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All elements of the skull of FMNH PR 2100 are represented on at least one side, and most are complete and undistorted. The only missing elements are the left premaxilla, right pterygoid, right ectopterygoid, and right vomer. In Fig. 1, the left premaxilla has been reconstructed on the basis of the preserved right premaxilla.
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The original theropod materials recovered from the Maevarano Formation, including isolated teeth and caudal vertebrae, were assigned to Megalosaurus crenatissimus (5). However, no single holotype specimen was named from this collection, and none of the specimens appear to be diagnostic to the generic level. A partial dentary was subsequently made the holotype of the species and was placed into a new genus, Majungasaurus (6). However, comparison of the latter specimen with dentaries of other abelisaurids failed to reveal diagnostic characters. The jaw is fragmentary, and the dentition does not appear to differ significantly from that in the Indian abelisaurid materials.
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The holotype specimen of Majungatholus atopus (MNHN.MAJ 4), housed in the Muséum National d'Histoire Naturelle, Paris, consists of a partial skull roof with a frontal "dome" (7). Several characteristics of this specimen (for example, lack of radiating arrangement of bony trabeculae in dome; dome occurring wholly within frontals rather than incorporating parietals; rugose ornamentation) are not present in any members of Pachycephalosauria. Conversely, MNHN.MAJ 4 has several theropod characteristics (for example, long, divided olfactory tracts) as well as derived features present in FMNH PR 2100. Most importantly, the frontal "dome" of the holotype is equivalent to the frontal "horn" in FMNH PR 2100.
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24
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2642605959
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Diagnostic characters for Majungatholus atopus: thickened, fused, highly pneumatic nasals with large, bilateral foramina; frontals with median hornlike projection; cervical ribs bifurcate distally; cervical ribs with multiple enlarged pneumatic foraminae proximally (diameter > 10 mm).
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J. A. Gauthier, Mem. Calif. Acad. Sci. 8, 1 (1986); T. Holtz, J. Paleontol. 68, 1100 (1994); P. Sereno, Neues Jahrb. Geol. Palaeontol. Abh., in press. Ceratosauria includes at least two groups, Coelophysoidea and Ceratosauroidea. Coelophysoids (for example, Coelophysis, Syntarsus, Dilophosaurus) are known from the Late Triassic and Early Jurassic of Africa, Europe, and North America. Ceratosauroids are thus far represented by two genera (Ceratosaurus and Elaphrosaurus), from the Late Jurassic of North America and Africa, and by the Cretaceous abelisaurids, known from South America, Madagascar, and the Indian subcontinent. Tetanurae includes more derived theropods such as Allosauroidea (allosaurids, spinosaurids, carcharodontosaurids) and Coelurosauria (for example, dromaeosaurids, ornithomimids, troodontids, and tyrannosaurids, as well as birds), which together are represented on all major landmasses.
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2642698442
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Synapomorphies of Abelisauridae observable in Majungatholus atopus include Cranial: External sculpturing of craniofacial elements; premaxilla with reduced to absent palatal process; maxilla-jugal contact elongate; rostral process of lacrimal strongly reduced or absent; lacrimal with pronounced suborbital process; broad lacrimal-postorbital contact; long axis of postorbital oriented rostroventral-caudodorsal; postorbital with pronounced suborbital process; high, transversely broad nuchal crest with large squamosal contributions; medial alveolar border of premaxilla, maxilla, and dentary bearing fused interdental plates with series of vertical ridges and grooves; hypertrophied external mandibular fenestra and associated weak contacts between dentary and postdentary elements. Postcranial: Postaxial epipophyses elongate.
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2642658980
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Synapomorphies of Majungatholus and Carnotaurus include Cranial: Postorbital rostrocaudally elongate; postorbital with stepped-down ventrolateral fossa; dorsally placed median fossa in saddle-shaped depression overlapping contact of frontals and parietals; enlarged, caudodorsally projecting parietal eminence supported ventrally by stout supraoccipital; laterally directed paroccipital processes with up-turned distal ends; stout, triangular splenial with straight caudal margin (73).
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The putative abelisaurid materials from Europe include a dentary and several isolated postcranial elements from the Late Cretaceous of France (24). Several of these appear to be derived from ceratosaurian theropods yet lack diagnostic abelisaurid features. The Moroccan materials (25) include isolated, partial jaw elements thought to resemble those of abelisaurids, yet once again there are no definitive characters supporting this designation.
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The described materials of Indian abelisaurids appear to represent at least two taxa, Indosaurus and lndosuchus, from the Late Maastrichtian Lameta deposits (27, 28). Both taxa were erected on fragmentary brain cases. Although the collection includes numerous additional cranial and postcranial elements, the great majority were surface collected as isolated specimens. Thus, assigning particular elements to one or the other taxon is problematic until associations are established, particularly between cranial and postcranial elements.
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note
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We thank B. Rakotosamimanana, P. Wright, B. Andriamihaja, the Institute for the Conservation of Tropical Environments staff, the people of Berivotra, and all expedition members for assistance with field work; G. Buckley, J. Flynn, and W. Simpson (FMNH) for preparation support; P. Currie and G. Olshevsky for anatomical and nomenclatural discussions; R. Coria and D. Fastovsky for manuscript reviews; K. Passaglia, V. Heisey, and J. Holstein for preparation of specimens; M. Stewart and J. Weinstein for photography; and B. Chen and R. Day for assistance with CT scanning. The figures were composed by L. Betti-Nash. Access to comparative materials was kindly provided by P. Taquet (Museum National d'Histoire Naturelle, Paris); S. Cumbaa, K. Shepherd, and C. Kennedy (Canadian Museum of Nature, Ottawa); S. Chatterjee (Texas Tech University, Lubbock); D. Rudra and S. Bandyopadhyay (Indian Statistical Institute, Calcutta); and S. Sen Gupta (Geological Survey of India, Calcutta). Supported by NSF and The Dinosaur Society.
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