New evidence of a giant bird from the Late Cretaceous of France

Abstract

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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Cambridge University Press 2012 1
doi:10.1017/S001675681200043X
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New evidence of a giant bird from the Late Cretaceous of France
ERIC BUFFETAUT∗†& DELPHINE ANGST‡
∗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)
Abstract
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,
vertebra
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: eric.buffetaut@sfr.fr
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

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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

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RAPID COMMUNICATION 3
(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
assemblages.
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,

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4RAPID COMMUNICATION
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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