Murusraptor barrosaensis, a new species in the megaraptorid clade.

Body reconstruction of Murusraptor barrosaensis (From Coria et al., 2016)

Body reconstruction of Murusraptor barrosaensis (From Coria et al., 2016)

Patagonia has yielded the most comprehensive fossil record of Cretaceous theropods from Gondwana, including Megaraptora, a clade of medium-sized and highly pneumatized theropods represented by Megaraptor, Orkoraptor and Aerosteon, and characterized by the formidable development of their manual claws on digits I and II and the transversely compressed and ventrally sharp ungual of the first manual digit (Novas et al, 2013). The enigmatic nature of this group has been a matter of discussion since the description of the first megaraptoran, Megaraptor namunhaiquii. For years, Megaraptor has been alternatively interpreted as belonging to different theropod lineages: as basal coelurosaurians (Novas,1998), basal tetanurans (Calvo et al.,2004; Smith et al., 2008), and allosauroids closely related with carcharodontosaurids (Smith et al., 2007; Benson et al., 2010; Carrano et al., 2012). The main reason for so many different interpretations is the incomplete nature of most available megaraptorid skeletons and the little information about their cranial anatomy.

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.

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 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 shares with all Megaraptoridae two unambiguous synapomorphies: teeth with no enamel wrinkles (interpreted as a reversion to primitive condition in Theropoda); and anterior caudal vertebrae with neural arch bearing prominent centrodiapophysial laminae that define a deep infradiapophysial fossa. 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 (Coria et al., 2016)

References:

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

Porfiri, J. D., Novas, F. E., Calvo, J. O., Agnolín, F. L., Ezcurra, M. D. & Cerda, I. A. 2014. Juvenile specimen of Megaraptor (Dinosauria, Theropoda) sheds light about tyrannosauroid radiation. Cretaceous Research 51: 35-55.

 

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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

A Tale of Two Exctintions.

The permian triassic boundary at Meishan, China (Photo: Shuzhong Shen)

The Permian Triassic boundary at Meishan, China (Photo: Shuzhong Shen)

Extinction is the ultimate fate of all species. The fossil record indicates that more than 95% of all species that ever lived are now extinct. Over the last 3 decades, mass extinction events  have become the subject of increasingly detailed and multidisciplinary investigations. In 1982, Jack Sepkoski and David M. Raup identified five major extinction events in Earth’s history: at the end of the Ordovician period, Late Devonian, End Permian, End Triassic and the End Cretaceous. These five events are know as the Big Five.

The end-Permian extinction is the most severe biotic crisis in the fossil record, with as much as 95% of the marine animal species and a similarly high proportion of terrestrial plants and animals going extinct . This great crisis occurred 252 million years ago (Ma) during an episode of global warming. The End-Triassic Extinction  is probably the least understood of the big five. Most mammal-like reptiles and large amphibians disappeared, as well as early dinosaur groups. In the oceans, this event eliminated conodonts and nearly annihilated corals, ammonites, brachiopods and bivalves. Although it’s almost impossible briefly summarize all the changes in biodiversity associated with both extinction events, we can describe their broad trends.

 

Flow chart summarizing proposed cause-and-effect relationships during the end-Permian extinction (From Bond and Wignall, 2014)

Flow chart summarizing proposed cause-and-effect relationships during the end-Permian extinction (From Bond and Wignall, 2014)

Both extinction events are commonly linked to the emplacement of the large igneous provinces of the Siberian Traps and the Central Atlantic Magmatic Province. Massive volcanic eruptions with lava flows, released large quantities of sulphur dioxide, carbon dioxide, thermogenic methane and large amounts of HF, HCl, halocarbons and toxic aromatics and heavy metals into the atmosphere. Furthermore, volcanism contribute gases to the atmosphere, such as Cl, F, and CH3Cl from coal combustion, that suppress ozone formation. Acid rain likely had an impact on freshwater ecosystems and may have triggered forest dieback. Mutagenesis observed in the Lower Triassic herbaceous lycopsid Isoetales has been attributed to increased levels of UV-radiation. Charcoal records point to forest fires as a common denominator during both events. Forest dieback was accompanied by the proliferation of opportunists and pioneers, including ferns and fern allies. Moreover, both events led to major schisms in the dominant terrestrial herbivores  and apex predators, including the late Permian extinction of the pariaeosaurs and many dicynodonts and the end-Triassic loss of crurotarsans (van de Schootbrugge and Wignall, 2016).

Aberrant pollen and spores from the end-Triassic extinction interval (scale bars are 20 μm). (a) Ricciisporites tuberculatus from the uppermost Rhaetian deposits at Northern Ireland (adapted from van de Schootbrugge and Wignall, 2016)

Aberrant pollen and spores from the end-Triassic extinction interval (scale bars are 20 μm). (a) Ricciisporites
tuberculatus and b) Kraeuselisporites reissingerii (adapted from van de Schootbrugge and Wignall, 2016)

During the end-Permian Event, the woody gymnosperm vegetation (cordaitaleans and glossopterids) were replaced by spore-producing plants (mainly lycophytes) before the typical Mesozoic woody vegetation evolved. The palynological record suggests that wooded terrestrial ecosystems took four to five million years to reform stable ecosystems, while spore-producing lycopsids had an important ecological role in the post-extinction interval. A key factor for plant resilience is the time-scale: if the duration of the ecological disruption did not exceed that of the viability of seeds and spores, those plant taxa have the potential to recover (Traverse, 1988). Palynological records from across Europe provide evidence for complete loss of tree-bearing vegetation reflected in a strong decline in pollen abundance at the end of the Triassic. In the Southern Hemisphere, the vegetation turnover consisted in the replacement to Alisporites (corystosperm)-dominated assemblage to a Classopollis (cheirolepidiacean)-dominated one.

Comparison of extinction rates for calcareous organisms during the end-Permian and end-Triassic extinction event (from van de Schootbrugge and Wignall, 2016)

Comparison of extinction rates for calcareous organisms during the end-Permian and end-Triassic extinction event (from van de Schootbrugge and Wignall, 2016)

Rapid additions of carbon dioxide during extreme events may have driven surface waters to undersaturation. Acidification affects the biogeochemical dynamics of calcium carbonate, organic carbon, nitrogen, and phosphorus in the ocean and interferes with a range of processes, including growth, calcification, development, reproduction and behaviour in a wide range of marine organisms like foraminifera, planktonic coccolithophores, pteropods and other molluscs,  echinoderms, corals, and coralline algae. Both extinction events led to near-annihilation of cnidarian clades and other taxa responsible for reef construction, resulting in ‘reef gaps’ that lasted millions of years. Black shales deposited across both extinction events also contain increased concentrations of the biomarker isorenieratane, a pigment from green sulphur bacteria, suggesting that the photic zone underwent prolonged periods of high concentrations of hydrogen sulphide. Following the end-Triassic extinction, Early Jurassic shallow seas witnessed recurrent euxinia over a time span of 25 million years, culminating in the Toarcian Oceanic Anoxic Event.

 

References:

BAS VAN DE SCHOOTBRUGGE and PAUL B. WIGNALL (2016). A tale of two extinctions: converging end-Permian and end-Triassic scenarios. Geological Magazine, 153, pp 332-354. doi:10.1017/S0016756815000643.

BACHAN, A. & PAYNE, J. L. 2015. Modelling the impact of pulsed CAMP volcanism on pCO2 and δ13C across the Triassic-Jurassic transition. Geological Magazine, published online

Retallack, G.J. 2013. Permian and Triassic greenhouse crises. Gondwana Research 24:90–103.