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A large heterocoelous cervical vertebra from the Late Cretaceous of Cruzy (Hérault, southern France) is described and referred to the giant bird Gargantuavis philoinos Buffetaut & Le Loeuff, 1998, confirming its avian nature. Gargantuavis appears to have been a long-necked bird with possibly a relatively small skull. Derived features such as heterocoely suggest that Gargantuavis was an advanced ornithuromorph, close to ornithurines.
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Geol. Mag.: page 1 of 4. c
Cambridge University Press 2012 1
New evidence of a giant bird from the Late Cretaceous of France
Centre National de la Recherche Scientifique, UMR 8538, Laboratoire de Géologie de l’Ecole Normale Supérieure,
24 rue Lhomond, 75231 Paris Cedex 05, France
Laboratoire de Géologie de Lyon, CNRS UMR 5276, 2 rue Dubois, 69622 Villeurbanne Cedex, France
(Received 15 February 2012; accepted 23 May 2012)
A large heterocoelous cervical vertebra from the Late
Cretaceous of Cruzy (Hérault, southern France) is described
and referred to the giant bird Gargantuavis philoinos
Buffetaut & Le Loeuff, 1998, confirming its avian nature.
Gargantuavis appears to have been a long-necked bird with
possibly a relatively small skull. Derived features such as
heterocoely suggest that Gargantuavis was an advanced
ornithuromorph, close to ornithurines.
Keywords: Aves, Late Cretaceous, France, Gargantuavis,
1. Introduction
The presence of a giant bird in the Late Cretaceous of
southern France was first reported by Buffetaut et al. (1995)
on the basis of a synsacrum fragment from Fox-Amphoux
(Var). Gargantuavis philoinos was erected by Buffetaut & Le
Loeuff (1998) on the basis of a synsacrum and parts of the
pelvis, from Campagne-sur-Aude (Aude), as the holotype,
with an incomplete femur from Villespassans (Hérault)
being referred to it because of anatomical congruence
of the femoral head. Since then, no additional material
referable to Gargantuavis had been identified, despite active
excavations at various sites in southern France, showing that
this bird is a rare component of the local Late Cretaceous
vertebrate assemblages (Buffetaut & Le Loeuff, 2010). The
discovery of a well-preserved cervical vertebra of a very
large bird in the Upper Cretaceous of Cruzy (Hérault) is
therefore worth reporting, especially in the context of a
recent controversy about the avian or pterosaurian nature
of Gargantuavis (Mayr, 2009; Buffetaut & Le Loeuff,
2010; Buffetaut, 2011a). The new specimen is referred to
Gargantuavis philoinos and provides additional evidence
about its osteology.
Institutional abbreviations: MC – Musée de l’Association
Culturelle, Archéologique et Paléontologique de l’Ouest
Biterrois, Cruzy (Hérault, France); MDE – Musée des
Dinosaures, Espéraza (Aude, France).
2. Geographical and geological setting
The vertebra was discovered in the course of excavations
at the Montplo-Nord locality, of late Campanian–early
Maastrichtian age as indicated by its vertebrate assemblage,
near the village of Cruzy (Hérault). There, continental red
Author for correspondence:
marls interbedded with sandstone lenses corresponding to
a floodplain environment yield abundant vertebrate remains.
The faunal assemblage has not yet been described in detail but
is generally similar to that from the nearby Massecaps locality
(Buffetaut et al. 1999). It includes freshwater bivalves, fish
(lepisosteids), turtles (the bothremydid Foxemys), crocodili-
ans, pterosaurs (azhdarchids) and dinosaurs (titanosaurid
sauropods, dromaeosaurid and abelisaurid theropods, nodo-
saurid ankylosaurs, rhabdodontid ornithopods). Previously
reported Gargantuavis specimens (Buffetaut et al. 1995;
Buffetaut & Le Loeuff, 1998) are from localities of similar
age and general geological setting in southern France
(Provence and Languedoc).
3. Description
Osteological nomenclature follows Baumel & Witmer
(1993). The bird vertebra from Montplo-Nord (MC-MN 478,
Fig. 1) is generally well preserved, having suffered only
slight distortion. Although the apophyses on the left side
are broken, those on the right side are in good condition,
despite some abrasion of the edges. As shown by vast calcite-
filled inner spaces revealed by the breaks on the left side,
the bone was heavily pneumatized. A pneumatic foramen
is visible on both sides, low on the neural arch, between
the bases of the pre- and postzygapophyses (Fig. 1b). Such
a foramen is commonly present in cervical vertebrae of
modern birds (O’Connor, 2004). There is no indication of
pneumatic foramina on the centrum. The centrum is low and
elongate, with a ventral face that is flat caudally and deeply
concave cranially between the Processi carotici, which are
well developed, forming well-defined subtriangular facets
that project medially (Fig. 1c). The caudal articular surface
of the centrum, located posterior to the caudal margin of the
postzygapophyses, is markedly saddle-shaped, being concave
craniocaudally and convex transversally, and narrower than
high (Fig. 1b, d). Unlike the condition in most birds, the
ventral part of this articular surface is remarkably narrow,
without any significant transversal expansion (Fig. 1c). The
cranial articular surface is strongly concave transversally and
convex craniocaudally (Fig. 1e). The vertebra from Montplo-
Nord thus shows the distinctive heterocoelous condition that
is found in modern birds and some Cretaceous forms (Marsh,
1879), most of which belong to Ornithuromorpha, although
the phylogenetic distribution of this character seems unclear
(O’Connor, Chiappe & Bell, 2011). The better preserved
right side of the vertebra shows an Ansa costotransversaria
enclosing the Foramen transversarium for the passage of
blood vessels. The Processus costalis is broken caudally.
