; White et al.,
2012; Thulborn, 2013, 2017; White et al., 2013a, 2013b, 2015a,
*Corresponding author. 2015b, 2016). The New South Wales record appears to be
Color versions of one or more of the figures in the article can be found restricted to the Cenomanian Griman Creek Formation and
online at www.tandfonline.com/ujvp.

Published online 10 Oct 2019

 Poropat et al.—Cretaceous megaraptorids from Australia (e1666273-2)
comprises indeterminate theropod, avetheropod, megaraptorid,
and enantiornithine remains (Molnar, 1999; |Bell et al., 2016,
2019; Brougham et al., 2019).
The Cretaceous theropod fossil record of Victoria is the most
diverse of any Australian state (Poropat et al., 2018). More than
a dozen sites in the upper Barremian–lower Aptian upper Strze-
lecki Group (Gippsland Basin) and the upper Aptian–lower
Albian Eumeralla Formation (Otway Basin) have produced ther-
opod body fossils, but only four sites—two in each basin—have
been extensively sampled (Table S1). In the upper Strzelecki
Group, these are Koonwarra, from which several feathers have
been recovered (Talent et al., 1966; Waldman, 1970; Vickers-
Rich, 1991; Kellner, 2002), and Flat Rocks, which has produced
evidence of Neovenatoridae, Megaraptora (including 90 teeth),
Maniraptora, and Enantiornithes (Close et al., 2009; Benson
et al., 2012). Of the two sites that have been systematically exca-
vated in the Eumeralla Formation, only the theropod fauna of
Dinosaur Cove (comprising two localities: Dinosaur Cove East
and Slippery Rock) has been extensively documented. This site
has produced nonavian and avian theropod footprints (Martin
et al., 2014), skeletal elements assigned to Tetanurae, Neovena-
toridae, Megaraptora, Coelurosauria, Tyrannosauroidea, and
Maniraptora (Benson et al., 2012; Novas et al., 2013), and the
holotype left femur of Timimus hermani (Rich and Vickers-
Rich, 1994). By contrast, only two theropod specimens from
Eric the Red West have been described: an indeterminate thero-
pod caudal vertebra (Benson et al., 2012) and a cervical vertebra
assigned by some workers to Spinosauridae (Barrett et al., 2011;
Benson et al., 2012) and by others to Averostra or Tetanurae
(Novas et al., 2013). Another caudal vertebra (NMV P229456),
incorrectly reported to have come from the Eric the Red West
site, has been described twice: initially as Theropoda indet.
(Benson et al., 2012) and later as a referred specimen of the
ornithopod Diluvicursor pickeringi (Herne et al., 2018) without
mention of the previous assignation. This specimen is, however,
from the Eric the Red Site, some 200 m east of the Eric the
Red West site.
Periodic excavations at the Eric the Red West site between
2013 and 2017 have led to the recovery of several additional ther-
opod specimens, such that this site has now produced as many
theropod elements as Dinosaur Cove (Table S1). In this paper,
we describe two teeth, a cervical rib, a caudal vertebra, a non-
ungual manual phalanx, two manual unguals, and an astragalus
from the Eric the Red West site. We also reappraise the ‘spino-
saurid’ cervical vertebra previously described from this site.
Several of these specimens, including the cervical vertebra,
appear to be referable to Megaraptoridae. However, despite
their common provenance and apparent size congruence, no
two specimens can be confidently attributed to a single theropod
individual: each was found isolated, and all were sourced from a
fluvial deposit that also contains isolated bones pertaining to
other vertebrate groups (Poropat et al., 2018). Irrespective of
how many individuals are represented, these theropod remains
demonstrate that a megaraptorid slightly smaller than, but mor-
phologically almost identical to, Australovenator lived in south-
east Australia during the late Early Cretaceous.
Institutional Abbreviations—AODF, Age of Dinosaurs Fossil,
Australian Age of Dinosaurs Museum of Natural History,
Winton, Queensland, Australia; LRF, Lightning Ridge Fossil,
Australian Opal Centre, Lightning Ridge, New South Wales, Aus-
tralia; MUCPv, Museo de la Universidad Nacional del Comahue,
Neuquén, Argentina; NMV, Museums Victoria (formerly
National Museum of Victoria), Melbourne, Victoria, Australia.
GEOLOGICAL SETTING
The Eumeralla Formation forms a significant component of the
onshore exposed Otway Basin in western Victoria (Fig. 1).
Despite the fact that it crops out extensively throughout the
Otway Ranges, the inland portion of the Eumeralla Formation
is deeply weathered and largely covered by forest and farmland.
Consequently, with few exceptions (e.g., Binns Road Quarry;
Tosolini et al., 1998), the only fresh fossiliferous outcrops of
Eumeralla Formation available are those exposed along the
coast by wave action (Flannery and Rich, 1981; Rich and Rich,
1989; Poropat et al., 2018).
The Otway Basin has a complex tectonic history characterized
by multiple phases of compression and extension (Perincek et al.,
1994; Hill et al., 1995; Perincek and Cockshell, 1995; Debenham
et al., 2019). The initiation of rifting between Victoria and Tasma-
nia during the Tithonian led to the formation of the Otway Basin
(Norvick and Smith, 2001), within which the Crayfish Group
accumulated throughout the Tithonian–Barremian (Krassay
et al., 2004). The Eumeralla Formation, which overlies the Cray-
fish Group (or basement highs), largely comprises blue-gray vol-
caniclastic claystones and sandstones (Noll and Hall, 2003;
Krassay et al., 2004). These volcanogenic sediments were
sourced from a contemporaneous volcanic system to the east
(Edwards and Baker, 1943; Edwards et al., 1996; Mitchell, 1997;
Duddy, 2003)—possibly a volcanic arc east of the Gippsland
Basin (Veevers et al., 1982, 1991), the Whitsundays Volcanic Pro-
vince (Bryan et al., 1997), or a more proximal rift-related system
(Gleadow and Duddy, 1980; Duddy, 1983, 2003). These were
deposited in a floodplain paleoenvironment characterized by
river channels and shallow lakes (Felton, 1997a, 1997b; Noll
and Hall, 2003; Krassay et al., 2004).
The Eric the Red West site, which is situated near the tip of
Cape Otway, is hosted within the lower Albian portion of the
upper Aptian–lower Albian Eumeralla Formation. To date, this
site has produced hundreds of skeletal elements pertaining to
actinopterygians, dipnoans, plesiosaurs, testudinatans, pterosaurs,
ornithopods, theropods, and mammals (Herne et al., 2018;
Poropat et al., 2018; S.F.P., T.H.R., and P.V.-R., pers. observ.).
However, few of these specimens have been described: an inde-
terminate theropod caudal vertebra (Benson et al., 2012); a spino-
saurid (Barrett et al., 2011; Benson et al., 2012) or averostran/
tetanuran cervical vertebra (Novas et al., 2013); the holotype
caudal series, crus, and pes of the ornithopod Diluvicursor picker-
ingi (Herne et al., 2018); and a mammalian dentary assigned to
Bishops cf. B. whitmorei (Rich et al., 2009). A partial mammalian
maxilla (NMV P231328) has also been figured (as Tribosphenida,
gen. et sp. nov.) but not described (Poropat et al., 2018).
The theropod specimens described in this paper were recov-
ered from ‘Alan’s no. 3 Hole’ within the Eric the Red West site,
approximately 30 m east-northeast (ENE) of the Diluvicursor
pickeringi holotype locality. The geology of the Eric the Red
West site immediately surrounding the Diluvicursor site (which
also yielded the theropod cervical and the Bishops dentary) was
briefly summarized by Rich et al. (2009), who determined that
several of the fossils were preserved in debris deposits that
hosted logs and clay clasts. These accumulated around an
upright tree stump in a braided river system. Furthermore, Rich
et al. (2009) noted that away from the debris accumulation, the
sedimentology was indicative of a high-energy river deposit. A
more detailed assessment of the depositional setting of this part
of the Eric the Red West site was provided by Herne et al.
(2018), who defined the fossiliferous stratum as the Eric the
Red West Sandstone and the underlying stratum as the Anchor
Sandstone. The specimens described in this paper were preserved
in the lowermost section of the Eric the Red West Sandstone,
which has an undulating contact with the Anchor Sandstone
below. This contact is readily observed when the rocks are
freshly exposed: the basal-most Eric the Red West Sandstone in
‘Alan’s no. 3 Hole’ incorporates rip-up clasts derived from the
Anchor Sandstone, plant debris including logs, and rounded
pebbles sourced from Palaeozoic strata.
 Poropat et al.—Cretaceous megaraptorids from Australia (e1666273-3)

FIGURE 1. Map of Cape Otway and surroundings of southwest Victoria, Australia, showing the outcrop extent of the upper Lower Cretaceous Eumer-
alla Formation and all known Cretaceous fossil vertebrate sites on this portion of the Victorian coast (except Eastern View, situated north of Lorne).
Geological data derived from Edwards et al. (1996).

