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Journal of the Geological Society A new giant ground from the Upper of southern France E. BUFFETAUT and J. LE LOEUFF Journal of the Geological Society 1998, v.155; p1-4. doi: 10.1 144/gsjgs.1 55.1 .0001

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Notes

© The Geological Society of London 2013 Downloaded from http://jgs.lyellcollection.org/ at University of Wisconsin - Madison on May 8, 2013 Jo urnal of the Geological Society, London, Vol. 155, 1998, pp. 1#4. Printed in Great Britain NsCtehiwnreeg t dllaeaicrsgces ooeyvnubes sriairo dcesrfsus m ap ofrufeotfv rhrideaorguna smlym eF nurr taec.nphcAo reb nteeto di tnnecf rrot ohmk menpolb t ewhateleseids Up gpeeo plfveo ia rfs cNonsisting of the synsacrum with part of the ilia is used to define a new t axon of bird, Gargantuavisp hiloinos, t o w hich a f emur is also referred. This bird is characterized by its v ery large size (comparable t o that of an ostrich), a broad pelvis w ith an anteriorly placed acetabulum, and a short robust f emur. The occurrence of this v ery large, flightless, bird in the Late Creta- ceous shows that, contrary t o w idespread opinion, the evolution of large terrestrial was not simply the result of adaptation during the Cenozoic t o ecological niches left empty by the extinction of the . It also strongly suggests that some of the large f ossil eggs from the Upper Cretaceous of southern France w ere laid by large ground birds rather t han dinosaurs.

Keywords: France, Upper Cretaceous, birds, eggs, dinosaurs.

The only bird specimen so far reported from the Upper Cretaceous of southern France was a synsacrum fragment from Provence (Buffetaut et al. 1995), w hich indicated a large form, but gave little more information about it. This paper describes new specimens found at two vertebrate sites in the Languedoc region of France, which provide additional information about the anatomy, size, and significance of these giant birds.

Systematic description Aves Gargantuavis p hiloinos gen. et sp. nov. Diagnosis: a v ery large bird with a broad pelvis, on which the acetabulum is placed in a v ery anterior position, at the level of the third and fourth synsacral transverse processes. The robust and relatively short synsacrum consists of ten completely fused vertebrae. The ilia do not meet each other dorsally. A well- developed antitrochanter is present posterodorsally to the relatively large acetabulum. Etymology: generic name from Gargantua, the giant of French folklore made famous by François Rabelais, and avis, 1

Latin for bird. Specific name from the Greek p hiloinos, #one who likes wine$, because the sites which have yielded remains of this bird are in the midst of v ineyards. Holotype: synsacrum with parts of the pelvis. Musée des Dinosaures, Espéraza, M D E-C3-525. Referreds pecimen: femur, Musée des Dinosaures, Espéraza, M DE-A08. This bone is referred to the same taxon as the pelvis because of agreement in size and compatible morphology (with, notably, a large trochanteric crest which probably matched a well developed antitrochanter on the ilium). Type locality: the holotype synsacrum was found at the Bellevue site at Campagne-sur-Aude (de´ partement A ude). The femur is from the Combebelle site (département Hérault). Both localities are in the Languedoc region of southern France. Type stratum: the Bellevue site is at the base of the continental Marnes de la Maurine Formation; it has yielded a rich v ertebrate fauna (Buffetaut et al. 1989; Le Loeuff 1995) dominated by titanosaurid sauropods and the ornithopod Rhabdodon. In southern France, this type of assemblage is characteristic of the late Campanian and early Maastrichtian, as opposed to late Maastrichtian assemblages in which hadrosaurs are dominant (Le Loeuff et al. 1994; Buffetaut et al. 1997). The Combebelle site has yielded a assemblage, with Rhabdodon and a titanosaurid, indicating a similar age.

