Neuroanatomy of the abelisaurid theropod Viavenator exxoni

Viavenator exxoni, Museo Municipal Argentino Urquiza

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 group was erected by Jose Bonaparte with the description of  Abelisaurus Comahuensis. These theropods exhibit spectacular cranial ornamentation in the form of horns and spikes and strongly reduced forelimbs and hands. In South America, braincase remains are known for Carnotaurus sastrei, Abelisaurus comahuensis, Aucasaurus garridoi, Ekrixinatosaurus novasi, Skorpiovenator bustingorryi, Eoabelisaurus and Viavenator exxoni.

The holotype of Viavenator exxoni (MAU-Pv-LI-530) was found in the outcrops of the Bajo de la Carpa Formation (Santonian, Upper Cretaceous), northwestern Patagonia, Argentina. Cranial elements of this specimen include the complete neurocranium: frontals, parietals, sphenethmoids, orbitosphenoids, laterosphenoids, prootics, opisthotics, supraoccipital, exoccipitals, basioccipital, parasphenoids and basisphenoids. The cranial endocast of Viavenator measures 157.7 mm from the olfactory bulbs to the foramen magnum, with a volume of approximately 141.6 cm3. The general shape of cranial endocast is elongate and narrow, similar to Aucasaurus and Majungasaurus. The widest part of the cranial endocast of Viavenator is at the level of the cerebral hemispheres. Four blood vessel foramina are recognized in the braincase: the caudal middle cerebral vein, the rostral middle cerebral vein, the cerebral internal carotid artery and the sphenoid artery.

Figure 1. Rendering of the type braincase of Viavenator exxoni (MAU-Pv-LI-530) in dorsal (A,B), and right lateral (C,D) view. Adapted from Carabajal y Filippi, 2017.

The forebrain of Viavenator include the olfactory tracts and olfactory bulbs, the cerebral hemispheres, optic nerves, the infundibular stalk, and the pituitary body. The CT scans show that the olfatory tracts are undivided. The olfactory bulbs are oval and are separated by a median septum at the anterior region of the sphenethmoids. The optic lobes are not clearly defined. The visible features of the hindbrain in the cranial endocast include the cerebellum, medulla oblongata, and cranial nerves V–XII. The cerebellum is not clearly expanded in the endocast; however, the floccular process of the cerebellum is well defined. The general morphology of both, brain and inner ear of Viavenator is markedly similar to that of Aucasaurus.
Neurosensorial capabilities of extinct animals can be inferred in part based on the relative development of certain regions of the brain. The flocculus of the cerebellum plays a role in coordinate eye movements, and tends to be enlarged in taxa that rely on quick movements of the head and the body. The flocculus of Viavenator is particularly large compared with Majungasaurus, suggesting that Viavenator relied more on quick movements of the head and sophisticated gaze stabilization mechanisms than the African form. The dimensions of the auditory sensory epithelium of Viavenator is similar to Majungasaurus, suggesting that they had similar hearing capabilities. In large dinosaurs, hearing was restricted to low frequencies with high frequency limit below 3 kHz.

References:

Paulina-Carabajal, A., Filippi, L., Neuroanatomy of the abelisaurid theropod Viavenator: The most complete reconstruction of a cranial endocast and inner ear for a South American representative of the clade, Cretaceous Research (2017), doi: 10.1016/j.cretres.2017.06.013

Leonardo S. Filippi, Ariel H. Méndez, Rubén D. Juárez Valieri and Alberto C. Garrido (2016). «A new brachyrostran with hypertrophied axial structures reveals an unexpected radiation of latest Cretaceous abelisaurids». Cretaceous Research 61: 209-219. doi:10.1016/j.cretres.2015.12.018

Solving the mystery of Megatherium diet.

Megatherium americanum, MACN.

Around 10,000 years ago, Argentina was home of numerous species of giant Xenarthrans, giant ground sloths (relative to tree sloth) and glyptodontids (relative to tiny extant armadillo). Sloths, characteristic of the mammal fauna of the Pleistocene of South America, show a great diversity with more than 80 genera, grouped in four families: Megatheriidae, Megalonychidae, Nothrotheriidae and Mylodontidae.