Most of the specimens described in this paper have been com-
pletely prepared out of their surrounding matrix. However, the
matrix that encased manual ungual phalanx I-2 (NMV
P239464) has been retained and registered under the same speci-
men number in the NMV collections. This gray sandstone matrix
typifies the basal Eric the Red West Sandstone: it is massive and
fine-grained, with few grains exceeding 0.5 mm in maximum
diameter, and also contains rounded mudstone rip-up clasts (up
to 27 mm long) and abundant carbonized plant fragments (Fig.
S1). The other manual ungual phalanx described herein (III-4;
NMV P252715A) also remains embedded within matrix, in
association with at least 10 additional isolated bones that
pertain to indeterminate theropods, ornithopods, and testudina-
tans (Poropat et al., 2018).
METHODS
Several of the theropod specimens described herein were
computed tomography (CT) scanned at St Vincent’s Hospital
Melbourne (Victoria, Australia) using a Revolution CT
scanner (GE Medical Systems). These specimens were
scanned at 140 kV and 401.5 mA, resulting in data with 0.625
cm voxel size. The CT data were imported into Mimics 21
(Materialise, Belgium) to enable digital preparation where
necessary and to enable three-dimensional (3D) models of
each element to be made. These 3D models were exported as
*.stl files, which were imported into Rhinoceros 6 (Robert
McNeel and Associates, U.S.A.) to enable the generation of
high-resolution two-dimensional renders of each element in all
six cardinal views.
SYSTEMATIC PALEONTOLOGY
THEROPODA Marsh, 1881
NEOTHEROPODA Bakker, 1986
TETANURAE Gauthier, 1986
TETANURAE, gen. et sp. indet.
Specimens—NMV P252700, right cervical rib; NMV P252704,
middle caudal vertebra; NMV P252405, right manual phalanx
II-1.
Horizon and Age—Eumeralla Formation, Otway Group,
Otway Basin. Early Albian (Wagstaff and McEwen Mason,
1989; Wagstaff et al., 2012; Korasidis et al., 2016).
Locality—All three specimens were found at the Eric the Red
West site, Cape Otway, Victoria (Rich et al., 2009). Specifically,
they were recovered from the eastern end of ‘Alan’s no. 3
Hole,’ a section of the site from which turtle and ornithopod
remains (among others) have also been extracted (Poropat
et al., 2018).
DESCRIPTION AND COMPARISONS
Right Cervical Rib
The cervical rib NMV P252700 is described with the long axis
horizontal (Fig. 2A–J; Table 1). In this orientation, the tubercu-
lum is situated dorsal to the capitulum and the elongate shaft pro-
jects posteriorly.
An anterolateral process was clearly present in NMV P252700.
Although broken, it seems unlikely that this process was much
more elongate than preserved. According to Carrano et al.
(2012), several non-tetanuran theropods possess elongate
 Poropat et al.—Cretaceous megaraptorids from Australia (e1666273-4)

FIGURE 2. Tetanurae indet. axial elements from the Eric the Red West site, Eumeralla Formation, Victoria, Australia. NMV P252700, right cervical rib
in A, B, dorsal, C, D, lateral, E, F, proximal, G, H, medial, and I, J, ventral views. NMV P252704, caudal vertebra in K, L, dorsal, M, N, posterior, O, P,
right lateral, Q, R, anterior, S, T, left lateral, and U, V, ventral views. A, C, E, G, I, K, M, O, Q, S, and U are photographs, whereas B, D, F, H, J, L, N, P, R,
T, and V are 3D renders derived from CT scan data. Scale bar equals 50 mm.

anterior processes on their cervical ribs, including Coelophysis
bauri (Colbert, 1989), Masiakasaurus knopfleri (Carrano et al.,
2011), and Majungasaurus crenatissimus (O’Connor, 2007). Dilo-
phosaurus wetherilli was scored by Carrano et al. (2012) as
showing this feature, despite Welles’ (1984) assertion that no
TABLE 1. Measurements (in mm) for NMV P252700, a right cervical
rib.
Dimension NMV P252700
Maximum proximodistal length
Dorsoventral height of proximal end
Proximal pneumatic foramen dorsoventral height
Proximal pneumatic foramen mediolateral width
Tuberculum dorsoventral height
Tuberculum anteroposterior length
cervical ribs were preserved, whereas Cryolophosaurus ellioti
was scored as having short processes despite the fact that Smith
et al. (2007) reported extremely elongate processes in this taxon.
Despite its relatively large size, it would appear that NMV
P252700 was not fused to its vertebra: the capitulum is preserved
as a complete, finished surface (Fig. 2G, H). However, fusion of
cervical ribs to vertebrae was observed in relatively few taxa by
Carrano et al. (2012): Coelophysis (Colbert, 1989), Ceratosaurus
(Madsen and Welles, 2000), Elaphrosaurus (Rauhut and Carrano,
2016), and Masiakasaurus (Carrano et al., 2011) among non-teta-
nurans and Neovenator (Brusatte et al., 2008) among the basal
115 tetanurans scored. Based on the interpreted absence of an
28 elongate anterolateral process, and the non-fusion of this
5 element to its vertebra, NMV P252700 appears to be referable
2 to Tetanurae.
13 The posterior shaft of NMV P252700 is long and slender, as is
7
that of the sole preserved cervical rib of the megaraptorid
 Poropat et al.—Cretaceous megaraptorids from Australia (e1666273-5)
Murusraptor barrosaensis (e.g., Coria and Currie, 2016) and the
few posterior cervical ribs of the juvenile Megaraptor namunhuai-
quii MUCPv-595 (Porfiri et al., 2014; S.F.P., pers. observ.). It is
likely that the posterior shaft of NMV P252700 was longer than
the vertebra with which it articulated. Based on the character
matrix of Brusatte et al. (2014), this feature excludes NMV
P252700 from Compsognathidae, Ornithomimosauria, and Man-
iraptora. Among non-avialan maniraptorans, some therizinosaurs
have cervical ribs that are longer than their respective cervical
vertebrae; however, the cervical ribs of Nothronychus mckinleyi
(Hedrick et al., 2015) and Falcarius utahensis (Zanno, 2010) are
morphologically divergent from NMV P252700 in other ways
(e.g., robusticity of the posterior shaft). Thus, NMV P252700
can be excluded from Therizinosauria, and from Maniraptori-
formes as a whole. Thus, it presumably pertains to Megalosauroi-
dea, Allosauroidea, or Tyrannosauroidea within Tetanurae.
Cervical ribs are known for several megalosauroid taxa, includ-
ing Afrovenator abakensis, Suchomimus tenerensis, and Spino-
saurus aegyptiacus (Sereno et al., 1994, 1998; Ibrahim et al.,
2014); however, they have not yet been described in detail for
any of these taxa. Nevertheless, in published schematics of all
three, the cervical ribs appear to possess elongate posterior
shafts and short, yet prominent, anterolateral processes. The
only figured cervical rib of Baryonyx walkeri abruptly tapers pos-
teriorly and has a deep pneumatic recess posterior to the capitu-
lum and tuberculum (Charig and Milner, 1997); both of these
features distinguish it from NMV P252700. An anterior cervical
rib of the megalosaurid Dubreuillosaurus valesdunensis (Allain,
2005:fig. 7b, c) is quite similar to NMV P252700. The morphology
of the anterolateral process and the degree of posterior tapering
in both are almost identical; however, the cervical rib of Dubreuil-
losaurus can be distinguished from NMV P252700 (Fig. 2E, F) by
its more developed anterior foramen. Furthermore, each of the
cervical ribs of Dubreuillosaurus bears a deep excavation on its
medial surface near the proximal end, a feature not evident in
NMV P252700.
Cervical ribs are known for many allosauroid taxa. The cervical
ribs of the metriacanthosaurids Yangchuanosaurus shangyouensis
and Sinraptor dongi appear to have a more elongate anterolateral
process than NMV P252700 (Dong et al., 1983; Currie and Zhao,
1993; Gao, 1993). Based on the description provided by Currie
and Zhao (1993), the morphology and position of the pneumatic
recesses on those of Sinraptor is also quite different from NMV
P252700 (Fig. 2C, D, G, H). The cervical ribs of the allosaurid
Allosaurus fragilis, particularly those figured by Madsen (1976:
pl. 38), are quite similar to NMV P252700 with respect to the mor-
phology of the anterior pneumatic foramen and the posterior
shaft. However, the anterolateral processes of these ribs, and
most of those figured by Gilmore (1920:fig. 18), are quite
rounded, in stark contrast to the rather pointed process of
NMV P252700 (Fig. 2A–D). Cervical ribs from the carcharodon-
tosaurid Acrocanthosaurus atokensis are preserved in at least two
exemplars (Harris, 1998; Currie and Carpenter, 2000). The pos-
terior cervical rib described by Currie and Carpenter (2000:fig.
5A) is similar overall to NMV P252700, although the anterolat-
eral process in this rib does not project beyond the margins of
the bases of the tuberculum and capitulum. By contrast, all of
the cervical ribs described and illustrated by Harris (1998) can
be distinguished from NMV P252700 (Fig. 2E, F) by the more
acute angle between the tuberculum and the capitulum, and the
resultant position of the anterolateral process ventral to both,
rather than between them, in the dorsoventral plane. The cervical
ribs of Concavenator corcovatus, in lateral view at least, appear
superficially similar to NMV P252700 despite the fact that most
of them have substantially longer anterolateral processes
(Ortega et al., 2010; Cuesta et al., 2019).
The sole preserved cervical rib of Neovenator lacks a pneumatic
opening on its lateral surface (Brusatte et al., 2008), as does NMV
P252700 (Fig. 2C, D). Nevertheless, much of the anterior part of
the lateral surface in NMV P252700 is concave. The ventral
margin of this concavity is delimited by an anteroposteriorly
oriented ridge, which runs posteriorly from the anterolateral
process along the anterior third of the shaft. A similar feature,
described as a robust ridge, was identified on the sole preserved
cervical rib of Neovenator (Brusatte et al., 2008); this might indi-
cate that it also had anterolateral processes on its cervical ribs.
The majority of the medial surface of NMV P252700 forms a
fossa (Fig. 2G, H), which appears to contrast with the situation
in Neovenator wherein the concavity is limited to the anterior-
most portion of the rib and was interpreted as a pneumatic
foramen (Brusatte et al., 2008). A pneumatic foramen is present
on the anterior surface of NMV P252700 (Fig. 2E, F), bordered
medially by a web of bone that connects the tuberculum with
the capitulum. Similar pneumatic foramina have been reported
in the cervical ribs of Aerosteon riocoloradensis (Sereno et al.,
2008) and Neovenator (Brusatte et al., 2008; n.b., these authors
described the cervical rib with the long axis of the shaft vertical,
and thus this feature was described as a dorsal pneumatopore)
and can also be seen in many other theropods, including Allo-
saurus (Madsen, 1976) and Tyrannosaurus rex (e.g., Brochu,
2003). The tuberculum of NMV P252700 is taller dorsoventrally
than it is long anteroposteriorly (Fig. 2G, H); the inverse is true
in Neovenator (Brusatte et al., 2008), the only preserved cervical
rib of which is also substantially larger than NMV P252700.
Cervical ribs have not been described in detail for any basal tyr-
annosauroids, although those of Guanlong wucaii appear to be
somewhat similar to NMV P252700 (Xu et al., 2006; Choiniere,
2010). By contrast, the cervical ribs of several tyrannosaurids
have been documented in detail, including Alioramus altai (Bru-
satte et al., 2012), Gorgosaurus libratus (Lambe, 1917), and Tyr-
annosaurus (Brochu, 2003). In all of these taxa, the posterior
shaft of the cervical ribs tapers more abruptly, and becomes sig-
nificantly more spindly distally, than in NMV P252700, although
we concede that it too might have tapered to a fine point when
complete.
Based on our comparisons of NMV P252700 with the cervical
ribs of several megalosauroid, allosauroid, and tyrannosauroid
taxa, we contend that it is most similar to megaraptorids.
However, our inability to determine the precise position of this
cervical rib within the cervical rib series, and our incomplete
understanding of non-maniraptoriform tetanuran cervical ribs,
renders such a precise assignation questionable. Thus, we inter-
pret NMV P252700 as a non-maniraptoriform tetanuran cervical
rib, while tentatively entertaining the possibility that it pertains to
a megaraptorid.
Finally, NMV P252700 preserves two apparent pathologies.
The first is a small protuberance near the posterior end of the
posterior shaft, which appears to represent a healed
lesion (Fig. 2G–J). The second is a small, shallow depression
on the lateral surface of the anterolateral process, within
which the cortical bone is depressed medially relative to the
lateral surface of the rib (Fig. 2C, D). This structure, which is
longer anteroposteriorly (4 mm) than tall dorsoventrally (3
mm), is followed 2 mm posterodorsally by another, slightly
longer depression (5 mm long anteroposteriorly × 3 mm dorso-
ventrally). Posterior to this, a natural sulcus on the lateral
surface of the shaft (ventral–posteroventral to the tuberculum)
appears to have been slightly exaggerated by compression
from the lateral side.
The two anterior depressions on the anterolateral process of
NMV P252700 are herein interpreted as bite marks (Fig. 2C,
D). Based on their arrangement and dimensions, it is plausible
that the bite was inflicted by a small theropod. Their spacing
suggests that the aggressor bit below the neck and in the direction
of the head. The absence of any healing traces suggests that this
bite was inflicted perimortem.
 Poropat et al.—Cretaceous megaraptorids from Australia (e1666273-6)

