Describer

Le Loeuff, 1995

Time

Cretaceous Late Campanian Maastrichtian

Classification

Saurischia Sauropodomorpha Sauropoda Titanosauria

Diet

Herbivore

Fossilsite

Marnes Rouges Inférieures Formation, Bellevue bonebed (Layer C3), Blanquette de Limoux vineyards, Campagne-sur-Aude, Aude Valley, Esperaza, France

Length

15 meter

Info

Genus - Typespecies

The first plated sauropod that has been found on the northern hemisphere was found in a former river bed.. It belongs to the midsized species of plant-eating sauropods. Its name stands for \\\" dinosaur of the vineyard.

Ampelosaurus atacis (nov. gen., nov. sp), a new titanosaurid (Dinosauria, Sauropoda) from the Late Cretaceous of the Upper Aude Valley (France)*

Jean Le Loeuff Translated by Matthew C. Lamanna July-August 2001

Introduction

Excavations led since 1989 at Campagne-sur-Aude (Aude Department, France, fig. 1) by a team from the Musée des Dinosaures in Esperaza (Jean Le Loeuff, Valérie Martin), the CNRS (Eric Buffetaut, Haiyan Tong, Lionel Cavin), and the Muséum d’Histoire Naturelle in Boulogne-sur-Mer (Michel Martin) permitted, in twelve months of fieldwork (with the involvement of several score of volunteers), the discovery of several hundred Maastrichtian reptile bones.

The Bellevue layer (layer C3 of Buffetaut et al., 1989) is situated at the base of the member of the Marnes Rouges of the Maurine of the lower Marnes Rouges Formation, of Maastrichtian age (Bilotte, 1985).

This continental formation is formed of fluviatile deposits (channels and floodplains) and has produced many dinosaur bones (Clottes and Raynaud, 1983; Buffetaut et al., 1989; Le Loeuff, 1991; 1992). The Bellevue layer has yielded in particular very numerous remains of a sauropod dinosaur of the family [Titanosauridae-Titanosauria], whose diagnosis is proposed here. Other vertebrates have been discovered: the fish Lepisosteus is represented by some scales, an indeterminate turtle by fragments of shell, a eusuchian crocodilian by some teeth and cranial fragments, a small theropod dinosaur by some isolated teeth, and the [Euornithopoda] incertae sedis Rhabdodon priscus (Matheron, 1869) by a dentary and many postcranial elements. Remains of plants and invertebrates have also been discovered.

Systematics

Sauropoda Marsh, 1878
Titanosauridae Lydekker, 1893
Ampelosaurus, nov. gen.


Derivatio nominis: from the Greek ampelous (fem) (the vineyard; cf. Elien, De la Nature des Animaux, 11, 32) and from the Latin saurus (reptile); the layer is situated in the southern border of the Blanquette de Limoux vineyard.


Species typicus: Ampelosaurus atacis, nov. sp.


Diagnosis: the same as that of the type species.


Ampelosaurus atacis nov. sp.

* Original citation: Le Loeuff, J. 1995. Ampelosaurus atacis (nov. gen., nov. sp.), un nouveau Titanosauridae (Dinosauria, Sauropoda) du Crétacé supérieur de la Haute Vallée de l’Aude (France).
Comptes Rendus de l’Academie des Sciences Paris 321 (ser. IIa): 693-699.


Derivatio nominis: from the Latin Atax, the Aude river.

Holotypus: three dorsal vertebrae in connection (C3-247).

Referred material: a tooth (C3-52); dorsal vertebrae (C3-38, C3-59; C3-92; C3- 93; C3-94; C3-148); caudal vertebrae (C3-24; C3-25; C3-26; C3-27; C3-46; C3-55; C3- 58; C3-63; C3-64; C3-65; C3-95; C3-96; C3-97; C3-98; C3-99; C3-100; C3-101; C3- 124; C3-127; C3-147); dorsal ribs (C3-60); chevrons (C3-72; C3-139); sternal plates (C3- XX; C3-80); scapulae (C3-21; C3-145); coracoids (C3-22; C3-161; C3-351); humeri (C3- 1; C3-79; C3-81; C3-86; C3-175; C3-312); ulnae (C3-56; C3-83; C3-300); radius (C3- 85); pubis (C3-57); ilium (C3-123); ischium (C3-84); femora (C3-20; C3-40; C3-44; C3- 61; C3-78; C3-201; C3-203; C3-261); tibiae (C3-137; C3-138; C3-144; C3-173); fibulae (C3-48; C3-137); phalanges (C3-88); dermal plates (C3-136; C3-192; C3-204; C3-205). All material is in of the collections of the Musée de Dinosaurs of Esperaza (Aude).