Description. The partial pelvis from Bellevue consists of the synsacrum and the attached ilia, which are somewhat distorted and incomplete. All the constituent v ertebrae of the synsacrum are completely fused, as in other birds and much more so than in any described dinosaur. Because of t his fusion, the exact number of vertebrae in the synsacrum is not easy to determine. On the basis of the transverse processes which meet the ilia, it appears that ten vertebrae are present, a number similar to that observed in some Cretaceous birds such as and , and higher than that of any known theropod dinosaur (Chiappe 1996). The centra are 30 mm wide at the broadest (anterior) part of the synsacrum, which makes it as broad as that of an adult ostrich, but its preserved length (probably close to the original length) is only 180 mm. The ventral surface of the synsacrum is markedly arched longi- tudinally. A nteriorly, it bears a median ridge, w hich is replaced by a median groove at the level of the acetabulum. The concave anterior articular f ace of the first synsacral vertebra is subcircular, rather than saddle-shaped as in most birds. The opening of the neural canal is relatively small and circular in outline. Dorsally, the fused neurapophyses of the synsacral vertebrae form a continuous low ridge. The transverse pro- cesses connecting the synsacrum with the ilia are robust. In the anterior part, they are deep, perpendicular to the axis of the synsacrum and consist of a dorsal and a ventral bar, whereas the more posterior ones are simple, thinner and oblique. The incomplete and crushed ilia show few details. The most striking feature is the very anterior position of the acetabulum, which is at the level of the third and fourth transverse processes. The acetabulum is about 4 0 mm in diameter and ventrolaterally oriented. On the left side, a large antitrochanter is preserved, again an avian feature (Chiappe 1996). The dorsal iliac crest is preserved on the right side and it is clear that the ilia did not meet medially above the synsacrum. A small part of the anterior end of the ilio-ischiatic fenestra is visible on the left side. This pelvis is characterized mainly by its great relative

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Fig. 1. Synsacrum and partial ilia (holotype) of Gargantuavis p hiloinos, n.g., n.sp. (Musée des Dinosaures, Espéraza, M DE-C3-525), from the Upper Cretaceous of Bellevue (Campagne-sur A ude, France) in ventral (a), dorsal (b), anterior (c) and right lateral (d) views. Remaining matrix is shown by cross-hatching. a, acetabulum; at, antitrochanter; fn, fused neural spines of synsacral v ertebrae; ic, iliac crest. Scale bar is 50 mm. Drawings by Guy Le Roux. breadth (150 mm at the level of the acetabulum), by the anterior position of the acetabulum, and by the failure of the ilia to meet dorsally. It is different in its robustness from that of flying birds and is also v ery much unlike the relatively narrow pelves of modern ratites, and the very narrow ones of phorusrhacids. In this respect, as in the anterior position of the acetabulum, it is somewhat reminiscent of the Early Tertiary Diatryma (Matthew & Granger 1917; A ndors 1992), but there are differences in the dorsal extent of the ilia and in the shape of the anterior articular surface of the synsacrum. This broad and heavily built pelvis is interpreted as that of a large, probably not fast-running, flightless bird. The previously reported synsacrum fragment from Provence (Buffetaut et al. 1995) is similar to the middle part of the Bellevue specimen. The femur from Combebelle referred to Gargantuavis philoinos is not very well preserved, being crushed and damaged by modern bush roots. The distal end is missing, although the distal widening of the shaft suggests that not much is lacking, but the proximal end is tolerably well preserved. It is a short stout bone, with a hollow but thick- walled shaft (thickness of the bony walls up to 8 mm). The articular head is well defined, rounded, with a diameter of 40 mm (which fits that of the acetabulum of the pelvis from Bellevue). A well marked #neck$ separates the articular head from the more lateral region, in which there is a trochanteric crest similar to that of most birds, with no indication of the posterior trochanter present in dromaeosaurids, Archaeop- teryx and enantiornithines (Chiappe 1996). This crest is rounded in outline, instead of proximally pointed as in most large flightless birds. The great development of the crest matches that of the antitrochanter on the pelvis from Bellevue, thus supporting an attribution to the same taxon. In proximal view, the proximal articular surface is similar in outline to that of ratites and Diatryma. The minimum circumference of the shaft is 148 mm, in the size range of the living ostrich. Equations devised to estimate the weight of vertebrates on the basis of the circumference of the femoral shaft (Anderson et al. 1985) can be applied to the Combebelle specimen. The resulting estimated weight is 141 kg, which places it within the weight range of ostriches. This is further evidence of the flightless nature of this bird, the estimated weight for the largest known flying bird, Argentavis magnificens, being only 71.9 kg (Campbell & Marcus 1992).

Downloaded from http://jgs.lyellcollection.org/ at University of Wisconsin - Madison on May 8, 2013 U P P E R C R ET AC EO U S GIAN T G RO U N D BI R DS 3

Upper Cretaceous of Combebelle (Villespassans, France), Musée des Dinosaures M D E-A08, in anterior (a), posterior (b) and proximal (c) views. Scale bar is 50 mm. Photos by Claude Abrial.