Middle Caudal Vertebra the neural spine is flanked on each side by subtle ridges. These
ridges connect the bases of each prezygapophysis to its ipsilateral
Specimen NMV P252704 is a poorly preserved middle caudal
postzygapophysis. In lateral view, the dorsal surfaces of these
vertebra (Fig. 2K–V; Table 2). The middle part of the centrum
ridges are shallowly concave.
sustained substantial damage in the field, whereas significant
The right postzygapophysis is far more complete than the left
parts of the transverse processes, zygapophyses, and neural
and appears to be missing little of its original surface. In addition,
spine were lost pre-fossilization. Nevertheless, NMV P252704
the ventrolaterally facing postzygapophyseal facet appears to be
is clearly morphologically consistent with the middle caudals
preserved. If true, this would suggest that the prezygapophyses
of numerous theropod taxa and is interpreted as a theropod
of the succeeding vertebra were not substantially longer than
middle caudal herein.
those preserved on NMV P252704; this might further imply that
The centrum is spool-shaped and amphicoelous, with shal-
the prezygapophyses of NMV P252704 are almost complete. The
lowly concave articular facets at both ends. The anterior articu-
bases of the postzygapophyses are confluent with each other and,
lar surface (Fig. 2M, N) is far more complete than the posterior
dorsally, with the underside of the neural spine. This combined
one (Fig. 2Q, R); however, based on the preservation of the
unit forms the dorsal margin of the posterior neural canal
latter, there can be little doubt it would have looked similar
opening (7.2 mm wide transversely × 4.6 mm tall dorsoventrally).
to the former when complete. Two subtle, parallel ridges situ-
Although caudal vertebrae are known for several neovenatorid
ated 3.5 mm from one another run anteroposteriorly along
and megaraptoran taxa, these often derive from different parts of
the ventral margin of the centrum (Fig. 2U, V); these presum-
the tail than NMV P252704. For example, the only described
ably track the path of the blood vessels that ran through the
caudal vertebra of Fukuiraptor kitadaniensis is from the posterior
hemal canal. Chevron facets were not observed at either end,
section of the tail (Azuma and Currie, 2000) and cannot be mean-
although this is presumably because the ventral surfaces of
ingfully compared with NMV P252704. The only preserved
both ends are heavily worn. No other prominent features are
caudal vertebrae of Siats meekerorum are situated either
present on the ventral or lateral surface of the centrum of
further anteriorly or posteriorly within the caudal series than
NMV P252704; indeed, the lateral surfaces are essentially flat,
NMV P252704 (Zanno and Makovicky, 2013). The anterior
particularly in the middle third.
caudals lack centra, thus meaningful comparison with NMV
As preserved, the transverse processes of NMV P252704 are
P252704 is not possible; by contrast, the posterior caudals have
anteroposteriorly elongate ridges (no more than 4 mm tall dor-
more prominent prezygapophyses and postzygapophyses than
soventrally along their length; Fig. 2O–P, S, T). The bases of
NMV P252704. A caudal vertebra from the Wayan Formation
these processes fade out before reaching either end. The left
(Albian–Cenomanian) of Idaho was referred to Neovenatoridae
transverse process (Fig. 2S, T) is more complete than the right
partly because of the presence of a single pneumatic foramen on
one (Fig. 2O, P): in the former, the anterior third is intact,
the lateral surface (Krumenacker et al., 2017); this feature dis-
whereas the posterior two-thirds (∼23 mm) constitute a
tinguishes it from NMV P252704.
broken surface. Thus, if the transverse processes formed promi-
Of the preserved vertebrae of Neovenator, NMV P252704 is
nent, lateral projections, such structures were restricted to the
most similar to caudal vertebra ‘O’ as per Brusatte et al. (2008).
posterior two-thirds of the vertebra. Based on the orientation
However, that vertebra—and all of the successive caudal vertebrae
of the cortical bone texture, it is likely that the transverse pro-
in Neovenator—has greatly enlarged, anteriorly projecting prezy-
cesses on this vertebra were low along their length, as in the
gapophyses that contrast with those of NMV P252704. Based on
middle caudal vertebrae of other theropods (e.g., Neovenator;
the bone texture of the prezygapophyseal bases in NMV
Brusatte et al., 2008).
P252704, the unpreserved, more anterior parts of the prezygapo-
The bases of both prezygapophyses are preserved, but the
physes were directed more dorsally than anteriorly. Consequently,
ends of both are broken; as such, their anterodorsal extent
they might have resembled the prezygapophyses of caudal ver-
cannot be determined with certainty. The prezygapophyseal
tebrae ‘L’ and ‘M’ of Neovenator as per Brusatte et al. (2008).
facets appear not to be preserved, so it can be assumed that
The caudal vertebrae of several megaraptorids have been
the prezygapophyses as a whole were at least slightly longer
reported. Although the absence of pneumatic foramina on the
in life. A thin plate of bone bridges the gap between the poster-
lateral surface of the centrum of NMV P252704 distinguishes it
omedial margins of the prezygapophyseal bases (Fig. 2Q, R).
from the anterior–middle caudal vertebrae of Aerosteon and
Concomitantly, this plate forms the roof of the anterior neural
Megaraptor (Calvo et al., 2004; Sereno et al., 2008; Porfiri et al.,
canal opening (7.3 mm wide transversely × 4.7 mm tall dorso-
2014), the vertebrae preserved in these taxa are from further
ventrally), as well as the floor of a 2.6 mm wide slit—the pre-
anterior in the tail than NMV P252704. It is probable that fora-
spinal fossa—that extends posteriorly between the
mina were not persistent along the entire caudal series in mega-
prezygapophyses for 16.6 mm then courses under the base of
raptorids. That said, the only known middle and posterior
the neural spine (Fig. 2K, L).
caudal vertebrae of Aerosteon possess pneumatic foramina,
The neural spine itself is restricted to the posterior two-thirds of
have a camellate internal texture, and are less proportionally
the vertebra and is exceedingly thin transversely (1.5 mm).
elongate than those of other theropods (Sereno et al., 2008).
Although it is broken along its entire dorsal surface, it seems unli-
The caudal vertebrae of Aoniraptor libertatem show significant
kely that it was substantially taller than preserved. In dorsal view,
serial variation. The middle–posterior caudal centra are elongate
relative to overall size and lack pneumatic foramina or fossae,
despite having camerate internal texture (Motta et al., 2016).
One caudal in particular, figured and described as a posterior
caudal despite having prominent transverse processes (Motta
TABLE 2. Measurements (in mm) for NMV P252704, a caudal vertebra.
et al., 2016:fig. 23), is quite similar to NMV P252704: the trans-
verse processes are restricted to the posterior two-thirds of the
Dimension NMV P252704
centrum, the postzygapophyses extend only slightly posterodorsal
Centrum Anteroposterior length 53 to the posterior margin of the centrum, the prezygapophyses are
Anterior articular facet Maximum height 17 relatively small and project anterodorsally rather than mostly
Maximum width 21
Posterior articular facet Maximum height 17
anteriorly, and a prespinal fossa is present between the prezyga-
Maximum width 21 pophyses. Nevertheless, assigning NMV P252704 to Megaraptor-
idae on the basis of these superficial similarities is not possible,
 Poropat et al.—Cretaceous megaraptorids from Australia (e1666273-7)