Locus typicus: Bellevue (Layer C3 of Campagne-sur-Aude; cf. Clottes and Raynaud , 1983; Buffetaut et al., 1989).

Stratum typicum: Marnes Rouges Inférieures Formation, member of the red marls of the Maurine (Bilotte, 1985).

Age: lower Maastrichtian (Bessière et al., 1989).

Diagnosis: teeth weakly spatulate; distal extremity of the neural spine of the dorsal vertebrae widened laterally; neural arch of the dorsal vertebrae strongly inclined toward the rear; absence of the distal expansion of the scapular blade; light ventral crest of the scapula; presence of osteoderms (plates, bulbs, spines).

Preliminary description

With the exception of the skull (represented by only one tooth), a great part of the skeleton of A. atacis is known from disarticulated bones from the Bellevue bonebed. Skull Only one titanosaur tooth, having a very particular morphology, has been recovered at Bellevue (fig. 2). The specimen (C3-52) measures 21 mm in height by a maximum width of 6 mm toward the middle of the crown. The axial part of the tooth is more or less cylindrical, with very thin rostral and caudal expansions, that give it a morphology similiar to a willow leaf. These expansions stop toward the base of the crown, causing a slight constriction at this level. This tooth is of the spatulate type, in opposition to the cylindrical teeth of the Laño titanosaur (Sanz, 1986; Astibia et al., 1990), more classic in this sauropod family. The surface of enamel is slightly distressed, as in the Laño titanosaur.

Axial skeleton

Three middle dorsal vertebrae in connection (C3-247) constitute the holotype of Ampelosaurus atacis. The centra, distinctly opisthocoelous, lengthen toward the rear. The pleurocoels are well defined, with an angular dorsal border. The neural spines are inclined strongly toward the rear. On the most posterior dorsal vertebra, a supplementary lamina is developed under the posterior infradiapophyseal lamina, and terminates at the level of departure of the infrapostzygapophyseal lamina.

The internal structure of the dorsal vertebrae is spongy with very large cells. Two dorsal vertebrae are preserved with a complete neural arch, but very crushed (C3-59). The neural spine, better conserved than that of the holotype, has a very characteristic shape: it is very widened distally, narrows downwards, and enlarges again at the level of the postzygapophyses (fig. 4). It does not possess hyposphene-hypantrum structures.

The caudal vertebrae are laterally compressed; they are all strongly procoelous. The anterior caudals possess very short prezygapophyses, with large prezygapophyseal facets. The diapophyseal rolls are very prominent, with tips directed toward the rear. The neural spine, very narrow and very high, is directed toward the rear; on the most anterior vertebrae, this spine possesses pre- and postspinal laminae. The middle caudal vertebrae are more laterally compressed; the centrum is proportionally more lengthened. The prezygapophyses are also more and more long, with facets less and less developed; correlatively, the postzygapophyses are situated more anteriorly on the centrum.

Appendicular skeleton

A right scapula has been discovered at Bellevue (C3-21), associated with a coracoid (C3-22). The craniocaudal length of the bone is of 72 cm. Contrary to other titanosaurs, the scapular blade of A. atacis is of roughly triangular shape, larger toward its base. Indeed, the scapular blade narrows distally and does not present a distal expansion, in contrast to other titanosaurs . The angle between the dorsal (or anterior) border of the scapular blade and the proximal expansion of the scapula is very obtuse, as in the Laño titanosaur. This angle is close to 900 in most titanosaurids.

The scapula presents a light ventral crest, behind the glenoid fossa; this crest is similar to that preserved on a scapulocoracoid from Mas d’Azil (Caoué layer: Collection Pouech (n0 Ca-1), Collège Jean XXIII, Pamiers). It is much less developed and more proximal than the posterior ventral scapular crest of the Laño titanosaur and that of \\\"Titanosaurus indicus\\\" of Fox- Amphoux (Lapparent, 1947).

Contrary to Magyarosaurus dacus and Saltasaurus loricatus, the scapula does not present the dorsal crest at the base of the scapular blade. The coracoid is of quadrangular shape, the coracoid foramen being situated in a very dorsal position compared to the glenoid fossa. A pubis (C3-57), 75 cm long, presents a large foramen, and a strong distal expansion, that recalls a titanosaurid from Brazil (cf. McIntosh, 1990, p. 371). The ilium (C3-123) does not present craniolateral development of the preacetabluar blade, contrary to the South American titanosaurids (Powell, 1992). Many femora are preserved; their length varies from 75 to 115 cm. The proximolateral deflection is well marked.