Comparisons. The combination of characters seen on the pelvis of Gargantuavis p hiloinos separates it f rom all previously described birds and j ustifies the erection of a new taxon. The number of synsacral v ertebrae, although not v ery large by comparison with modern birds, which have 11 to 23 synsacral vertebrae, is slightly larger than in Patagopteryxa nd in known enantiornithines (Chiappe 1996). Among known Cretaceous birds, the great relative breadth of the pelvis is found only in Patagopteryx( Bonaparte & Alvarenga 1992; Chiappe 1996), and among later birds it is reminiscent mainly of robust extinct ground birds such as Diatryma (Matthew & Granger 1917; Andors 1992), but in the latter, as in most modern birds, the ilia meet dorsally above the neural spines of the syn- sacral v ertebrae, unlike the condition in Gargantuavis and Patagopteryx. The very anterior position of the acetabulum is remarkable; again, the condition in Gargantuavis is rather similar to that in Patagopteryx, and approximated, but not to the same extent, by Diatryma. The systematic position of Gargantuavis is debatable, although the number of synsacral vertebrae clearly places it in the unnamed group of Ornith- othoraces comprising the , Patagopteryx and the (Chiappe 1996). The absence of a posterior trochanter on the femur suggests that it is more derived than the enantiornithines, but the relatively large size of the acetabulum (Chiappe 1996) may suggest that it is not an ornithurine. As mentioned above, it resembles the smaller, chicken-sized Patagopteryx deferrariisi (Bonaparte & Alvarenga 1992; Chiappe 1996), from the Upper Cretaceous of , in several respects, but it is much larger and more robust, with a larger number of synsacral v ertebrae. Whether the resemblances indicate really close relationships between Gargantuavis and Patagopteryx, which is more derived than the enantiornithines but less so than the ornithurines (Chiappe 1995, 1996), is uncertain, but possible.

Conclusions: giant birds and dinosaurs. Gargantuavisphiloinos is not the first flightless bird to be reported from the Cretaceous. Both Patagopteryx (Bonaparte & A lvarenga 1992; Chiappe 1996) and (Marsh 1880) (and related marine birds), from the Upper Cretaceous of North America, were also flightless. However, the adaptations of Gargantuavis were different from those of the diving Hesperornithiformes, with their narrow pelvis, and it was much larger than either Hesperornis or Patagopteryx. Gargantuavis is so far the oldest known flightless bird to have reached a very large size (approximately that of an ostrich, although its proportions were probably different). A ccording to a widely held v iew (Romer 1966; K urt e´n 1971; Beaumont 1974; Benton 1990), large flightless birds (such as the Gastornithidae and the Phorusrhacidae) evolved in the early Tertiary to fill terrestrial ecological niches left v acant by the extinction of the dinosaurs at the end of the Cretaceous. The occurrence of Gargantuavis together with dinosaurs in the Late Cretaceous of Europe shows that the extinction of the dinosaurs was in fact not a prerequisite for the evolution of large ground birds. The occurrence of giant ground birds in western Europe during the Late Cretaceous may be linked to the absence there of ostrich-like ornithomimid dinosaurs, which were widespread during the same time interval both in A sia and in North America, and may have occupied a similar ecological niche. However, we need more giant bird material from Europe in order to assess the extent of convergent adaptation between those birds and ornithomimids. Whether there are any direct phylogenetic links between Gargantuavis and later large flightless birds seems unlikely in v iew of the above-mentioned peculiarities of the former, and the resemblances with Diatryma are probably explainable as convergences linked to similar adaptations in large and heavily built flightless birds. The discovery of Gargantuavis has a bearing on the interpretation of the large fossil eggs found in great abundance in the Upper Cretaceous of France. This large flightless bird must have laid large eggs on the ground. It is therefore likely that some of the above-mentioned eggs actually were laid by Gargantuavis rather than by dinosaurs, as is usually assumed#all the more so that eggshell fragments with a bird-like microstructure have been reported from the Upper Cretaceous of southern France (Cousin 1997). We therefore urge caution when discussing the chronology of dinosaur extinction in western Europe on the basis of eggshell remains alone.