FIGURE 3. Tetanurae indet., NMV P252405, right manual phalanx II-1 from the Eric the Red West site, Eumeralla Formation, Victoria, Australia, in A,
B, medial, C, D, proximal, E, F, lateral, G, H, dorsal, I, J, distal, and K, L, ventral views. A, C, E, G, I, and K are photographs, whereas B, D, F, H, J, and L
are 3D renders derived from CT scan data. Scale bar equals 50 mm.

because several of these features are more widely distributed
among Theropoda.
Right Manual Phalanx II-1
Specimen NMV P252405 is an elongate phalanx with ginglymoid
articular facets at both ends (Fig. 3; Table 3). It is nearly complete,
lacking only a small part of the ventral surface of the proximal
end. The bone texture of the mid-shaft and articular surfaces is
smooth, whereas the non-articular portions of the proximal and
distal ends are characterized by longitudinally striated bone texture.
The proximal end of NMV P252405 is oval in proximal view
and is bisected by a vertical midline ridge, either side of which
lies a concave articular facet (Fig. 3C, D). In medial (Fig. 3A, B)
and lateral (Fig. 3I, J) views, the proximal end is strongly
TABLE 3. Measurements (in mm) for NMV P252405, a right manual
phalanx II-1.
Dimension NMV P252405
Proximodistal length
Proximal mediolateral breadth
Proximal dorsoventral height
Mid-shaft minimum circumference
Distal mediolateral breadth
Distal dorsoventral height
concave dorsoventrally; this is partially attributable to the pres-
ence of a mediolaterally narrow extensor tubercle. The proximal
articular facet of NMV P252405 implies that the distal articular
facet of the preceding metacarpal or phalanx was functionally
ginglymoid. Furthermore, the lateral articular facet is slightly
deeper than the medial one on the proximal end of NMV
P252405. The dorsal surface of the shaft of the phalanx is
almost entirely mediolaterally convex, except at the distal end
where a prominent extensor fossa is present immediately prox-
imal to the distal articular facet (Fig. 3G–H). Proximodistally
elongate vascular canals terminating in foramina are present
on the medial and lateral surfaces of the proximal end of
NMV P252405 (Fig. 3A, B, I–J).
The ginglymoid distal end of NMV P252405 is divided into two
convex articular facets separated by a very slightly dorsolaterally–
ventromedially inclined midline ridge (Fig. 3E, F). The medial
condyle is taller dorsoventrally (26 vs. 24 mm) but shorter proxi-
modistally (26 vs. 27 mm) than the lateral condyle in NMV
P252405. The medial collateral pit (11 mm long proximodistally
× 6 mm tall dorsoventrally) is slightly smaller than the lateral
one (12 mm long proximodistally × 6 mm tall dorsoventrally),
although the former is deeper than the latter.
We interpret NMV P252405 as a manual phalanx because it is
96 proximodistally elongate, the ventral surface of its shaft is
31 strongly convex, the distal end is taller dorsoventrally than wide
37 mediolaterally, and the distal condyles are only shallowly separ-
72 ated ventrally (cf. Allosaurus; Madsen, 1976:pls. 43–45, 53–55).
27
27
The central position of the collateral pits on the medial and
lateral surfaces of the distal condyles indicates that this element
 Poropat et al.—Cretaceous megaraptorids from Australia (e1666273-8)
is not a distal-most non-ungual phalanx, because in such pha-
langes the collateral pits are often shifted dorsally. In light of
this, it is likely that NMV P252405 is a manual phalanx II-1. If
so, then it is markedly divergent from the corresponding
manual phalanges of the megaraptorids Australovenator (White
et al., 2012) and Megaraptor (Calvo et al., 2004), which are pro-
portionally less elongate proximodistally and have strongly
arched ventral margins in medial and lateral views.
Given the difficulty associated with determining the anatomical
placement of NMV P252405, it seems prudent to regard it as Teta-
nurae indet. Although there are similarities between NMV
P252405 and the Australovenator phalanx originally identified
as right manual III-1 (White et al., 2012) but later reinterpreted
as left pedal III-3 (White et al., 2016), in the latter element the
collateral pits are positioned dorsally rather than centrally. We
regard NMV P252405 as Tetanurae indet., because it clearly
does not pertain to Ceratosauria (Ruiz et al., 2011; Burch and
Carrano, 2012; Pol and Rauhut, 2012; Carrano and Choiniere,
2016; Guinard, 2016).
TETANURAE Gauthier, 1986
MEGARAPTORA Benson, Carrano, and Brusatte, 2010
MEGARAPTORIDAE Novas, Agnolín, Ezcurra, Porfiri, and
Canale, 2013
Brief Comment on the Relationships of Megaraptoridae—The
phylogenetic position of Megaraptoridae within Theropoda
remains contentious (Novas et al., 2013; Bell et al., 2016;
Ezcurra and Novas, 2016; Novas et al., 2016; Aranciaga
Rolando et al., 2019). Some researchers have classified the
slightly more inclusive clade Megaraptora as a subset of Neovena-
toridae nested within Allosauroidea (Benson et al., 2010; Carrano
et al., 2012; Zanno and Makovicky, 2013), whereas others have
included Megaraptoridae within the Coelurosauria (Novas
et al., 2015; Porfiri et al., 2018; Aranciaga Rolando et al., 2019;
Samathi et al., 2019), sometimes within Tyrannosauroidea
(Novas et al., 2013; Porfiri et al., 2014). The true phylogenetic pos-
ition of Megaraptoridae will need to be determined before the
paleobiogeographic implications of its constituents are fully
understood. However, it should be noted that both Tyrannosaur-
oidea (Table S3) and Neovenatoridae (Table S4) are almost
exclusively Laurasian when megaraptorids are exempted from
consideration; isolated neovenatorid remains have been reported
from the upper Strzelecki Group of southeast Australia, whereas
tyrannosauroids have been reported from the Eumeralla For-
mation of southeast Australia and the Santana Formation of
northeast Brazil (Table S5). It is possible that some or all of
these specimens might, in fact, be megaraptorans (or
megaraptorids).
Recently, Motta et al. (2016) suggested that similarities
between the Argentinean megaraptoran Aoniraptor and the
African theropods Bahariasaurus ingens (Stromer, 1934) and
Deltadromeus agilis (Sereno et al., 1996) might be indicative of
a close phylogenetic relationship. Motta et al. (2016) united
these taxa as Bahariasauridae and also alluded to the then-unde-
scribed Gualicho shinyae (Apesteguía et al., 2016) as another
possible member of this clade (Table S6). However, as noted by
Porfiri et al. (2018), the reduced forelimbs and functionally didac-
tyl manus of Gualicho are in stark contrast to the massive fore-
limbs, tridactyl manus, and hypertrophied unguals of
Megaraptor and Australovenator.
MEGARAPTORIDAE, gen. et sp. indet.
Specimen—NMV P221081, cervical vertebra.
Horizon and Age—Eumeralla Formation, Otway Group,
Otway Basin. Early Albian (Wagstaff and McEwen Mason,
1989; Wagstaff et al., 2012; Korasidis et al., 2016).
Locality—Found at the Eric the Red West site, Cape Otway,
Victoria, ca. 5 m from the Diluvicursor pickeringi holotype indi-
vidual (Herne et al., 2018).
DESCRIPTION AND COMPARISONS
Cervical Vertebra
This cervical vertebra, NMV P221081, was originally assigned
to Spinosauridae (Barrett et al., 2011; Benson et al., 2012). Refer-
ral to Tetanurae was justified by the presence of a single pneu-
matic foramen on each lateral surface of the centrum, whereas
referral to Allosauroidea or Megalosauroidea was inferred on
the basis of the strongly opisthocoelous centrum (Barrett et al.,
2011; Benson et al., 2012). The subdivision of the pneumatic
foramen by a short, simple, internal lamina, and the elongate
nature of the centrum (length:height ratio >2.0), was cited to
support its referral to Spinosauridae, whereas the non-camellate
nature of the internal bone was used to exclude it from Carchar-
odontosauria (Barrett et al., 2011; Benson et al., 2012). More
recently, the spinosaurid affinities of this cervical vertebra have
been questioned and an averostran or tetanuran classification
supported (Novas et al., 2013).
At the time that Barrett et al. (2011) described NMV P221081,
very few megaraptorid cervical vertebrae were known (Calvo
et al., 2004). The most critical specimen described since is
MUCPv-595, a juvenile exemplar of Megaraptor (Porfiri et al.,
2014). Personal observation of this specimen (by S.F.P., 2018) con-
firmed the following features identified (but not illustrated) in the
original description (Porfiri et al., 2014): the cervical centra are
opisthocoelous with hemispherical condyles and flat ventral sur-
faces (except cervical IX); and the internal texture is camellate,
with smaller chambers at the articular ends and on the apophyses
and larger ones within the centrum. However, paired elliptical
pneumatic foramina are not present dorsal to each parapophysis
(Fig. 4A, B): this feature is absent on one or both sides of some
cervicals and is not observable in others (e.g., the right side of cer-
vical VI in MUCPv-595 has two pneumatic foramina, whereas the
left has only one; S.F.P., pers. observ.; Fig. 4A, B). Specimen NMV
P221081 is similarly asymmetrical (Fig. 4C): the left side bears two
pneumatic foramina (one of which is hosted largely on the dorsal
surface of the parapophysis and is barely visible in lateral view),
whereas the right bears a single pneumatic foramen. Each large
foramen is subdivided internally by a single simple strut (as
described by Barrett et al. [2011]). Aranciaga Rolando et al.
(2019:124) stated that an apomorphy of Megaraptoridae was
“postaxial cervical vertebrae, with two pleurocoels separated by
an anterodorsally oriented thin lamina.” In light of this, the fact
that even one side of NMV P221081 presents two pneumatic for-
amina suggests that it might be referable to Megaraptoridae.
The centrum of cervical VII of MUCPv-595 (55 mm sans
cotyle) is more than twice as long anteroposteriorly as either of
its articular surfaces is tall dorsoventrally (anterior: 24 mm; pos-
terior: 29 mm; S.F.P., pers. observ.). Thus, the length:height ratio
of this cervical was 1.90 (based on the posterior height) or 2.29
(anterior), versus ∼2.00 (posterior) in NMV P221081. This indi-
cates that elongate cervical vertebrae sensu Barrett et al. (2011)
were present in at least one section of the only known articulated
megaraptorid cervical series.
Novas et al. (2013) likened the internal texture of NMV
P221081 to that of the then-unpublished juvenile Megaraptor
(Porfiri et al., 2014). As related by Porfiri et al. (2014), the internal
texture of the cervical vertebrae of MUCPv-595 varied within
each element; the same is true in NMV P221081, as noted by
Novas et al. (2013). Consequently, the internal texture of NMV
P221081 does not exclude it from Carcharodontosauria (contra
Barrett et al. [2011]) and might in fact suggest affinities with
Megaraptoridae.
 Poropat et al.—Cretaceous megaraptorids from Australia (e1666273-9)