Osteoderms

Four osteoderms measuring between 25 and 28 cm long are referred to Ampelosaurus; they present different morphologies (spines, plates, bulbs). As in many titanosaurids, A. atacis was therefore covered in real bony armor (cf. Le Loeuff et al., in press, for a description of these structures and their paleobiologic significance).

Discussion

Ampelosaurus atacis is one of the best represented European sauropods; this form measured up to 15 m long. A. atacis is evidently a titanosaur, with [word missing] strongly procoelous caudal vertebrae, an absence of hyposphene-hypantrum structures in the dorsal vertebrae, and the bony texture of the presacral vertebrae composed of very large cells (Powell, 1986; 1987; 1992; McIntosh, 1990; Le Loeuff, 1993).

The titanosaur of the Haute Vallée de l’Aude is distinctly different from the other forms of the Upper Cretaceous of Europe. It is distinguished at this time from Magyarosaurus dacus (in this form: pleurocoels of the dorsal vertebrae very reduced; caudal vertebrae compressed dorsoventrally, dorsomedial crest of the scapula more developed, very reduced proximolateral expansion of the femur) and from the Laño titanosaur: (cylindrical teeth, caudal vertebrae compressed dorsoventrally, posterior ventral crest of the scapula).

Conclusion

The preliminary description of Ampelosaurus atacis (a mongraph is in preparation) confirms the diversity of the titanosaurs of the European Maastrichtian. Three forms are currently correctly defined (A. atacis and the Laño titanosaur in the lower Maastrichtian, and Magyarosaurus dacus in the upper Maastrichtian).

Hypselosaurus priscus (Matheron, 1869), from the Maastrichtian of Provence, based on non-diagnostic remains, is a nomen dubium (Le Loeuff, 1993). Many other isolated remains discovered in the lower Maastrichtian of the south of France and the north of Spain cannot be referred to these forms (Le Loeuff, 1993): the diversity of the titanosaurs in the beginning of the Maastrichtian was therefore certainly more important than is currently known. Their decline in the upper Maastrichtian, parallel to the development of the Hadrosauridae (Le Loeuff et al., 1993; 1994) is probably due to a climatic deterioration in the last millions of years of the Cretaceous, with the passage of a subtropical climate to a more temperate climate in the south of Europe (a region situated then toward 350 north latitude: cf. Bardossy and Dercourt, 1990). Although no French site of the upper Maastrichtian has produced remains of [Titanosauridae-Titanosauria] to this day, their survival until the extreme end of the Maastrichtian is nevertheless attested to in Transylvania, with Magyarosaurus dacus, and in Catalonia (Casanovas et al., 1987).

Acknowledgments: Jean-Luc Le Douarec, Colette Rives, and Gérard Chauvet (GAEC of Bellevue) have kindly allowed us to work on the property. The excavations have been driven thanks to the involvement of about a hundred volunteers of the l’Association Dinosauria since 1989. The logistical support of SIVOM of the Haute Vallée de l’Aude was indispensable. The preparation of the material has been assured since 1992 by Valerie Delforno (Musée des Dinosaures, Esperaza). Numerous discussions with Eric Buffetaut (Paris), Valerie Martin (Esperaza), Jaime Powell (Tucuman), and José-Luis Sanz (Madrid) improved this article considerably.

_{Source: Polyglot Paleontologist}

---

Le Loeuff, J. (2005) Osteology of Ampelosaurus atacis (Titanosauria) from Southern France. in: Thunder Lizards - The Sauropodomorph Dinosaurs. Eds.: Tidwell, V. & Carpenter, K. Part 1 Ch. 4 pp. 115-137

Abstract: More than 500 titanosaur bones referred to Ampelosaurus atacis (Le Loeuff, 1995) have been excavated since 1989 at the Upper Cretaceous lacality of Bellevue in the Upper Aude Valley (Aude department, southern France). The skeleton of A. atacis is described and compared to other Europoean titanosaurids. Evidence is accumulating that Late Cretaceous sauropods in Europe were more diverse than previously realized.