For more than a century different hypotheses on the dietary preferences of giant ground sloths have been proposed. In 1860, Owen gave an extensive explanations about their possible diet and behavior. He based his conclusions on the morphology of the skull, combined with peculiarities of the rest of the skeleton, but always by analogy with living tree sloth. He wrote: “Guided by the general rule that animals having the same kind of dentition have the same kind of food, I conclude that the Megatherium must have subsisted, like the Sloths, on the foliage of tree…”. In 1926, Angel Cabrera discussed the diet of Megatherium, rejecting some theories on myrmecophagy or insectivory, and agreed with Owen’s statements about a folivorous diet.

Megatherium americanum lower right tooth series. Scale bar: 5 cm (From M.S. Bargo and S.F. Vizcaíno, 2008)

The dietary preferences of extinct mammals can usually be evaluated through their tooth morphology, but the application of stable isotopes on fossil bones has yielded very important information to solve debates about the diet of extinct large mammal groups, by comparing the carbon and nitrogen isotopic composition of their bone collagen with those of coeval herbivorous and carnivorous taxa. Another isotopic approach is to mesure the difference between the carbon isotopic abundances of the collagen and the carbonate fractions of skeletal tissues. An animal with a herbivorous diet, exhibits significantly larger differences than a carnivore. The values measured on bone collagen from Megatherium, clearly fall in the same range as the large herbivores such as the equid Hippidion, the notoungulate Toxodon and the liptoptern Macrauchenia, for which there is no doubt about their herbivorous diet. Therefore, the hypotheses of insectivory or carnivory for these extinct mammals are not supported by the isotopic data.

 

References:

Hervé Bocherens et al. Isotopic insight on paleodiet of extinct Pleistocene megafaunal Xenarthrans from Argentina, Gondwana Research (2017). DOI: 10.1016/j.gr.2017.04.003

Bargo, M.S., Vizcaíno, S.F., 2008. Paleobiology of Pleistocene ground sloths (Xenarthra, Tardigrada): biomechanics, morphogeometry and ecomorphology applied to the masticatory apparatus. Ameghiniana 45: 175-196

Introducing Isaberrysaura

Isaberrysaura skull in lateral view and maxillary teeth (Adapted from Salgado et al., 2017)

Isaberrysaura mollensis gen. et sp. nov. is the first dinosaur recovered in the marine-deltaic deposits of the Los Molles Formation (Neuquén Province, Argentina), and the first neornithischian dinosaur known from the Jurassic of South America. So far, the South American record of Jurassic ornithischian dinosaurs was limited to a few specimens belonging to Heterodontosauriformes, a clade of small-sized forms that survived in Europe up to the Early Cretaceous. The name Isaberrysaura is derived from “Isa Berry” (Isabel Valdivia Berry, who reported the initial finding) and the Greek word “saura” (lizard).

The holotype of Isaberrysaura is an incomplete articulated skeleton with an almost complete skull, and a partial postcranium consisting of 6 cervical vertebrae, 15 dorsal vertebrae, a sacrum with a partial ilium and an apparently complete pubis, 9 caudal vertebrae, part of a scapula, ribs, and unidentifiable fragments. One of the most notable features of the discovery is the presence of permineralized seeds in the middle-posterior part of the thoracic cavity. The seeds were assigned to the Cycadales (Zamiineae) on the basis of a well-defined coronula in the micropylar region. The findings suggest the hypothesis of interactions (endozoochory) between cycads and dinosaurs, especially in the dispersion of seeds.

Gut content of Isaberrysaura mollensis gen. et sp. nov. (a–c), seeds of cycads (c), and other seeds (s); rib (r). From Salgado et al., 2017

The cranium of Isaberrysaura is reminiscent of that of the thyreophorans. The skull is estimated to be 52 cm long and 20 cm wide across the orbits. The jugal is triradiate and the nasals are ~20 cm long. There are two supraorbital bones; one is elongated (~10 cm), as in stegosaurs, and the other element interpreted as a posterior supraorbital is located on the posterior margin of the orbit. It has at least six premaxillary teeth, and there is no diastema between the premaxillary and the maxillary tooth row. Despite the many similarities between Isaberrysaura and the thyreophorans, the phylogenetic analysis indicates that Isaberrysaura is a basal ornithopod, suggesting that both Thyreophora and neornithischians could have achieved significant convergent features.