FIGURE 4. Megaraptorid cervical vertebrae.

A, B, Megaraptor namunhuaiquii (MUCPv-
595) cervical vertebrae V–VII in A, left
lateral and B, right lateral views, showing the
presence of a single pneumatic foramen on
the left side of cervical vertebrae VI and VII
and the presence of a pair of pneumatic fora-
mina on the right side of cervical vertebra VI.
C, ?Megaraptoridae, gen. et sp. indet. (NMV
P221081; formerly Spinosauridae, gen. et sp.
indet.: Barrett et al., 2011), middle cervical ver-
tebra in left lateral view, showing the presence
of a pair of pneumatic foramina on the left
side. Scale bar equals 50 mm (A, B) and 20
mm (C).

Overall, NMV P221081 is similar to cervical vertebrae V–VII
of MUCPv-595 (Fig. 4). The presence of two pneumatic foramina
on the left side (centrum + parapophysis) of NMV P221081 sup-
ports its referral to Megaraptoridae, rather than Spinosauridae.
Consequently, in agreement with Novas et al. (2013), spinosaurids
are not yet represented in the Australian fossil record.
Horizon and Age—Eumeralla Formation, Otway Group,
Otway Basin. Early Albian (Wagstaff and McEwen Mason,
1989; Wagstaff et al., 2012; Korasidis et al., 2016).
Locality—All specimens were found at the Eric the Red West
site, Cape Otway, Victoria, specifically from the eastern end of
‘Alan’s no. 3 Hole.’

MEGARAPTORIDAE cf. AUSTRALOVENATOR

WINTONENSIS Hocknull, White, Cook, Tischler, Calleja,
Sloan, and Elliott, 2009
Specimens—NMV P239459, tooth; NMV P252264, tooth;
NMV P239464, right manual ungual I-2; NMV P252715A, left
manual ungual III-4; NMV P253701, right astragalus.
DESCRIPTION AND COMPARISONS
Teeth
To date, two megaraptorid teeth, NMV P239459 (Fig. 5A) and
NMV P252264 (Fig. 5B–E), have been recovered from the Eric
the Red West site (Table 4). The description of these teeth
 Poropat et al.—Cretaceous megaraptorids from Australia (e1666273-10)

FIGURE 5. Megaraptoridae cf. Australovenator wintonensis teeth from the Eric the Red West site, Eumeralla Formation, Victoria, Australia. A, NMV
P252264 in lingual view; B–E, NMV P239459 in B, labial, C, distal, D, lingual, and E, mesial views. All images are photographs. Scale bar equals 10 mm
(A) and 6.66 mm (B–E).