We are grateful to the Association Culturelle et A rchéologique Cruziate for providing us with the femur from Combebelle, and to the many v olunteers who took part in the excavations at Bellevue, as well as to the owners of the site. We thank E. Gaffney (American Museum of Natural History, New Y ork), A. Milner (The Natural History Museum, London) and C. Lefèvre (Museum National d$Histoire Naturelle, Paris) for access to specimens in their care. A . Andors (New Y ork), L. Chiappe (New Yo rk) and L. Martin (Lawrence) offered valuable advice. This work was supported by the Institut National des Sciences de l$Univers (Paris) and the Musée des Dinosaures (Espéraza).

References

A#$%&+0#, J .F., H233-M2&45#, A. & R6++%33, D.A. 1985. Long-bone circum- ference and weight in mammals, birds and dinosaurs.J o urnal of Zoology, A207, 53761 . A#$0&+, A.V. 1992. Reappraisal of the Eocene groundbird Diatryma (Aves: Anserimorphae). Natural History Museum of Los Angeles County Science Series, 36, 10971 25.

B%2680#4, G. $% 1974. Guide des Vertébrés fossiles. Delachaux & Niestlé, Neuchätel.

Downloaded from http://jgs.lyellcollection.org/ at University of Wisconsin - Madison on May 8, 2013 4 E . B U F F ETA U T & J . L E LO E U F F

B%#40#, M.J . 1990. Vertebrate Palaeontology. Unwin Hyman, London. B0#292&4%, J .F. & A3<2&%#>2, H. 1992. A new flightless landbird from the Cretaceous of Patagonia. Natural History Museum of Los Angeles County Science Series, 36, 51764. B6??%4264, E., C3044%+, P., C6#@, G., D6A&0AB, S., L% L0%6??, J ., M2&45#, M ., P0C%33, J .E., R2@#26$, C. & T0#>, H. 1989. Les gisements de dinosaures maastrichtiens de la haute vallée de lDAude (France): premiers résultats des fouilles de 1989. Comptes Rendus de lDAcadémie des Sciences, Paris, 309, II, 1723.

EE, L% L0%6??, J ., C2<5#, L., D6??26$, S., GF%%&G&2#4, E., L26&%#4, Y. , M2&45#, M., R2>%, J .C., T0#>, H. & V 2++%, D. 1997. Late Cretaceous non-marine v ertebrates from southern France: a review of recent finds. Geobios Mémoire Spécial, 20, 9571 02.

EE, EE, M%AF5#, P. & M%AF5#-S23%++@, A. 1995. A large French Cretaceous bird. Nature, 377, 110. C289G%33, K. E. & M2&A6+, L. 1992. The relationship of hindlimb bone dimensions to body weight in birds. Natural History Museum of Los Angeles County Science Series, 36, 395741 2. CF5299%, L.M. 1995. The first 85 million years of avian evolution. Nature, 378, 3497355.

EE 1996. Late Cretaceous birds of southern South America: anatomy and systematics of Enantiornithes and Patagopteryx deferrariisi. Münchner Geowissenschaftliche A bhandlungen, A 30, 2037244. C06+5#, R. 1997. Nouvelle découverte de fragments de coquilles dDoeufs structure ornithoïde, dans les sédiments du Maastrichtien continental de Rennes-le-Château (Aude). Bulletin de la Sociétég éologique de Normandie et des amis du Muséum du Havre, 84, 1371 5. K6 &4 %´#, B. 1971. The Age of Mammals. Weidenfeld & Nicolson, London. L% L0%6??, J . 1995. Ampelosaurus atacis (nov.gen., nov.sp.), un nouveau Titanosauridae (Dinosauria, Sauropoda) du Crétacé supérieur de la Haute Vallée de lDAude (France). Comptes Rendus de lDAcadémie des Sciences, Paris, 321, IIa, 6937699. à ##, B$%%&+0$+, E. & M02+34, M. 1994. The last stages of dinosaur faunal history in Europe: a succession of Maastrichtian dinosaur assemblages from the Corbières (southern France). Geological Magazine, 131, 6255 630. M0267, O.C. 1880. Odontornithes: a monograph on the extinct toothed birds of North America. Government Printing Office, Washington. M0++7&8, W.D. & G2049&2, W. 1917. The skeleton of Diatryma, a gigantic bird f rom the Lower Eocene of Wyoming. Bulletin of theAmerican Museum of Natural History, 37, 3075326. R<>&2, A.S. 1966. Vertebrate Paleontology (3rd edition). University of Chicago Press, Chicago.

Received 11 J uly 1997; revised typescript accepted 18 A ugust 1997. Scientific editing by Nick Rogers.