The Zygapophysis cranialis is large, with a fairly deep Downloaded: 30 Jul 2012 IP address:
Figure 1. (Colour online) Large bird vertebra, referred to Gargantuavis philoinos, from the Late Cretaceous of Cruzy (Hérault, France),
MC-MN478. Photographs (left) with explanatory drawings (right), in dorsal (a), right lateral (b), ventral (c), caudal (d) and cranial (e)
views. Ac – Ansa costotransversaria; Cto – Cristae transversae obliquae; Fp – Foramen pneumaticum; Ft – Foramen transversarium;
Fv – Foramen vertebrale; Pc – Processi carotici; Ps – Processus spinosus; Td – Torus dorsalis; Zca – Zygapophysis caudalis; Zcr –
Zygapophysis cranialis. Scale bar =10 mm.
elongate depression on its lateral surface (Fig. 1a). The
zygapophysial facet is oval, flat, and slopes at about 45.
The Zygapophysis caudalis is incompletely preserved, part of
its dorsal margin being damaged. The zygapophysial facet is
large and somewhat concave. There is a well-developed Torus
dorsalis (although much less developed than the epipophyses
of non-avian theropods). The Processus spinosus is tall
(although its distal end is broken) and short craniocaudally
(Fig. 1b). Its caudal margin is sloping, whereas its cranial
margin is vertical. Both its cranial and caudal faces are
concave, probably for the insertion of tendons. Posterior to
the Processus spinosus, a long and narrow median groove is
limited laterally by the Cristae transversae obliquae, which
merge caudally with the zygapophyses (Fig. 1a). Anterior
to the Processus spinosus, a broad shelf extends onto the
medial side of the zygapophyses and overhangs the Foramen
vertebrale, which is cranially low, with an oval outline, and
caudally higher and narrower.
4. Identification and comparisons
MC-MN 478 clearly belongs to a bird, as shown by its
heterocoelous condition, which is an avian character (Marsh,
1879). Incipiently heterocoelous cervical vertebrae have
been reported in troodontid and dromaeosaurid dinosaurs
(Xu & Norell, 2004), but MC-MN 478 shows an advanced
heterocoely that seems to have no equivalent in any known
non-avian dinosaur. Similarities with troodontid vertebrae
(Makovicky & Norell, 2004) include the presence of
Processi carotici and a distinct Torus dorsalis on the postzy-
gapophyses, but the markedly saddle-shaped, posteriorly
projecting caudal articular surface of the centrum of MC-MN
478 has no equivalent in troodontids. The lack of pneumatic
openings on the centrum also seems to separate MC-MN 478
from deinonychosaurs. Its elongate centrum, the presence of
a Foramen transversarium and of well-developed Processi
carotici, and the complete absence of a hypapophysis show
that MC-MN 478 belongs to the middle part of the neck.
The articular surfaces of MC-MN 478 show that in this
region dorsal bending was possible, whereas ventral bending
must have been nearly impossible because the long saddle-
shaped caudal articular surface was blocked by the strongly
developed and medially projecting Processi carotici. This
indicates that the vertebra belongs to Boas’s section II of
the neck, in which only dorsal flexion is possible (Boas,
1929). However, because the various sections of the neck as
defined by Boas (1929) do not contain the same number of
vertebrae in all groups of birds, the exact position of MC-MN
478 in the vertebral column cannot be precisely determined.
Lateral bending must have been extremely limited because
the cranial articular surface is deeply concave transversally
and must have severely restricted lateral movements of the
elongate caudal articular area of the vertebra immediately
anterior to it.