follows the methodology and terminology outlined by Hendrickx Both teeth are moderately narrow labiolingually (crown:base
et al. (2015). ratio ≈0.5) and have smooth enamel. The precise cross-sectional
Both NMV P239459 and NMV P252264 suffered damage from shape of the base of NMV P252264 cannot be determined
hand tools when they were discovered; consequently, they have because of its incomplete state of preparation, although the pres-
been preserved with Paraloid B72 and repaired with superglue. ence of a subtle depression on the presumed lingual surface might
The teeth were prepared using fine needles, meaning that it is be indicative of it having a figure-8 cross-sectional shape at the
possible that some fine striations were introduced during prep- base. This was tentatively identified as a synapomorphy of Mega-
aration. Only one half of NMV P252264, the larger tooth, has raptoridae by White et al. (2015a). By contrast, the preserved part
been prepared (Fig. 5A), whereas the smaller tooth, NMV of NMV P239459 is oval in cross-section. However, given that the
P239459, is fully prepared (Fig. 5B–E). Neither tooth appears crown of NMV P239459 is incomplete, it is possible that further
to have been subjected to significant postmortem distortion. basally it, too, would have had a figure-8 cross-sectional shape.
Specimens NMV P239459 and NMV P252264 are ziphodont In both teeth, denticles are present only on the distal carina.
lateral teeth with distally recurved apices, convex mesial The apical portion of the mesial surface in NMV P239459 has
margins, and concave distal carinae. The latter is substantially been obliterated by an extensive wear facet. By contrast, the
more complete than the former, shows a greater degree of curva- mesial surface of NMV P252264 is largely unworn yet lacks den-
ture, and also preserves the apical-most part of the root base ticles regardless. However, the apical-most portion of the mesial
(Fig. 5A). This suggests that NMV P252264 was forcibly dislodged surface has been affected by a wear facet, and the exposed
from its alveolus post mortem, presumably during hydraulic surface also appears to be slightly pitted. Thus, it is possible
transport. By contrast, the root of NMV P239459 is entirely that the apical-most section of the mesial surface of NMV
lacking, and the broken basal end is infilled with matrix. A P252264 bore a carina and possessed denticles when freshly
subtle depression is present on the presumed lingual surface of erupted, but that these wore away. The distal carina of NMV
each tooth—in the case of NMV P252264, this is the only P252264 preserves 46 denticles in total, with four denticles per
surface fully exposed. The subtle, presumably labial, deflection millimeter on average; this matches the condition of the second
of the distal carina of NMV P252264 supports the interpretation dentary tooth in Australovenator (denticles are damaged and sub-
that the exposed surface is the lingual one. sequently not observable in the other teeth in this specimen) but
represents a greater denticle density overall than the shed teeth
from the Australovenator type locality (White et al., 2015a). In
TABLE 4. Measurements (in mm) for NMV P239459 and NMV NMV P252264, the denticles appear to be slightly less densely
P252264, teeth, following the terminology of Hendrickx et al. (2015). packed along the apical segment of the distal carina than they
are further basally. This contrasts with the isolated teeth from
Dimension NMV P239459 NMV P252264 the Australovenator type locality, which were described as
having denticles least densely packed at the mid-height of the
Crown height (CH) 11.5 15.8 crown, with denticle density increasing both apically and basally
Crown:base length (CBL) 6.2 8.1
Crown:base width (CBW) 3.8 ∼4 (White et al., 2015a). Although the lingual surface of the crown
Apical length (AL) 12 17.2 apex preserves a wear facet, this does not intersect the distal
Crown:base ratio (CBR) 0.61 ∼0.5 carina. The well-worn mesial surface of NMV P239459 lacks den-
Crown:height ratio (CHR) 1.86 1.95 ticles altogether, whereas its distal carina preserves 11 denticles
Denticles per mm (total) 28/6.9 mm = 4.06 46/11.4 mm = 4.04 within the apical-most 3 mm (3.66/mm); basal to this, an
Denticles per mm (apex) 11/3 mm = 3.67 31/7.8 mm = 3.97
Denticles per mm (base) 17/3.9 mm = 4.36 15/3.6 mm = 4.17
additional 17 broken denticle bases are present within a 3.9-mm
section (4.36/mm). Although the average denticle density is
Note that NMV P239459 is broken; thus, the measurements are only four per millimeter overall, the denticles on NMV P239459 are
indicative of the preserved portion. more densely packed basally than apically, as in NMV P252264.
 Poropat et al.—Cretaceous megaraptorids from Australia (e1666273-11)
Denticles are relatively uniform in size, shape, and spacing along
each tooth. The denticles on both teeth are small, subquadrangu-
lar, oriented more or less perpendicular to the carina, and are
chisel-shaped sensu Hendrickx et al. (2015), with uniformly
convex external margins. Shallow interdenticular slits are
present between all denticles, the interdenticular diaphyses are
narrow, and the interdenticular spaces are shallow. No interdenti-
cular sulci were observed.
White et al. (2015a) concluded that megaraptorid teeth gener-
ally lacked denticles on the mesial carina but noted that denticles
were sometimes present toward the apex. Indeed, in both Mega-
raptor (Porfiri et al., 2014) and Orkoraptor burkei (Novas et al.,
2008), the mesial carinae lack denticles altogether. The situation
in Murusraptor is somewhat more complex: in most teeth, the
mesial carinae lack denticles altogether, but in the six teeth
where the crown apex was preserved, the apical-most section of
the mesial carina supported six to 10 denticles—often completely
worn (Coria and Currie, 2016). This might indicate that all of the
teeth of Murusraptor had a small number of denticles on the
apical portion of the mesial carina, but that these were often
worn off by tooth-tooth contact. However, personal observation
of these teeth (by S.F.P., 2018) revealed a high degree of size
and morphological disparity—it is probable that they derive
from more than one megaraptorid individual and that non-mega-
raptorid theropod teeth are also present in the sample. At least
three of the 10 isolated theropod teeth recovered from the Aus-
tralovenator type locality in the Winton Formation (AODF 822,
AODF 824, AODF 825) preserve denticles along their mesial
carinae, but the rest were too poorly preserved for the presence
or absence of denticles to be determined, even when the mesial
carina itself was present (White et al., 2015a). Unfortunately,
none of the in situ teeth in the right dentary of Australovenator
were sufficiently well preserved to allow the presence of mesial
denticles to be determined (White et al., 2015a). In stark contrast
to the teeth of Gondwanan megaraptorids, all preserved teeth of
the megaraptoran Fukuiraptor (Azuma and Currie, 2000; Currie
and Azuma, 2006; Molnar et al., 2009) and the neovenatorid Neo-
venator (Brusatte et al., 2008) have fully denticulate mesial
carinae. Finally, it should be noted that the only tooth attributed
to the megaraptorid Aerosteon (Sereno et al., 2008) has since
been interpreted as that of an abelisaurid (Novas et al., 2013;
Coria and Currie, 2016); consequently, it is excluded from the
comparisons undertaken herein.
Both NMV P239459 and NMV P252264 preserve wear facets
on the mesial sides of their crown apices (Fig. 5A, B, E). In the
case of NMV P252264, the wear facet (4 mm long apicobasally)
only intersects the mesial surface on the crown apex; otherwise,
it is situated entirely on the (presumed) lingual surface. By con-
trast, in NMV P239459, the wear facet is much more extensive
(8 mm long apicobasally) and has obliterated much of the
mesial surface. In mesial view, the midline of the wear facet lies
to one side of the midline (presumed herein to be labial) of the
tooth as a whole.
Right Manual Ungual I-2
Specimen NMV P239464 is a hypertrophied, sickle-shaped
ungual phalanx (Fig. 6; Table 5). It is almost identical to, but
slightly smaller than, manual ungual I-2 of Australovenator
(Hocknull et al., 2009; White et al., 2012). It is also very similar
to, albeit significantly smaller than, the manual ungual I-2 of
Megaraptor (Novas, 1998; Calvo et al., 2004).
Apart from Australovenator (Hocknull et al., 2009; White et al.,
2012) and Megaraptor (Novas, 1998; Calvo et al., 2004; Porfiri
et al., 2007), manual ungual I-2 is represented in relatively few
published megaraptorid, megaraptoran, or neovenatorid speci-
mens. However, some specimens have been described in an
unpublished thesis (Lamanna, 2004), and others have been
illustrated in a published paper (Casal et al., 2016); these
unguals show few differences from those described (S.F.P., pers.
observ.). The sole preserved ungual phalanx of Chilantaisaurus
tashuikouensis (Hu, 1964) has been interpreted as manual
ungual I-2 (Benson and Xu, 2008) but was more recently reinter-
preted as manual ungual II-3 (White et al., 2012). We follow the
latter interpretation; thus, NMV P239464 will not be compared
with Chilantaisaurus.
A fragmentary theropod specimen (LRF 100–106) from the
Cenomanian Griman Creek Formation of Lightning Ridge,
New South Wales, was recently described and assigned to Mega-
raptoridae (Bell et al., 2016). Included in this specimen is the
proximal part of a manual ungual, tentatively identified in text
“ … as belonging to digit I or II based on its large size … ”
(Bell et al., 2016:477) and in the figure caption as “ … manual
ungual (?)I-2 … ” (Bell et al., 2016:478) but not definitively
assigned to the left or right side because of the incomplete
nature of the vascular grooves. Based on comparisons with
NMV P239464 and Australovenator (Hocknull et al., 2009;
White et al., 2012), we agree that this specimen represents a
megaraptorid manual ungual I-2 and suggest that it is from
the left side, based on the relative positions and trajectories of
the vascular grooves. Consequently, we compare it with NMV
P239464 herein. Although the proximal end of LRF 100–106
ungual is substantially taller (68 mm) than NMV P239464 (52
mm) and Australovenator (59.84 mm), it is not as large as an
incomplete manual ungual II-3 (NMV P186153) from the
upper Strzelecki Group of the Gippsland Basin (Victoria),
assigned to Neovenatoridae by Benson et al. (2012). Morpho-
logically, NMV P186153 is almost identical to manual ungual
II-3 of Australovenator and is herein referred to Megaraptori-
dae. The proximal end of NMV P186153 is less than two-
thirds complete but still almost as tall (58 mm) as that of
manual ungual II-3 in Australovenator (White et al., 2012).
Thus, it is unlikely that LRF 100–106 represents the
largest known Australian theropod (contra Bell et al., 2016,
pp. 473, 483).
Specimen NMV P239464 is strongly compressed mediolater-
ally: the ratio of the dorsoventral height to the mediolateral
width of the proximal end is 2.42. Values of this ratio equal to
or greater than 2.3 were regarded as a neovenatorid synapomor-
phy (Benson et al., 2010). However, although the value of this
ratio in Megaraptor exceeds this value (2.75; Benson et al.,
2010), that of Australovenator falls just short (2.29; White
et al., 2012). Similarly, in LRF 100–106, the value of this ratio
is lower than 2.3 (2.19; Bell et al., 2016), although this is pre-
sumably a consequence of the incomplete preservation of its
extensor tubercle.
The bone texture of the proximal articular surface of NMV
P239464 is smooth. However, that of its immediate surrounds—
proximal to the vascular grooves—is characterized by axially
oriented striations and is rougher to the touch than the bone of
the blade of the claw, which is smooth and only finely (albeit hap-
hazardly) striated.
The ginglymoid proximal articular surface of NMV P239464
consists of two shallowly concave articular facets, separated by
a narrow, dorsoventrally oriented ridge (Fig. 6F–G). The lateral
articular facet is taller dorsoventrally (37 vs. 33 mm) and nar-
rower mediolaterally (10 vs. 12 mm) than the medial facet, see-
mingly distinguishing this ungual from that of Australovenator
(Hocknull et al., 2009; White et al., 2012) in which the inverse
is true; however, the proximal articular surface of NMV
P239464 has suffered some damage. Dorsal to the articular
facets, a bulbous extensor tubercle is present; ventrally, the tuber-
cle tapers and merges with the narrow ridge that lies between the
articular facets, as in Australovenator (White et al., 2012). Ventral
and distal to the articular surface is a sulcus that effectively rep-
resents the unification of the two vascular grooves at the proximal
 Poropat et al.—Cretaceous megaraptorids from Australia (e1666273-12)