References:

Salgado, L. et al. A new primitive Neornithischian dinosaur from the Jurassic of Patagonia with gut contents. Sci. Rep. 7, 42778; doi: 10.1038/srep42778 (2017)

From Argentina with Love: Top Fossils of 2016

Geographic provenance and speculative reconstruction of the gigantic titanosaurian sauropod dinosaur Notocolossus gonzalezparejasi gen. et sp. nov. (From González Riga  et al., 2016; Credit: Scientific Reports)

Since the discovery of dinosaur remains in the Neuquen basin in 1882, Argentina has gained the title of Land of the Giants. And 2016 has brought us amazing fossil discoveries. From Notocolossus to Gualicho, my fossil pick for this year are:

  • Notocolossus

Notocolossus gonzalezparejasi gen. et sp. nov. from the Upper Cretaceous of Mendoza Province, Argentina is one of the largest known dinosaurs. The name derived from the Greek notos (southern) and the Latin colossus, in reference to the gigantic size and Gondwanan provenance of the new taxon. The species name honours Dr. Jorge González Parejas, who provided legal guidance on the research, protection, and preservation of dinosaur fossils from Mendoza Province. The holotype of Notocolossus (UNCUYO-LD 301) consists of a partial skeleton lacking the skull. It contains an anterior dorsal vertebra, an anterior caudal vertebra, the right humerus (with 1.76 m in length), and the proximal end of the left pubis. The pes of  Notocolossus is comparatively shorter and more mediolaterally symmetrical than those of other titanosaurs, and indeed, most other sauropods. Notocolossus also presents truncated unguals, characteristics otherwise unknown in the Sauropoda.

Cranium of Sarmientosaurus musacchioi in right lateral view. Scale bar = 10 cm. (From Martínez et al., 2016)

Cranium of Sarmientosaurus musacchioi in right lateral view. Scale bar = 10 cm. (From Martínez et al., 2016)

  • Sarmientosaurus

Another remarkable new species of titanosaurian sauropod was Sarmientosaurus musacchioi. The holotypic and only known specimen consists of an articulated, virtually complete skull and part of the cranial and middle cervical series. The new titanosaur comes from the Lower Member of the Upper Cretaceous Bajo Barreal Formation on the Estancia Laguna Palacios near the village of Buen Pasto in south-central Chubut Province, central Patagonia, Argentina. It is the most basal known titanosaur to be represented by a well-preserved skull. Furthermore, the cranial endocast preserves some of the most complete information about the brain and sensory system for any sauropod.

Scapulocoracoid of Viavenator exxoni gen. et sp. nov. MAU-Pv-LI-530. in lateral view. Scale bar: 10 cm

Scapulocoracoid of Viavenator exxoni in lateral view. Scale bar: 10 cm (a, acromion; cf, coracoid foramen; gc, glenoid cavity; pvp, posteroventral process. From Filippi et al., 2016)

  • Viavenator

The holotype of Viavenator exxoni was found in the outcrops of the Bajo de la Carpa Formation (Santonian, Upper Cretaceous), northwestern Patagonia, Argentina. The new taxon belongs to the South American clade of abelisaurid and possesses, among other characteristics, hypertrophied structures in the presacral axial skeleton. The name derives from the latin word ‘Via’ (road) and ‘venator’ (hunter), meaning the hunter of the road; ‘exxoni’ is in recognition of Exxonmobil’s commitment to the preservation of paleontological heritage of the La Invernada area, Rincón de los Sauces, Neuquen, Patagonia Argentina.

Right postorbital (holotype) of Taurovenator violentei gen. et sp. nov. A, lateral view

Right postorbital (holotype) of Taurovenator violentei gen. et sp. nov. A, lateral view. Scale bar: 3 cm (From Motta et al., 2016)

  • Taurovenator.

Taurovenator violantei gen. et sp. nov. was is a medium-sized carcharodontosaurid theropod from the Huincul Formation (Upper Cretaceous) in northwestern Río Negro province, Patagonia, Argentina. The generic name derives from the Latin words “tauro” (Bull) and “venator” (Hunter). The specific name honours Enzo Violante, owner of the farm where the specimen was discovered. Taurovenator is similar in gross morphology to Giganotosaurus, Carcharodontosaurus, and Mapusaurus, but shows two unique features: the presence of a horn-like structure in the orbital brow and the presence of an excavation housed at the posterodorsal surface of the eye socket.