Comparison with Gargantuavis philoinos is not easy
because there are no cervical vertebrae in the original
material. In size, however, MC-MN 478 is compatible with
the type synsacrum of Gargantuavis philoinos (MDE-C3–
525), as suggested by comparisons with vertebral columns
of modern large terrestrial birds. In ratites there is a gradual
decrease in the diameter of the vertebral centra from the
synsacrum to the cervical vertebrae (Mivart, 1874, 1877):
the width of the articular face of an ‘intermediate’ cervical
vertebra is roughly one third that of the cranial articular
face of the synsacrum. Assuming a comparable decrease in
Gargantuavis, the 40 mm wide cranial articular face of the
type synsacrum is in good agreement with the 13 mm wide
articular faces of the centrum of MC-MN 478. The vertebra
from Montplo-Nord is roughly the size of a corresponding
vertebra of a cassowary (Casuarius), in good agreement
with size estimates based on the original giant bird material
from southern France, which suggested a cassowary-sized Downloaded: 30 Jul 2012 IP address:
(Buffetaut et al. 1995) or ostrich-sized (Buffetaut & Le
Loeuff, 1998) bird. Because MC-MN 478 is compatible in
size with previously described specimens of Gargantuavis
philoinos and has been found in sediments of the same age
in the same geographical area, it seems reasonable to refer
it to this taxon, rather than postulating the existence of two
distinct giant birds in the local Late Cretaceous vertebrate
Comparisons with other Cretaceous birds are limited
because details of vertebral structure are available for few
of them (many of the more complete Cretaceous birds are
known from specimens that are crushed flat on slabs of rock).
Buffetaut & Le Loeuff (1998) suggested possible affinities
between Gargantuavis and Patagopteryx deferrariisi, a hen-
sized flightless bird from the Late Cretaceous of Patagonia
(Bonaparte & Alvarenga, 1992; Chiappe, 2002), because
of similarities in pelvic construction. However, MC-MN
478 shows little resemblance to the cervical vertebrae of
Patagopteryx, which have a low neural spine and a caudal
articular area that does not project beyond the level of the
postzygapophyses (Chiappe, 2002). Similarly, MC-MN 478
differs from enantiornithine cervical vertebrae from the Late
Cretaceous of Argentina (Walker & Dyke, 2009) in the strong
posterior elongation of its caudal articular region. MC-MN
478 appears to be more markedly heterocoelous than the
cervical vertebrae of Patagopteryx and enantiornithines.
The cervical vertebra from Montplo-Nord is clearly
distinct from those of the Late Cretaceous Ichthyornis,
which are not fully heterocoelous (Marsh, 1879, 1880;
Clarke, 2004). In the large marine Late Cretaceous bird
Hesperornis, similarly positioned cervical vertebrae are fully
heterocoelous (Marsh, 1880), but differ from MC-MN 478 in
their transversally narrow, anteroposteriorly elongate neural
spine and in their caudal articular surface, which is more
expanded transversally and does not project much beyond
the level of the postzygapophyses.
Comparisons with large extinct birds from the Cenozoic do
not reveal notable similarities. MC-MN 478 differs from the
much more robust and less elongate cervical vertebrae of the
Early Tertiary gastornithids (Gervais, 1873; Lemoine, 1878;
Matthew & Granger, 1917). This confirms that Gargantuavis
and gastornithids are not closely related (Buffetaut, 2002).
The Dromornithidae from the Cenozoic of Australia (Murray
& Vickers-Rich, 2004) have short cervical vertebrae with
low neural spines, unlike MC-MN 478. In the mainly South
American Phorusrhacidae, there is no posterior elongation of
the caudal articular area and the neural spine is low (Sinclair
& Farr, 1932).
MC-MN 478 is somewhat reminiscent of modern ratites
(Mivart, 1874, 1877) in the elongation of the centrum. Its
proportions are more reminiscent of the cervical vertebrae
of the emu (Dromaius) and the cassowary (Casuarius)than
of those of the ostrich (Struthio) and the rhea (Rhea), which
are more elongated (Mivart, 1877). However, its posteriorly
projecting and narrow caudal articular region and its tall
and anteroposteriorly compressed Processus spinosus are
significant differences.
To sum up, MC-MN 478 is not especially reminiscent of
any Cretaceous bird in which the cervical vertebrae are well
known, nor does it closely resemble the vertebrae of more
recent large birds. This may suggest endemic evolution on the
relatively isolated Late Cretaceous Ibero-Armorican island.
5. Conclusions
It was recently suggested that the remains attributed to
Gargantuavis in fact belong to a giant pterosaur (Mayr,
2009). This is untenable because of highly significant
differences between the specimens referred to Gargantuavis
and the corresponding elements in pterosaurs (Buffetaut
& Le Loeuff, 2010; Buffetaut, 2011a; Mourer-Chauviré
et al. 2011). The typically avian vertebra from Montplo-
Nord, which confirms the presence of a very large bird in the
Late Cretaceous of France, can in all likelihood be referred
to Gargantuavis philoinos and provides new evidence about
its anatomy. Its size and morphology indicate a bird with
probably a long, rather slender neck, unlike the Early Tertiary
gastornithids, which had a robust and relatively short neck
(and a very large head). The shape of the articulations
suggests that, at least at the level of this vertebra, bending of
the neck was very limited, except in a dorsal direction.
The vertebra from Montplo-Nord shows an advanced
heterocoelous condition that distinguishes it from various
groups of basal birds in which heterocoely apparently was
not fully developed (O’Connor, Chiappe & Bell, 2011).