FIGURE 6. Megaraptoridae cf. Australovenator wintonensis, NMV P239464, right manual ungual I-2 from the Eric the Red West site, Eumeralla For-
mation, Victoria, Australia, in A, dorsal, B, C, lateral, D, E, medial, F, G, proximal, H, I, ventral, and J, distal views. A, B, D, F, H, and J are photographs,
whereas C, E, G, and I are 3D renders derived from CT scan data. Scale bar equals 50 mm.
 Poropat et al.—Cretaceous megaraptorids from Australia (e1666273-13)
TABLE 5. Measurements (in mm) for NMV P239464 and NMV
P252715A, manual unguals.
Manual ungual I-2 Manual ungual III-4
Dimension (NMV P239464) (NMV P252715A)
Proximodistal length 201 73
(outer curve)
Proximodistal length 166 60.5
(straight line from tip to
base)
Proximal mediolateral 21.5 12
breadth
Proximal dorsoventral 52 (excluding 31
height tubercle)
end (the ‘ventral mediolateral groove’ of White et al., 2012; the
‘deep sulcus’ of Bell et al., 2016), and immediately proximal to
this a roughened, shallow concavity is present, as is the case in
Australovenator (the ‘rugose area’ of White et al., 2012). Distal
to the aforementioned sulcus, a prominent, median flexor tuber-
cle is present (Fig. 6H, I), similar to that of Australovenator
(White et al., 2012), Megaraptor (Novas, 1998; Calvo et al.,
2004; Porfiri et al., 2007), and LRF 100–106 (Bell et al., 2016).
The flexor tubercle is flanked on either side by shallowly
concave flexor facets, with the medial flexor facet (Fig. 6E)
deeper than the lateral one (Fig. 6C). Specimen NMV P239464
lacks a ventral flexor facet; this distinguishes it from Australove-
nator (White et al., 2012).
The dorsal surface of NMV P239464 (Fig. 6A) is more broadly
and symmetrically convex than the ventral one, which tapers to a
keel that projects slightly proximomedially–distolaterally (Fig.
6H). The same keel is visible on the ventral surface of manual
ungual I-2 in Australovenator and Megaraptor (Novas, 1998;
Porfiri et al., 2007; White et al., 2012); however, Novas et al.
(2016:57) erred in stating that “the ventral keel gradually dis-
places laterally, joining the lateral margin of the claw on its
most proximal portion.” Presuming that the holotype manual
ungual I-2 of Australovenator (Hocknull et al., 2009; White
et al., 2012) is from the right side and that the Megaraptor holo-
type manual ungual I-2 is from the left—which seems certain in
FIGURE 7. Megaraptoridae cf. Australovenator wintonensis, NMV
P252715A, left manual ungual III-4 from the Eric the Red West site,
Eumeralla Formation, Victoria, Australia. A, photograph of NMV
P252715A in situ in ‘The Block’ (NMV P252715). B–G, 3D renders of
NMV P252715A derived from CT scan data, in B, proximal, C, dorsal,
D, medial, E, ventral, F, distal, and G, lateral views. Scale bar equals 20
mm.
light of the morphology of the unguals that constitute part of an
articulated right manus referred to Megaraptor (Calvo et al.,
2004; S.F.P., pers. observ.), the ventral keel is flush with the
medial margin at the proximal termination and therefore projects
proximomedially–distolaterally.
As in Australovenator (Hocknull et al., 2009; White et al.,
2012) and Megaraptor (Novas, 1998; Calvo et al., 2004), the
medial surface of NMV P239464 (Fig. 6D, E) is flatter than
the more strongly dorsoventrally convex lateral surface (Fig.
6B, C). In addition, the lateral vascular groove is deeper than
the medial one in both NMV P239464 and Australovenator
(Hocknull et al., 2009; White et al., 2012). The curvature of
the vascular groove on the medial surface essentially follows
the curvature of the ungual itself when viewed medially, such
that it terminates slightly dorsal to the mid-height of the distal
tip. Although the vascular groove on the lateral surface
follows a similar trajectory, it does so further dorsally such
that it terminates on the dorsal surface of the distal tip. The
distal end of the lateral vascular groove cannot be observed
in lateral view but is visible in distal view (Fig. 6J). The holotype
manual ungual of Megaraptor, originally described as a right
pedal ungual (Novas, 1998), was later shown to be a manual
ungual (Calvo et al., 2004) from the left side (Lamanna,
2004). As such, although it was originally claimed (Novas,
1998) and later reiterated (Calvo et al., 2004; Bell et al., 2016;
Aranciaga Rolando et al., 2019) that the medial vascular
groove was situated further dorsally on the ungual than the
lateral vascular groove, the inverse is true: the lateral vascular
groove was situated further dorsally than the medial one
(Calvo et al., 2004:fig. 9a).
Several nutrient foramina are preserved on NMV P239464,
notably at the proximal end of each vascular groove. The
medial surface of manual ungual I-2 in Megaraptor preserves a
nutrient foramen in exactly the same position as observed in
NMV P239464 (Novas, 1998); however, the lateral side does not
(S.F.P., pers. observ.).
Left Manual Ungual III-4
The theropod manual ungual NMV P252715A preserved on
‘The Block’ (Poropat et al., 2018) is currently only partially
exposed (Fig. 7A). However, CT scans of the specimen demon-
strate that it is almost identical to, albeit slightly smaller than
and a mirror image of, the right manual ungual III-4 of Austra-
lovenator (Fig. 7B–E; Table 5). The description herein is based
mostly on the CT data. The ginglymoid proximal end is ovate,
with the long axis vertical and the dorsal margin more
tapered than the ventral one (Fig. 7A). The ratio of dorsoven-
tral height to mediolateral width of the proximal end of NMV
P252715A is 2.58, which is higher than the value of this ratio
for the corresponding element in Australovenator (White
et al., 2012). As in Australovenator, the medial articular facet
is taller than the lateral (White et al., 2012). The extensor tuber-
cle is weakly developed, whereas the flexor tubercle is pro-
nounced and dorsoventrally deep, albeit morphologically
simple (lacking the complex of tubercles and fossae present in
manual phalanx I-2). It is possible that the height of this
feature has been exaggerated through taphonomic distortion—
the CT data suggest that this part of the ungual is broken—or
because of damage from rock saw cuts.
Both medial and lateral vascular grooves are present on NMV
P252715A, with the medial groove (and the ridge that floors it)
more prominent. The grooves run parallel with each other, not
showing asymmetrical paths as in manual phalanx I-2. In all of
these respects, NMV P252715A corresponds well with the mor-
phology of manual phalanx III-4 in Australovenator (White
et al., 2012). In Megaraptor, the lateral vascular groove was
reported to be absent on this element (Calvo et al., 2004);
 Poropat et al.—Cretaceous megaraptorids from Australia (e1666273-14)

FIGURE 8. Megaraptoridae cf. Australovenator wintonensis, NMV P253701, right astragalus from the Eric the Red West site, Eumeralla Formation,
Victoria, Australia, in A, B, lateral, C, D, anterior, E, F, medial, G, H, posterior, I, J, proximal, and K, L, distal views. A, C, E, G, I, and K are photo-
graphs, whereas B, D, F, H, J, and L are 3D renders derived from CT scan data. Abbreviation: PALP, proximal anterolateral process. Scale bar equals 20
mm.