Different appendicular elements of Murusraptor in their original burial positions (From Coria et al., 2016)

Different appendicular elements of Murusraptor in their original burial positions (From Coria et al., 2016)

  • Murusraptor

Murusraptor barrosaensis, from the Upper Cretaceous of Neuquén Province, Argentina, belongs to a Patagonian radiation of megaraptorids together with Aerosteon, Megaraptor and Orkoraptor. Murusraptor, meaning “Wall Raptor”, was discovered in a canyon wall in 2001 during an expedition to Sierra Barrosa in northwestern Patagonia. The holotype specimen includes much of the skull, axial skeleton, pelvis and tibia. The braincase is intact and most of the sutures are still visible, indicating that this was not a fully mature animal. Murusraptor barrosaensis is unique in having anterodorsal process of lacrimal longer than height of preorbital process; sacral ribs hollow and tubelike; short ischia distally flattened and slightly expanded dorsoventrally. Murusraptor also exhibits some characters that are interpreted as convergencies of this taxon with non-tyrannosauroid theropods, including lacrimal with a small pneumatic recess; and a highly pneumatic braincase.

Gualicho shinyae, at the Centro Cultural de la Ciencia.

Gualicho shinyae, at the Centro Cultural de la Ciencia.

  • Gualicho

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. The 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.

 

References:

Bernardo J. González Riga et al. A gigantic new dinosaur from Argentina and the evolution of the sauropod hind foot, Scientific Reports (2016). DOI: 10.1038/srep19165

Martínez R.D.F. et al. 2016. A Basal Lithostrotian Titanosaur (Dinosauria: Sauropoda) with a Complete Skull: Implications for the Evolution and Paleobiology of Titanosauria. PLoS ONE 11 (4): e0151661; doi: 10.1371/journal.pone.0151661

Leonardo S. Filippi, Ariel H. Méndez, Rubén D. Juárez Valieri and Alberto C. Garrido (2016). «A new brachyrostran with hypertrophied axial structures reveals an unexpected radiation of latest Cretaceous abelisaurids». Cretaceous Research 61: 209-219. doi:10.1016/j.cretres.2015.12.018

Matías J. Motta, Alexis M. Aranciaga Rolando, Sebastián Rozadilla, Federico E. Agnolín, Nicolás R. Chimento, Federico Brissón Egli, and Fernando E. Novas (2016). «New theropod fauna from the Upper Cretaceous (Huincul Formation) of northwestern Patagonia, Argentina». New Mexico Museum of Natural History and Science Bulletin 71: 231-253

Rodolfo A. Coria, Philip J. Currie. A New Megaraptoran Dinosaur (Dinosauria, Theropoda, Megaraptoridae) from the Late Cretaceous of Patagonia. PLOS ONE, 2016; 11 (7): e0157973 DOI: 10.1371/journal.pone.0157973

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

A brief introduction to the Early dinosaurs from Argentina.

eoraptor-skeleton

Articulated skeleton of Eoraptor lunensi (From Sereno 2013)

The oldest record of Argentinean dinosaurs comes from the Ischigualasto Formation, NW Argentina, dated from 231.4 Ma to 225.9 Ma. Adolf Stelzner in 1889 published the first data on the geology of Ischigualasto, but it was not until 1911 that Guillermo Bodenbender briefly refers to the fossils of the site. In the early 40′s, Joaquin Frenguelli, initiates a geological survey in the western margin of the basin. Later, in 1943, Angel Cabrera described fragmentary therapsid fossils. However, intensive paleontological study of the Ischigualasto and Chañares Formations, began only in the late 1950s.

The Ischigualasto Formation has 300–700 m of mudstone, sandstone, conglomerate, and basalt, and consists of four lithostratigraphic members which in ascending order include the La Peña Member, the Cancha de Bochas Member, the Valle de la Luna Member, and the Quebrada de la Sal Member. Eight valid species of dinosaurs are known from the Ischigualasto Formation: Pisanosaurus mertii, Herrerasaurus ischigualastensis, Sanjuansaurus gordilloi, Eodromaeus murphi, Eoraptor lunensis, Panphagia protos, and Chromogisaurus novasi.

 

Skull of Herrerasaurus ischigualastensis (Sereno, 2013)

Skull of Herrerasaurus ischigualastensis (Sereno, 2013)

Pisanosaurus mertii is a small specimen, know by an incomplete maxilla and lower jaw fragments bearing teeth, vertebrae, incomplete hind limb, and the impression of the pelvis. Described in 1967 by Rodolfo CasamiquelaPisanosaurus is considered as the oldest known ornithischian.