Amended diagnosis: Axial part of the teeth cylindrical, with thin rostral and caudal expansions; constriction between the crown and the root; presence of an accessory spinal limina in posterior cervicals; posterior centroparapophysial lamina very developed in dorsal vertebrae; well-developed accessory lamina between the postzygapophysis and the posterior centradiapophyseal lamina in dorsal vertebrae\\\'dorsal neural spines directed backward; two sacro-caudals in the sacrum; first three caudals procoelus with very eleongated prezygapophyses; scapular blade unexpanded distally; shaft of the ischium enexpanded; and presence of osteoderms including spines.

Description: Skull

Cranium and teeth: A partial cranium consisting of frontals, parietals, supraoccipital, exoccipitals, basisocipital, prootics, laterosphenoids, orbitosphenoids, and the dorsal part of the parasphenoid-basisphenoid complex (C3-761). Almost no sutures can be distinguished. most of teh bones being completely fused. The most conspicious features of this cranium are the narrow supratemporal fenestra, the thick parietals forming a marked promontory similar to that of Antarctosaurus wichmannianus (cf. Huene, 1929), and a strong nuchal crest. A partial dentary (MDE-C3-396) preserves nine alveoli.

The tooth (C3-52) is 21 mm. high and has a maximal width of 6 mm. The axial part of the crown is roughly cylindrical, with thin rostral and caudal expansions that stop at te base of the crown. Libially, there is an apical wear effect. This slightly spatulate tooth is quitte different from the more usual peglike teeth of [Titanosauridae-Titanosauria]. It is very different from the cylindrical titanosaurid teeth recovered in Romania and Spain, and reffered to the genera Magyarosaurus dacus and Lirainosaurus astibiae respectively.

The general morphology of the cranial elements of A. atacis (position of the foramina for cranial nreves, thickened parietals) is very similar to that of other titanosaurids such as Antarctosaurus wichmannianus and the braincase from Dongargaon (See Berman and Jian, 1982) and (Wilson et al., 2005 [GSP-UM 7000, an isolated braincase that includes elements of the skull roof (frontal, parietal) and basicranium (supraoccipital, exoccipital-opisthotic, basioccipital, basisphenoid, prootic, laterosphenoid, orbitosphenoid). A cast of the specimen is housed in the University of Michigan Museum of Paleontology (UMMP 11303)]). Another partial titanosaurid skull was described from southern France by Le Loeuff at al., (1989) and shows some differences with Ampelosaurs (including the precense of facets arround the occipital condyle), suggesting the presence of at least one other titanosaur species in the Upper Cretaceous of Southern France. (See Le Loeuff, 1998).

Klein, N., Sander, M. & LeLoeuff, J. (2006) An unusual bone histology and growth pattern in Ampelosaurus atacis, a Titanosaurid from South France. JVP 26(3) Abstracts pp.85

Several humeri and femora of the titanosaur sauropod Ampelosaurus atacis from the Maastrichian of South France were sampled by core drilling and cross sectioning for paleohistological study. The material was collected within the scope of a large project about sauropod bone histology. The comparison with other sampled sauropods (Neosauropoda including several Diplodocoidea and Macronaria) or any other dinosaur currently studied by bone histology, shows that Ampelosaurus atacis is clearly different/unique in its bone histology.

Whereas all studied sauropods and even closer relatives like Alamosaurus and Phuwiangosaurus show the laminar fibro-lamellar bone typical for most dinosaurs, Ampelosaurus atacis indeed retains the laminar vascular organization, but completely lacks the fibrous bone tissue component, even in early ontogenetic stages. The dominant bone tissue here is the parallel-fibered bone and lamellar bone. However, contrary to e.g. reptiles which also grew with parallel-fibered or lamellar bone, A. atacis growth was not cyclically interrupted by LAGs or other growth marks, except in the outer cortex of some ontogenetically old specimens. Furthermore, A. atacis long bones are characterized by remodelling (secondary osteons) starting very early in ontogeny.

Current phylogenetic hypothesis place Ampelosaurus as a derived titanosaur. Thus, its unique bone histology must have evolved from the standard sauropod long bone histology that always indicates high growth rates. The lack of the fibrous component in the laminar bone of A. atacis suggests a greatly reduced bone apposition rate, which in turn would implicate an clearly decreased growth rate and delayed maturity relative to other sauropod dinosaurs. Such a delayed maturity is sometimes seen in insular forms such as the extinct moa (Dinornis) of New Zealand, although the bone histology of moa is different from Ampelosaurus. This is consistent with the Maastrichtian paleogeography of southwestern Europe, which indicates that A. atacis inhabited large islands.

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