Buffetaut & Le Loeuff (1998) concluded that Gargantuavis
philoinos could be placed within Ornithothoraces and was
more advanced than Enantiornithes because of femoral
morphology, but was probably not an ornithurine because
of the large size of the acetabulum (possibly more apparent
than real because in the type of Gargantuavis philoinos
the ilium is incompletely preserved). The cervical vertebra
from Cruzy supports placement of Gargantuavis within
Ornithuromorpha but does not suggest close proximity to
Patagopteryx. In terms of heterocoely, it is more advanced
than Ichthyornis, which may suggest that it is close to
or within Ornithurae (defined as ‘a dichotomy between
Neornithes and Hesperornithiformes’, and as the sister-group
of Ichthyornis, by O’Connor, Chiappe & Bell, 2011). This
also raises the question of the acquisition of heterocoely
in Cretaceous birds, which is not fully understood and
may have involved some parallel evolution. However, a
more precise assessment of the phylogenetic position of
Gargantuavis philoinos requires a re-examination of all
specimens referred to this taxon, which is beyond the scope of
this paper. With the vertebra from Montplo-Nord confirming
its avian status, Gargantuavis philoinos remains the only
giant ground bird hitherto reported from the Late Cretaceous,
since Samrukia nessovi, a purported huge bird from the
Santonian of Kazakhstan (Naish et al. 2012), is in fact a
pterosaur (Buffetaut, 2011b).
The ecological place of Gargantuavis philoinos in the
ecosystems of the Late Cretaceous Ibero-Armorican island
(Pereda-Suberbiola, 2009) remains uncertain. Contrary to a
suggestion by Buffetaut & Le Loeuff (1998), it does not
seem to have occupied an ecological niche similar to that
of ornithomimosaurs in other parts of the world, because
it appears to have been a graviportal rather than cursorial
form (Buffetaut & Le Loeuff, 2010). The new vertebra from
Cruzy, in addition to the previously described material, may
suggest a bird somewhat similar in proportions to some of
the medium-sized graviportal moas of New Zealand, such as
Pach y o r n i s (Bunce et al. 2009), although lack of evidence
about the skull limits the reliability of the comparison.
Be that as it may, the new specimen confirms that a very
large ground bird co-existed with dinosaurs on the Ibero-
Armorican island. In the present state of our knowledge,
no such giant Late Cretaceous birds have yet been reported
from other parts of the world, including areas with excellent
Late Cretaceous vertebrate records, and this suggests that
the occurrence of Gargantuavis in France probably reflects
endemicity (Pereda-Suberbiola, 2009).
Acknowledgements. This work was supported by the
INTERRVIE Programme of the Centre National de la
Recherche Scientifique and by the Association Culturelle, Downloaded: 30 Jul 2012 IP address:
Archéologique et Paléontologique de l’Ouest Biterrois,
whose members are thanked for their help in the field. We are
grateful to Jean-Claude Guilhaumon (Cruzy) for permission
to dig on his land. Thanks to Christine Lefèvre for access
to material in the comparative anatomy collection of the
Muséum National d’Histoire Naturelle, Paris, and to Jean Le
Loeuff (Musée des Dinosaures, Espéraza) for access to the
type material of Gargantuavis philoinos. Federico Agnolin
and Gareth Dyke provided useful reviews.
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... The Montplo-Nord locality has already yielded a cervical vertebra (Buffetaut & Angst, 2013) and pelvic elements (Buffetaut & Angst, 2016b) of Gargantuavis philoinos. Although the specimens were not found in direct association, they come from the same sedimentary layer and were found a short distance from each other and may belong to a single individual. ...
... Mainly on the basis of pelvic and femoral characters, Buffetaut & Le Loeuff (1998) suggested that it was more advanced than enantiornithines but less derived than ornithurines and noted possible similarities with Patagopteryx deferrarisii, from the Late Cretaceous of Argentina. Because of the highly heterocoelous condition of the cervical vertebra from Montplo-Nord, Buffetaut & Angst (2013, 2016a considered that Gargantuavis could be placed within Ornithuromorpha (the group that includes all birds more advanced than Enantiornithes), was not very closely related to Patagopteryx, and was close to or within Ornithurae (the group that includes Neornithes (modern birds) and some more basal Cretaceous groups such as Hesperornithiformes and Ichthyornithiformes). Beyond confirming some previous observations (e.g. the absence of a posterior trochanter, which separates it from other Cretaceous birds such as enantiornithines), the wellpreserved femur from Montplo-Nord provide important new evidence about the systematic position of Gargantuavis. In particular, the morphology of the distal end, the evolutionary significance of which was highlighted by Elzanowski (2008), shows that this bird was more advanced than many other Mesozoic avians. ...
... Whether this groove is an original feature of the bone or an artefact is unclear. However that may be, direct comparison between MC MN1335 and Patagopteryx femora confirms what was already suggested by the cervical vertebra from Montplo-Nord (Buffetaut & Angst, 2013), viz. that Gargantuavis is not especially closely related to Patagopteryx. ...
