A brief history of Mesozoic theropods research in Gondwana

Snout of the ceratosaurian Genyodectes serus

In the last decades, the study of Gondwanan non-avian theropods has been highly prolific, showing that the group reached a great taxonomic and morphological diversity comparable to that of Laurasia. The Mesozoic Gondwanan neotheropod record includes: coelophysoids, basal averostrans, ceratosaurids, abelisauroids, megalosauroids, carcharodontosaurids, megaraptorans, basal coelurosaurs, compsognathids, alvarezsauroids, unenlagiids, and basal avialans, as well as putative tyrannosauroids, ornithomimosaur-like forms, and troodontid. Therefore, the Gondwanan fossil record has been crucial to understand the evolution and global biogeography of dinosaurs during the Mesozoic.

The first probable theropod remains from Gondwana were discovered in Colombia by Carl Degenhard, a German engineer, in 1839. At that time the word “dinosaur” did not even exist yet. Although Degenhard identified them as bird footprints, his brief description suggests that they were tracks of bipedal dinosaurs. But it was not until 1896 that the first Gondwanan theropod was named by the French palaeontologist Charles Depéret as “Megalosaurus” crenatissimus from the Upper Cretaceous of Madagascar. Several theropod remains were described from India, Africa, and South America during the 19th century. These early fragmentary discoveries lead the authors of the late XIX and early XX centuries to interpret them as belonging the same lineages present in Europe and North America.

Elaphrosaurus bambergi (Museum für Naturkunde 4960, holotype) from the Upper Jurassic of Tanzania (Janensch, 1920)

In 1901, A. Smith Woodward described Genyodectes, based on fragmentary skull bones, including portions of both maxillas, premaxillae,  parts of the supradentaries, and some teeth, discovered by Santiago Roth in Chubut, at the end of the 1880s. Genyodectes remained as the most completely known  theropod from South American until the 1970s. In 2004, O. Rauhut concluded that Genyodectes is more closely related to Ceratosaurus than the more derived abelisaurs.

Between 1915 and 1933, the most relevant Gondwanan theropod discoveries were produced by the work of the German palaeontologists Frederich von Huene, Ernst Stromer, and Werner Janensch, including for the first time the publication of very informative partial skeletons, such as those of Spinosaurus aegyptiacus and Elaphrosaurus bambergi (Stromer, 1915; Janensch, 1920). Despite its low fossil record, Spinosaurus is one of the most famous dinosaur of all time. This gigantic theropod possessed highly derived cranial and vertebral features sufficiently distinct for it to be designated as the nominal genus of the clade Spinosauridae. But during and after the Second World War the influence of the German palaeontology in the research of Gondwanan theropods abruptly declined.

Skull and neck of Carnotaurus sastrei

By the 1960s, the Argentine biologist Osvaldo Reig, together with Rodolfo Casamiquela and José Bonaparte, began to explore the Mesozoic rocks of Argentina looking for fossil tetrapods. In 1985, Bonaparte published a note presenting Carnotaurus sastrei as a new genus and species and briefly describing the skull and lower jaw. It was collected in the lower section of La Colonia Formation, Chubut Province. The discoveries of Bonaparte and his collaborators resulted in the recognition of the Patagonian theropod record as the most relevant and informative among Gondwanan continents. Some of the theropod species discovered in Patagonia are known on the basis of skulls and fairly complete skeletons offering insights into the anatomy and phylogeny of abelisaurids, carcharodontosaurids, and maniraptorans.

References:

Martín D. Ezcurra, and Federico L. Agnolín (2017). Gondwanan perspectives: Theropod dinosaurs from western Gondwana. A brief historical overview on the research of Mesozoic theropods in Gondwana. Ameghiniana 54: 483–487. Published By: Asociación Paleontológica Argentina https://doi.org/10.5710/102.054.0501

Novas, F.E., et al., Evolution of the carnivorous dinosaurs during the Cretaceous: The evidence from Patagonia, Cretaceous Research (2013), http://dx.doi.org/10.1016/j.cretres.2013.04.001 

Buffetaut, E. 2000A forgotten episode in the history of dinosaur ichnology; Carl Degenhardt’s report on the first discovery of fossil footprints in South America (Colombia, 1839). Bulletin de la Société Géologique de France 171: 137140Google Scholar

 

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A brief introduction to the Carnotaurus family tree.