Herrerasaurus ischigualastensis was described by Osvaldo Reig in 1963. The taxon is one of the best known Triassic dinosaurs and the largest dinosaur of the Ischigualasto Formation. Herrerasaurus was fully bipedal, with strong hind limbs, short thighs and long feet. The skull has a rectangular profile and a transversely narrow snout (Sereno and Novas, 1992). The presence of two sacral vertebrae and lack of brevis fossa made Herrerasaurus, and other herrerasaurids, a controversial group.

Sanjuansaurus gordilloi is similar to Herrerasaurus ischigualastensis, although more gracile and possessing short and straight pubis among other differences (Alcober & Martínez, 2010). It’s known from one specimen that preserves left maxilla, partial axial column, scapulae, left ulna, ungual of manual digit III, partial left ilium and pubis, both femora and tibiae, right fibula, right astragalus and calcaneum, and left metatarsal.

Skull and skeleton of Eodromaeus murphi (PVSJ 560). Scale bar equals 10 cm.

Skull and skeleton of Eodromaeus murphi
(PVSJ 560). Scale bar equals 10 cm.

Eodromaeus murphi is a small species with a total length of about 1.2 metres, known from five specimens. The trunk was long and slender, and forelimbs were shorter than the hindlimbs. The skull is relatively low and lightly built with a relatively spacious antorbital fenestra.  A phylogenetic analysis places Eodromaeus within Theropoda as the sister taxon to Neotheropoda

Eoraptor lunensis is known from eight specimens, including the holotype that preserves most of the skeleton. Eoraptor had a slender body with an estimated weight of about 10 kilograms. The lightly built skull has a slightly enlarged external naris and the premaxilla is observed to have a slender posterolateral process. The long bones of the hind limb have more robust shafts than those of Eodromaeus, although in both genera the tibia remains slightly longer than the femur (Sereno et al., 2013). Initially considered a basal theropod, the sauropodomorph affinity of Eoraptor has been strengthened after the publication of its anatomy in 2013.

Panphagia protos is a small species, known from one partial skeleton including several skull bones, lower jaw, and partial axial skeleton. The specimen is an immature individual with an estimated body length of approximately 1.30 m. It was originally proposed as the most basal sauropodomorph (Martinez and Alcober, 2009)

Chromogisaurus novasi is also similar in size to Eoraptor lunensis. It’s known from a partial skeleton lacking the skull. It includes elements of the front and hind limbs, the pelvis and two caudal vertebrae.

References:

Martín D. EZCURRA & Ricardo N. MARTÍNEZ (2016), Dinosaur precursors and early dinosaurs from Argentina., In book: Historia Evolutiva y Paleobiogeografía de los Vertebrados de América del Sur, Publisher: Contribuciones del MACN, Editors: F. Agnolíin, G.L. Lio, F. Brissón Egli, N.R. Chimento, F. Novas, pp.97-107

Reig, O.A. (1963). “La presencia de dinosaurios saurisquios en los “Estratos de Ischigualasto” (Mesotriásico Superior) de las provincias de San Juan y La Rioja (República Argentina)”. Ameghiniana (in Spanish). 3 (1): 3–20.

Sereno, P.C.; Novas, F.E. (1992). “The complete skull and skeleton of an early dinosaur”. Science. 258 (5085): 1137–1140.

Ricardo N. Martinez; Paul C. Sereno; Oscar A. Alcober; Carina E. Colombi; Paul R. Renne; Isabel P. Montañez; Brian S. Currie (2011). “A Basal Dinosaur from the Dawn of the Dinosaur Era in Southwestern Pangaea”. Science. 331 (6014): 206–210. doi:10.1126/science.1198467

Martinez RN, Alcober OA (2009) A Basal Sauropodomorph (Dinosauria: Saurischia) from the Ischigualasto Formation (Triassic, Carnian) and the Early Evolution of Sauropodomorpha. PLoS ONE 4(2): e4397. doi:10.1371/journal.pone.0004397

Ezcurra, M. D. 2010. “A new early dinosaur (Saurischia: Sauropodomorpha) from the Late Triassic of Argentina: a reassessment of dinosaur origin and phylogeny.” Journal of Systematic Palaeontology 8: 371-425.