... The giant bird Gargantuavis philoinos Buffetaut and Le Loeuff, 1998 is known from a limited number of skeletal elements from a small number of localities in southern France (Buffetaut et al., 2015a) and northern Spain (Buffetaut et al., 2015b). One of these localities is Montplo-Nord, near the village of Cruzy (H erault), which has yielded a cervical vertebra (Buffetaut & Angst, 2013). In the present paper, we describe additional Gargantuavis material from Montplo-Nord, consisting of an incomplete left ilium and a synsacrum fragment. ...
... The new specimens from Montplo-Nord thus confirm that the pelvis of Gargantuavis was extensively pneumatised, with well-developed pneumatic spaces in both the synsacrum, as already noted in specimens from Fox-Amphoux (Buffetaut et al., 2015a), and the ilium. Pneumatisation is also present in the cervical vertebra from Montplo-Nord described by Buffetaut and Angst (2013) (2009) distinguished several pneumaticity levels in birds, from reduced to hyperpneumatic. Pneumatisation of the pelvis occurs in the 'expanded' and 'hyperpneumatic' patterns. ...
... In addition, the occurrence at Montplo-Nord of pelvic elements clearly referrable to Gargantuavis philoinos supports the attribution to that taxon of the large avian cervical vertebra described from that locality by Buffetaut and Angst (2013). ...
A synsacrum fragment and an incomplete ilium of the giant bird Gargantuavis philoinos are described from the upper Campanian/lower Maastrichtian Montplo-Nord locality at Cruzy (Hérault, southern France). The ilium provides new information about the pelvic morphology of Gargantuavis. Both the synsacrum and ilium are extensively pneumatised. This extensive pneumatisation may be a plesiomorphic feature, but may also have had a weight-reduction function in a large flightless bird.
... On the basis of more complete material, including a partial pelvis and a femur, found at other localities farther west (Bellevue, Aude and Combebelle, Hérault), the taxon Gargantuavis philoinos was erected by . More recently, a large cervical vertebra from the Montplo-Nord locality at Cruzy (Hérault) was referred to Gargantuavis philoinos (Buffetaut & Angst, 2013). Despite claims to the contrary (Mayr, 2009), the avian nature of Gargantuavis is clearly supported by osteological (Buffetaut & Le Loeuff, 2010) and histological (Chinsamy et al., 2014) evidence. ...
... The acetabulum appears to be rather large, which is a basal character in birds (O'Connor et al., 2011), but it is difficult to really estimate its size relative to that of the ilium, since no complete ilia have yet been found. The advanced heterocoelous cervical vertebra from Cruzy suggests a position within Ornithuromorpha and close to Ornithurines (Buffetaut & Angst, 2013), but the pelvis, as mentioned above, shows various basal features. It should be admitted that much remains obscure about Late Cretaceous basal ornithuromorphs, and it may be difficult to get a better idea of the systematic position of the probably endemic (Buffetaut & Angst, 2013) Gargantuavis until more complete material is found. ...
... The advanced heterocoelous cervical vertebra from Cruzy suggests a position within Ornithuromorpha and close to Ornithurines (Buffetaut & Angst, 2013), but the pelvis, as mentioned above, shows various basal features. It should be admitted that much remains obscure about Late Cretaceous basal ornithuromorphs, and it may be difficult to get a better idea of the systematic position of the probably endemic (Buffetaut & Angst, 2013) Gargantuavis until more complete material is found. ...
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Two incomplete pelves of the giant bird Gargantuavis philoinos are described from Late Cretaceous deposits at Fox-Amphoux (Var, southeastern France). They consist of synsacra with attached parts of the ilia. One of them has undergone considerable dorsoventral compression, which makes it very similar in appearance to the holotype pelvis of Gargantuavis philoinos from Campagne-sur-Aude (Aude, southern France). The second specimen has suffered some lateral distortion but is uncrushed dorsoventrally. Because of this, its avian characters (including an arched synsacrum and widespread pneumatisation) are especially clear. These new specimens confirm the avian nature of Gargantuavis and reveal new details about its pelvic anatomy, but provide little new evidence about its systematic position within Aves. The geographical distribution and general rarity of Gargantuavis are discussed.
... Another femur from Cruzy was figured by Angst and 61 and differs from the specimen from Villespassans in its smaller size and 62 more slender overall shape. From Cruzy, which also yielded pelvic remains, a referred 63 cervical vertebra is known (Buffetaut and Angst, 2013). 64 The referred femora ; and the 65 cervical vertebra (Buffetaut and Angst, 2013) cannot be assigned to G. philoinos with 66 certainty, since these bones were not found in direct association with pelvic elements. ...
... From Cruzy, which also yielded pelvic remains, a referred 63 cervical vertebra is known (Buffetaut and Angst, 2013). 64 The referred femora ; and the 65 cervical vertebra (Buffetaut and Angst, 2013) cannot be assigned to G. philoinos with 66 certainty, since these bones were not found in direct association with pelvic elements. The The synsacrum includes nine vertebrae, which are fully co-ossified (Fig. 3C). ...