 

Skull and neck of Carnotaurus sastrei

Skull and neck of Carnotaurus sastrei (From Novas et al., 2013)

The Abelisauridae represents the best-known carnivorous dinosaur group from Gondwana. Their fossil remains have been recovered in Argentina, Brazil, Morocco, Niger, Libya, Madagascar, India, and France. The oldest records of abelisauroid theropods are from the Early Jurassic. These ceratosaurian theropods exhibit spectacular cranial ornamentation in the form of horns and spikes; and strongly reduced forelimbs and hands. The group was erected by Jose Bonaparte with the description of  Abelisaurus Comahuensis. Although represented by relatively well-known skeletons, the phylogenetic relationships within abelisaurids remain debated. The Argentinean record of abelisauroid theropods begins in the Middle Jurassic (Eoabelisaurus mefi) and spans most of the Late Cretaceous, from Cenomanian (Ilokelesia, Xenotarsosaurus, and Ekrixinatosaurus) to Campanian–Maastrichtian (Abelisaurus, Carnotaurus, Aucasaurus, and Noasaurus).

Abelisauroidea has been divided into two main branches: the Noasauridae and the Abelisauridae. The Noasauridae are known from Cretaceous beds in northern Argentina, Madagascar, India, and Niger. They are small and slender with sizes that range from 1 to 3 metres in length. The best-preserved and most complete noasaurid is Masiakasaurus knopfleri from the Maastrichtian of Madagascar. The Abelisaurids are medium to large, robust animals, such as the Carnotaurus and the Majungasaurus of Madagascar. The group exhibits short, round snouts; thickened teeth; short, stocky arms; and highly reduced forearms.

Masiakasaurus on display at the Royal Ontario Museum.

Masiakasaurus on display at the Royal Ontario Museum.

Carnotaurus sastrei is the most advanced member of Abelisauridae. It was collected in the lower section of La Colonia Formation, Chubut Province, Argentina, by an expedition led by Argentinian paleontologist José Bonaparte. In 1985, Bonaparte published a note presenting Carnotaurus sastrei as a new genus and species and briefly describing the skull and lower jaw. The skull of Carnotaurus is complete, measuring 60 cm from the tip of the premaxillae to the distal tip of the paroccipital process. The most distinctive features of Carnotaurus are the two robust conical horns that extend from the frontals. The horns are dorsoventrally compressed, and 146 mm long on both sides. The dorsal surface of each horn is ornamented with a series of longitudinal grooves. Because relatively few abelisaurid braincases are known, the description of the Carnotaurus braincase is important for understanding the variability of this structure within the clade (Carabajal 2011). C. sastrei would have had a comparatively weak muscle-driven bite.

The forelimbs of Carnotaurus show an extreme reduction, proportionally greater than the reduction observed in tyrannosaurids, although the radius, ulna and humerus are very robust. The hand has four digits, including a large, conical-shaped metacarpal IV lacking an articulation for a phalanx.

 

References:

Novas, F.E., et al., Evolution of the carnivorous dinosaurs during the Cretaceous: The evidence from Patagonia, Cretaceous Research (2013), http://dx.doi.org/10.1016/j.cretres.2013.04.001

Bonaparte, José F.; Novas, Fernando E.; Coria, Rodolfo A. (1990). “Carnotaurus sastrei Bonaparte, the horned, lightly built carnosaur from the Middle Cretaceous of Patagonia”, Contributions in Science (Natural History Museum of Los Angeles County) 416.

Mazzetta, Gerardo V.; Fariña, Richard A.; Vizcaíno, Sergio F. (1998). “On the palaeobiology of the South American horned theropod Carnotaurus sastrei Bonaparte”, Gaia 15: 185–192.

Ruiz, Javier; Torices, Angélica; Serrano, Humberto; López, Valle (2011). “The hand structure of Carnotaurus sastrei (Theropoda, Abelisauridae): implications for hand diversity and evolution in abelisaurids”. Palaeontology 54 (6): 1271–1277.

Introducing Gualicho.