however, the lateral surface of this specimen is not completely
preserved (S.F.P., pers. observ.), meaning that it is likely that it,
too, had a lateral vascular groove.
Right Astragalus
The right astragalus NMV P253701 is almost complete (Fig. 8;
Table 6), lacking only the apex of the ascending process, parts of
the posterior margin of the base, and the apex of the proximal
anterolateral process (the ‘cranio-proximal process’ of Hocknull
et al., 2009; White et al., 2013b). Although it was clearly not
fused to the calcaneum or fibula, the relatively small size of
NMV P253701 suggests that it represents an immature individual;
thus, whether or not fusion to either of these elements took place
at osteological maturity cannot be determined. In overall mor-
phology, NMV P253701 is very similar to the astragalus of
Australovenator (Hocknull et al., 2009; White et al., 2013b),
despite being only 40% the size (mediolateral width of base in
NMV P253701 = 41.63 mm vs. 105 mm in AODF 604; White
et al., 2013b).
The astragalar body is mediolaterally concave and proximodis-
tally convex, giving it a saddle-like appearance in anterior view. In
distal view (Fig. 8K, L), the maximum depth of the concavity lies
distinctly lateral to the midline. The distal condyles are markedly
 Poropat et al.—Cretaceous megaraptorids from Australia (e1666273-15)
TABLE 6. Measurements (in mm) of NMV P253701, a right astragalus.
NMV
Dimension P253701
Total proximodistal height 39.8*
Base Anteroposterior length 20.6
Mediolateral width 41.6
Proximodistal height (measured with the 12.5
dorsal surfaces of the posterior and
anterior margins aligned)
Ascending Proximodistal height (measured from 21.4*
process dorsal surface of the medial edge of the
base, with the process vertical)
Anteroposterior length (measured 5.2
proximal to the proximal anterolateral
process)
Mediolateral width (measured at base of 31.8
process)
Ratio of proximodistal height (ascending process): 1.712*
proximodistal height (base)
An asterisk (*) indicates a measurement made on an incomplete
structure/surface.
deflected anteriorly, as in other tetanurans (Carrano and
Sampson, 2008; Carrano et al., 2012). The medial margin is
broadly rounded when viewed anteriorly (Fig. 8C, D) or poster-
iorly (Fig. 8G, H), whereas the lateral margin is essentially
straight and vertical, providing an articular surface for the calca-
neum (Fig. 8A, B). The articular surface for the calcaneum is gen-
erally smooth but bears several proximodistally oriented subtle
ridges and grooves (presumably for the adherence of a cartilagi-
nous cap). A horizontal groove is present on the proximal part of
the anterior surface of the astragalar base (Fig. 8C, D); this
feature is commonly present in Coelurosauria (Brusatte et al.,
2014) and is also seen in Australovenator (Hocknull et al., 2009;
White et al., 2013b) and Fukuiraptor (Azuma and Currie,
2000). No distinct tendoneal groove is present on the anterior
surface of the base, thereby distinguishing NMV P253701 from
many alvarezsaurian and paravian theropods (Brusatte et al.,
2014). Although it is broken, it is clear that the astragalar body
of NMV P253701 originally possessed a proximal anterolateral
process (Fig. 8A–D), as in Australovenator (Hocknull et al.,
2009; White et al., 2013b) and Fukuiraptor (Azuma and Currie,
2000). The posteromedial part of the base of the astragalus was
inadvertently lost to a saw cut in the field (Fig. 8G–L); neverthe-
less, it is likely that relatively little was lost. Given that both pos-
terolateral and posteromedial crests are present on the astragalar
base of neotheropods generally (Carrano et al., 2012), and that
NMV P253701 possesses a posterolateral one, it is highly likely
that the missing posteromedial margin formed a crest as well.
The base of the ascending process in NMV P253701 is laterally
offset from the astragalar body, as is typical of most coelurosaurs
except Confuciusornis sanctus (Brusatte et al., 2014); Sinraptor
(Currie and Zhao, 1993) and Allosaurus (Madsen, 1976) do not
show this morphology. The apex of the ascending process is
incomplete but was clearly laminar, as in other tetanurans
(Carrano and Sampson, 2008; Carrano et al., 2012), and broad
mediolaterally. Furthermore, the ascending process of NMV
P253701 was substantially taller than the astragalar body, which
distinguishes it from several basal coelurosaurs (e.g., Coelurus
fragilis, Tugulusaurus faciles, Zuolong salleei), the basal tyranno-
sauroid Guanlong, therizinosaurs, and most alvarezsaurians (Bru-
satte et al., 2014). By contrast, an ascending process at least 1.6
times taller than the astragalar body unites NMV P253701 with
several theropods scored by Carrano et al. (2012), including the
spinosaurid Suchomimus (Sereno et al., 1998), the megaraptoran
Fukuiraptor (Azuma and Currie, 2000), and the megaraptorids
Aerosteon (Sereno et al., 2008) and Australovenator (Hocknull
et al., 2009; White et al., 2013b). The anterior surface of the
base of the ascending process (Fig. 8C, D) appears to lack the
fossa seen in many tetanurans (Carrano et al., 2012); however,
taphonomic distortion (particularly with respect to the angle of
the ascending process relative to the astragalar body) might
have subdued this feature. Nevertheless, several features can be
observed on the anterior surface: for example, at the base of
the ascending process, and near the base of the proximal antero-
lateral process, a sulcus is present (Fig. 8C, D). Medial to this,
another shallower but more mediolaterally elongate sulcus is
present. The medial half of the anterior surface of the ascending
process is mostly smooth; however, the lateral half is covered in
proximodistally oriented grooves that appear to radiate from a
subtle sulcus situated immediately medial to the anterolateral
process. The deepest such groove is the medial-most; immediately
lateral to it, close to its base, lies a nutrient foramen. By contrast,
the posterior surface of the ascending process is almost comple-
tely smooth, undulating only slightly either side of a proximodis-
tally elongate concavity (Fig. 8G, H). At the base of this concavity,
just proximal to the astragalar body on the posterior surface, lies a
round sulcus; however, it is unclear whether or not this is a natural
feature. Immediately lateral to this sulcus lies a nutrient foramen.
DISCUSSION
Megaraptorid theropods were a prominent component of Bar-
remian–Turonian ecosystems in Australia and Albian–Santonian
terrestrial ecosystems in South America (Tables S1, S7). By con-
trast, non-megaraptorid megaraptorans—the ?upper Barremian
Phuwiangvenator yaemniyomi (and possibly Vayuraptor non-
gbualamphuensis; Samathi et al., 2019), the Albian Fukuiraptor
kitadaniensis (Azuma and Currie, 2000; Currie and Azuma,
2006; Molnar et al., 2009), the Cenomanian Siats meekerorum
(Zanno and Makovicky, 2013), and the Turonian or younger Chi-
lantaisaurus tashuikouensis (Benson and Xu, 2008)—are from the
mid-Cretaceous of eastern Asia and western North America. This
suggests that Megaraptora were widespread before the Barre-
mian, but that Megaraptoridae had diverged by the Barremian
(Bell et al., 2016).
The manual ungual I-2 described herein highlights the remark-
able morphological conservatism of megaraptorid forelimbs—
previously implied by the ulna from Dinosaur Cove (Smith
et al., 2008)—across a vast geographic range (Australia + South
America) and throughout a significant geological time span
(lower Albian–Coniacian). This geographic range includes
several high paleolatitude sites, including Flat Rocks (∼78.31°S
at 120 Ma), Eric the Red West (∼75.99°S at 110 Ma), and Dino-
saur Cove (∼75.93°S at 110 Ma) in Victoria (Torsvik et al.,
2012; Van Hinsbergen et al., 2015). Although the evolutionary
pressure(s) underlying the development—and retention—of
these forelimbs is unknown, and despite the fact that no direct
evidence of megaraptorid predation has been reported to date,
prey acquisition might have been a significant contributing
factor. In the Cenomanian–lowermost Turonian Winton For-
mation, shed megaraptorid teeth are quite commonly found in
association with sauropod remains (White et al., 2015a)—both
adult and subadult (S.F.P. and M.A.W., pers. observ.), implying
that these theropods fed on sauropods either through direct pre-
dation or opportunistic scavenging. However, irrespective of their
trophic relationship in northeast Australia, megaraptorids evi-
dently could not have fed on sauropods in southeast Australia
because of the absence of the latter in the fauna (Benson et al.,
2012; Poropat et al., 2018). The Australian megaraptorid record,
including the new material described herein, demonstrates that
these theropods thrived both in the absence of sauropods in
late Early Cretaceous Victoria (Benson et al., 2012; Poropat
et al., 2018) and in their presence in the early Late Cretaceous
 Poropat et al.—Cretaceous megaraptorids from Australia (e1666273-16)
of New South Wales (Molnar and Salisbury, 2005) and Queens-
land (Coombs and Molnar, 1981; Molnar and Salisbury, 2005;
Hocknull et al., 2009; Poropat et al., 2015a, 2015b, 2016, 2017).
CONCLUSIONS
The discovery of new remains of Megaraptoridae in the Eumer-
alla Formation at the Eric the Red West site, Cape Otway, Vic-
toria, highlights the success of this enigmatic theropod group in
Australia during the mid-Cretaceous. Reappraisal of the sup-
posed ‘spinosaurid’ cervical vertebra from this site indicates
that it might pertain to Megaraptoridae. The discovery of a tren-
chant, mediolaterally compressed manual ungual I-2 demon-
strates that megaraptorids in southeast Australia had developed
the claws characteristic of this clade by at least the early
Albian. That Megaraptor (Turonian–Coniacian; Portezuelo For-
mation), which lived in Argentina tens of millions of years after
the Victorian megaraptorids, was equipped with a virtually iden-
tical manual ungual on each first digit implies that such claws con-
ferred a significant adaptive advantage on their owners, the
precise nature of which is unknown. Finally, the morphological
similarity between the early Albian megaraptorid remains from
Victoria and those of the stratigraphically younger megaraptorids
of New South Wales (Cenomanian) and Queensland (Cenoma-
nian–earliest Turonian) is suggestive of connectivity between
mid-Cretaceous faunas throughout eastern Australia. Further-
more, the presence of megaraptorids at several sites in southeast
Australia implies that, unlike their titanosauriform sauropod
cousins, they flourished at high latitudes.
ACKNOWLEDGMENTS
The authors would like to thank all of those who have been part
of the Dinosaur Dreaming crew at Eric the Red West;
F. Karakitsos, L. Kool, A. Tait, W. Turner, W. White, and
J. Wilkins for discovering and collecting the specimens described
herein; L. Kool and the late D. Pickering for preparing the speci-
mens; T. Ziegler (Melbourne Museum) for facilitating access to
these specimens and for discussion regarding the possible bite
marks; A. Gonzales and C. Paragnani for assistance with the
photographs; B. Freshwater (St Vincent’s Hospital, Melbourne)
for CT scanning the specimens; O. Panagiotopoulou (Monash
University) for assistance with the processing of the CT and syn-
chrotron data; The Paleontological Society for an Arthur James
Boucot Research Grant (awarded to S.F.P. in 2017), which facili-
tated firsthand observation of the Australovenator type specimen;
D. Elliott and T. Sloan (Australian Age of Dinosaurs Museum)
for allowing access to the Australovenator type specimen in
their care; the Winston Churchill Memorial Trust for a Churchill
Fellowship (awarded to S.F.P. in 2017) that enabled firsthand
observation of several Argentinian megaraptorid specimens;
and R. Coria and L. Coria (Museo Municipal Carmen Funes),
J. Calvo and J. Mansilla (Centro Paleontológico Lago Barreales),
J. Porfiri and D. dos Santos (Universidad Nacional del Comahue),
and R. Martínez and M. Luna (Universidad Nacional de la Pata-
gonia San Juan Bosco) for allowing S.F.P. to make firsthand obser-
vations of Argentinian megaraptorid specimens in their care. The
authors would also like to thank P. Bell (University of New
England), F. Agnolin (Museo Argentino de Ciencias Naturales
‘Bernardino Rivadavia’), an anonymous reviewer, and editor
L. Zanno for their constructive comments that greatly improved
the manuscript.
ORCID
Stephen F. Poropat http://orcid.org/0000-0002-4909-1666
Patricia Vickers-Rich http://orcid.org/0000-0003-1850-8067
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Submitted October 8, 2018; revisions received June 20, 2019;
accepted July 10, 2019.
Handling editor: Lindsay Zanno.