We describe a well-preserved pelvis from the Maastrichtian Sanpetru Formation of the Hateg Basin in Romania. The fossil closely resembles the pelvis of Gargantuavis philoinos from the Ibero-Armorican Peninsula, but differs in a smaller size and a few morphological features. It constitutes the first record of Gargantuavis outside the Ibero-Armorican Island and is more complete than any of the previously known Gargantuavis pelves. The new fossil allows the recognition of characteristics previously unknown for Gargantuavis. These include the presence of large supratrochanteric processes, the absence of a widened midsection of the synsacrum (which indicates the absence of a glycogen body), and the absence of fusion between the pelvic bones at the level of the acetabulum. The latter two features suggest that Gargantuavis is not closely related to the Ornithurae and the taxon may even fall outside the Ornithothoraces, the clade including Enantiornithes and Ornithuromorpha. Recognition of Gargantuavis in the fauna of the Hateg Island is of particular significance, because various theropods have been described from the Upper Cretaceous of Romania. The Romanian pelvis is of similar size to Elopteryx nopcsai, which was described as avian and is based on hindlimb elements, and it also shows some similarities to the pelvis of the unusual theropod Balaur bondoc. The new fossil furthermore disproves the hypothesis that the flight capabilities of Gargantuavis were lost in an insular environment of the Ibero-Armorican Island, and raises the possibility that Gargantuavis, Elopteryx, and Balaur belong to a distinctive theropod clade of the Late Cretaceous European archipelago.
... Angst). cassowary (Buffetaut and Angst, 2013). The first evidence of this bird was a synsacrum fragment from south-eastern France (Buffetaut et al., 1995). ...
... Subsequently, the taxon Gargantuavis philoinos was erected on the basis of a partial pelvis and an incomplete femur from other localities in southern France by Buffetaut and Le Loeuff (1998). During the next twenty years, several other specimens attributed to this bird were found in the same general region of France (Buffetaut and Angst, 2013, 2016a, 2016b. However, no specimen had hitherto been 0753-3969/© 2017 Elsevier Masson SAS. ...
Gargantuavis philoinos is a large terrestrial bird, initially described from several bones (synsacrum, cervical vertebra, femora) discovered in the Late Cretaceous deposits of southern France. The synsacrum described here comes from the late Campanian (Late Cretaceous) site of Laño, in north-western Spain, and is similar in all respects to the other synsacra of Gargantuavis philoinos described from France. This is therefore the first specimen attributed to this species to be reported outside France. This find increases the diversity of the fossil assemblage from Laño and the geographical distribution of this bird, which existed on the Ibero-Armorican island, which included southern France and the Iberian Peninsula, during the Late Cretaceous.
... This vertebra was identified as belonging to a giant bird that would be included within Ornithuromorpha and closely related to Ornithurae, due to its marked heterocoelous articulation. MPZ 2019/264 and MC-MN 478 are similar in size (cassowary-size according to Buffetaut and Angst, 2013) and in that the caudal articular surfaces of the centra project further than the postzygapophyses (Fig. 7C2). MC-MN 478 differs from MPZ 2019/264 in the presence of epipophyses above the postzygapophyses (A1, character 782:1; character 1083:0) (Fig. 7C2, C3) and in the scarce craniocaudal, and ample dorsoventral, development of the dorsal spine (Fig. 7A, C). ...
Throughout the evolutionary history of Avialae, several members of this clade have evolved into giant forms, in different time periods and ecological contexts. In Europe, the first birds that show this condition, the Gargantuaviidae, occur during the Late Cretaceous (late Campanian–early Maastrichtian), but it is during the Paleogene when more groups evolve large forms. However, until now, there was no record of any giant bird during the late Maastrichtian of Europe, close to the K/Pg boundary. Here we describe a cervical vertebra (MPZ 2019/264) from Beranuy (Huesca, NE Spain), which is the first fossil evidence of a giant bird from the late Maastrichtian of Europe, within Chron C29r. The vertebra displays some features, such as a well-marked heterocoelous articulation, lateral pneumatic foramina, ventral carotid processes, and a low neural spine, that support its inclusion within the clade Ornithuromorpha. This phylogenetic assignment is supported by two cladistic analyses. The vertebra is clearly different from the one assigned to Gargantuavis, meaning that it belonged to a distinct taxon. Although the kinship between these two taxa of giant birds is still unclear, this finding demonstrates that large-sized birds were part of the ecological communities of the Ibero-Armorican island from the late Campanian to the Late Maastrichtian, being present during the last hundreds of thousands of years prior to the K/Pg extinction event.