Gualicho shinyae, at the Centro Cultural de la Ciencia.

Gualicho shinyae, at the Centro Cultural de la Ciencia.

The Cretaceous beds of Patagonia have yielded the most comprehensive record of Cretaceous theropods from Gondwana and includes at least five main theropod lineages: Abelisauroidea, Carcharodontosauridae, Megaraptora, Alvarezsauridae, and Unenlagiidae. The best represented theropod clades in the Late Cretaceous terrestrial strata of the Neuquén Basin are the Abelisauroidea and the Carcharodontosauridae. The  Abelisauroidea has been divided in two main branches: the Noasauridae which includes the small-sized abelisauroids, and the Abelisauridae which comprises medium to large-sized animals, like the popular Carnotaurus sastrei. The group exhibits strongly reduced forelimbs and hands, stout hindlimbs, with a proportionally robust and short femur and tibia.  The Carcharodontosauridae includes the largest land predators in the early and middle Cretaceous of Gondwana, like the popular, Giganotosaurus carolinii. The group evolved large skulls surpassing the length of the largest skull of Tyrannosaurus rex.  Another common trait is the fusion of cranial bones. Gualicho shinyae gen. et sp. nov, a partially articulated mid-sized theropod (about 7.6m long and 450kg in weight) represents a new tetanuran theropod taxon from the Huincul Formation.

Articulated right foot of the holotype of Gualicho shinyae during excavation (from Apesteguía et al., 2016)

Articulated right foot of the holotype of Gualicho shinyae during excavation (from Apesteguía et al., 2016)

The new specimen exhibits a new and unusual combination of characters observed in various remotely related clades including ceratosaurs, tyrannosaurids, and megaraptorans. The didactyl manus with a semilunate distal carpal are indicative of derived tetanuran affinities, while the expanded posterior margin of the metatarsal III proximal articulation, are shared with ceratosaurs. The reduced forelimbs with didactyl manus are similar to those of the tyrannosaurids. However, in tyrannosaurids, the carpal elements are reduced and proximodistally flattened, whereas in Gualicho the semilunate and scapholunare carpals retain a more complex shape typical of the carpal elements of most non-coelurosaurian tetanurans. In addition, the manus of Gualicho differs from tyrannosaurids in having a proportionately more robust metacarpal I with a rectangular, rather than triangular, proximal articulation in end view (Apesteguía et al., 2016).

Left humerus of the of the holotype specimen of Gualicho shinyae (MPCN PV 0001) in (A) anterior, (B) posterior, (C) proximal, and (D) distal views. Abbreviations: dpc, deltopectoral crest; ics, intercondylar sulcus; it, internal tuberosity; msh, scar for insertion of m. scapulohumeralis (From Apesteguía et al., 2016).

Left humerus of the of the holotype specimen of Gualicho shinyae (MPCN PV 0001) in (A) anterior, (B) posterior, (C) proximal, and (D) distal views. Abbreviations: dpc, deltopectoral crest; ics, intercondylar sulcus; it, internal tuberosity; msh, scar for insertion of m. scapulohumeralis (From Apesteguía et al., 2016).

Gualicho shares several derived characters with the African theropod Deltadromeus, including reduced distal humeral articulations, and an expanded lobe bearing a medial trough on the proximocaudal aspect of the fibula. The faunal resemblances between strata in the Neuquén and San Jorge Basins of Patagonia and North African Cenomanian beds are intriguing, but difficult to interpret due to a lack of well sampled, age equivalent strata elsewhere.

Gualicho was discovered on a paleontological expedition led by Sebastian Apesteguía in 2007. The name derived from the Gennaken (Northern Tehuelche languaje) watsiltsüm, an old goddess now considered a source of misfortune. The name was chosen to reflect the difficult circumstances surrounding the discovery and study of the specimen. The specific name honors Ms. Akiko Shinya, Chief Fossil Preparator at the Field Museum.

References:

Apesteguía S, Smith ND, Juárez Valieri R, Makovicky PJ (2016) An Unusual New Theropod with a Didactyl Manus from the Upper Cretaceous of Patagonia, Argentina. PLoS ONE 11(7): e0157793. doi: 10.1371/journal.pone.0157793