... The giant bird Gargantuavis philoinos is known from a few Late Cretaceous localities in France and Spain (Buffetaut & Angst 2016a, b;); in addition, a pelvis resembling that of Gargantuavis has recently been described from the Late Cretaceous of Romania (Mayr et al. in press). Its osteology is still incompletely known, as only elements of the pelvis [synsacrum and incomplete ilia Buffetaut et al. 1995;Buffetaut & Le Loeuff 1998;Buffetaut et al. 2015;Buffetaut & Angst 2016b), a cervical vertebra (Buffetaut & Angst 2013) and two femora (Buffetaut & Le Loeuff 1998;Buffetaut & Angst 2019)] had hitherto been reported. Therefore, any new discovery of Gargantuavis bones is worth reporting. ...
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An incomplete femur from the Bastide-Neuve site (Fox-Amphux, Var) is referred to the giant bird Gargantuavis philoinos, various remains of which have already been reported from that locality. It shows close similarities with a more complete Gargantuavis femur previously described from the Montplo-Nord locality (Cruzy, Hérault). An estimate based on the minimum circumference of the femur from Bastide-Neuve indicates a mass of 75 kg, between the weights of the living cassowary and ostrich.
... Northwest of Cruzy, at the site of Sainte-Foy, a very large form represented by one femur and several teeth very similar to Ischyrochampsa meridionalis has been studied (Buffetaut, 2001Buffetaut, , 2005 Martin & Buffetaut, 2005). South of Sainte-Foy, at the site of Montpl^ o Nord, some undetermined crocodylomorph remains have been found (Buffetaut & Angst, 2013). Another important site is Quarante, which is located southeast of Cruzy and from which two skull fragments (one dentary and the occipital region of one skull) assigned to Acynodon iberoccitanus have been studied (Martin, 2007). ...
A total of 227 theropod teeth have so far been recovered from the upper Campanian Laño site (northern Iberian Peninsula). The teeth were studied for their qualitative and quantitative features. From the theropod sample found at Laño, seven morphotypes attributed to five taxa are identified: a medium to large abelisaurid (Arcovenator sp.) and four small coelurosaurians (Dromaeosauridae indet., Paraves indet., cf. Paronychodon sp. and cf. Richardoestesia sp.) Together with the ground bird Gargantuavis and a possible ornithomimosaur, the theropod fauna of Laño might be composed of two medium–large-sized non-avian theropods, four small-bodied non-avian theropods and a large terrestrial bird. This makes the Laño site the richest and most diverse latest Cretaceous theropod site in Europe. Furthermore, the Laño site and the Upper Cretaceous localities of Europe that have yielded theropod remains suggest that the medium–large-sized theropods were abelisaurids or indeterminate theropods. The small theropods are more abundant, diverse and represented by different dromaeosaurids, Paronychodon, Richardoestesia or related forms, troodontids and, probably, by other paravians. Of the birds, enantiornithines, gargantuaviids and ornithurines are also common in the European Upper Cretaceous sites. The theropod assemblage of Laño, together with the taxa of other Upper Cretaceous sites, supports the idea that several theropod dispersal events took place during the Cretaceous. This resulted in a mixture of European endemic, Asiamerican and Gondwanan forms. This study also supports the hypothesis that the intra-Maastrichtian faunal turnover that occurred in the Ibero-Armorican landmass seems to have had no apparent effect on theropods.
We agree with Mayr et al., 2020, that a pelvis from the Maastrichtian of Romania closely resembles Garantuavis philoinos, from the Late Cretaceous of the Ibero-Armorican island, showing that gargantuaviids were not restricted to the latter landmass. However, we disagree with many other assertions and interpretations of Mayr et al.: contrary to their claims, lines of arrested growth are known in the bones of many modern birds; not all terrestrial birds have a narrow pelvis; the pubic and ischiadic peduncles are broken on all Gargantuavis pelves from the Ibero-Armorican island; an enlargement of the middle part of the synsacrum (to accomodate the glycogen body) is not present in all Ornithothoraces, since it is absent in Hesperornithiformes (probably because of the low number of preacetabular synsacral vertebrae, as in Gargantuavis); the palaeobiogeographical scenario proposed by Mayr et al. is not supported by palaeogeographical reconstructions of Cretaceous Europe. We conclude that Gargantuavis was a basal ornithurine, at an evolutionay level similar to that of Hesperornithiformes, in all likelihood the result of insular evolution (involving the crossing of sea barriers) on the Late Cretaceous European archipelago.
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A family of giant, flightless ground birds, the Gastornithidae, has been known for a long time from the early Tertiary (late Paleocene to middle Eocene) of both Europe and North America. The giant ground bird Gargantuavis was recently described from the Upper Cretaceous (probably early Maastrichtian) of France. The question may therefore be asked whether there is any close phylogenetic relationship between Gargantuavis and the Gastornithidae, which would suggest survival of giant birds across the Cretaceous-Tertiary boundary. A close anatomical comparison, however, reveals that Gargantuavis is a much more primitive bird than the Gastornithidae, and that they do not belong to the same lineage, resemblances probably being due to convergent adaptation to a similar mode of life. Although it cannot be demonstrated at the moment that Gargantuavis became extinct at the Cretaceous-Tertiary (K-T) boundary, this is a distinct possibility. If this is the case, the mass extinction of the K-T boundary will appear to have stopped early giantism and flightlessness in birds; these were followed by renewed and similar giantism and flightlessness in a different group of birds in the Paleocene.
Troodontidae is a clade of small, lightly built maniraptorans known from Cretaceous deposits of Asia and North America. These theropods have serrated teeth, raptorial hands, and an enlarged sickle-shaped claw on the foot. This chapter discusses the taxonomy and diagnostic features of troodontids. It also discusses phylogenetic hypotheses of troodontid relationships. Diagnostic remains of troodontids are largely restricted to Central Asia and China, and the origin and most of the evolutionary history of the clade was endemic to that continent. Only a single derived troodontid taxon, Troodon formosus, occurs in North America.
The continental tetrapod assemblages from the Santonian-Maastrichtian of Europe consist of dinosaurs (theropods: Abelisauroidea, Alvarezsauridae, Dromaeosauridae, ?Oviraptorosauria, ?Troodontidae, and birds: Enantiornithes, basal Ornithurae; sauropods: Titanosauria; ankylosaurs: Nodosauridae; ornithopods: Hadrosauridae, Rhabdodontidae; and neoceratopsians), pterosaurs (Azhdarchidae), crocodyliforms (eusuchians: Alligatoroidea, Gavialoidea, ?Hylaeochampsidae; sebecosuchian-like ziphosuchians; and, probably, basal neosuchians), choristoderes (?Champsosauridae), squamates (lacertilians: Iguanidae s.l., Paramacellodidae, Polyglyphanodontinae, Varanoidea; snakes: Madtsoiidae; possible amphisbaenians), turtles (cryptodires: Chelydroidea, Kallokibotionidae, Solemydidae; pleurodires: Bothremydidae, Dortokidae), lissamphibians (Albanerpetontidae; anurans: Discoglossidae, Palaeobatrachidae; caudates: Batrachosauroididae, Salamandridae), and mammals (multituberculates: Kogaionidae, ?"Paracimexomys group"; marsupials: Herpetotheriidae; eutherians: "Zhelestidae"). The palaeobiogeographical affinities of the Late Cretaceous continental tetrapods of Europe are complex. The faunas are commonly considered to show a mixed pattern resulting from the addition of "Asiamerican" and Gondwanan forms to European taxa. Albanerpetontids, both paramacellodid and polyglyphonodontine lizards, and hadrosaurid dinosaurs are taxa with Palaeolaurasian (or, in some case, even Neopangean) affinities. Other forms, such as paleobatrachid and batrachosauroidid lissamphibians, solemydid turtles, alligatoroid crocodyliforms, and nodosaurid dinosaurs can be considered as Euramerican taxa. Kallokibotionid and dortokid turtles, rhabdodontid dinosaurs and kogaionid mammals are endemic to Europe. The Gondwanan taxa have been regarded as African immigrants that reached southern Europe via the Mediterranean Tethyan sill. Abelisaurid and titanosaurid dinosaurs, trematochampsid crocodyliforms, podocnemidid and bothremydid turtles, and boid and madtsoiid snakes constitute the basic pattern of the so-called "Eurogondwanan fauna". However, the validity of some of these taxa is a disputed matter (Titanosauridae, Trematochampsidae), and the presence of other taxa in the Late Cretaceous of Europe is based on controversial data (Boidae, Podocnemididae). Only Abelisauroidea, Madtsoiidae and Bothremydidae (and, yet for confirming, Sebecosuchia) provide evidence of interchanges between Africa and Europe. At least abelisauroids might have reached Europe before the Late Cretaceous. In conclusion, most of the continental tetrapod taxa from the latest Cretaceous of Europe show affinities with those of northern Hemisphere landmasses. Latest Cretaceous trans-Tethyan dispersal events between Africa and Europe remain poorly documented.
Gargantuavis philoinos was described as a giant terrestrial bird on the basis of various postcranial elements (synsacrum and pelvis, femur) from Late Cretaceous (Campanian-Maastrichtian) localities in Southern France. It has recently been suggested that these remains in fact belong to giant pterosaurs. A detailed comparison between bones referred to Gargantuavis and the corresponding skeletal elements of pterosaurs reveals considerable differences and confirms the avian nature of Gargantuavis. The broad pelvis of Gargantuavis is similar to that of various extinct graviportal terrestrial birds.
Samrukia nessovi was described as a giant bird on the basis of a pair of mandibular rami from the Late Cretaceous of Kazakhstan. Anatomical comparison shows that the specimen bears no distinctive avian characters, and that its purported autapomorphies, as well as all its other characters, are in fact well-known pterosaurian features. The published phylogenetic analysis placing Samrukia within Aves is flawed because it did not include pterosaurs. Samrukia nessovi is clearly a large pterosaur, not a giant bird.