Introducing Kaijutitan, the strange beast.

The entrance to the town of Rincón de los Sauces.

Since the discovery of dinosaur remains in the Neuquen basin in 1882, Argentina has gained the title of Land of the Giants. The tittle was reinforced by the discoveries of titanosaurs like Argentinosaurus, Dreadnoughtus, Notocolossus, Puertasaurus, and Patagotitan. The study of this diverse group of sauropod dinosaurs embrace an extensive list of important contributions, which started with Richard Lydekker’s pioneering work on Patagonian dinosaurs, and by the classic Friedrich von Huene monograph on Argentinean saurischians and ornithischians.

Titanosaurus were a diverse group of sauropod dinosaurs represented by more than 30 genera, which included all descendants of the more recent common ancestor of Andesaurus and Saltasaurus. The group includes the smallest (e.g. Rinconsaurus, Saltasaurus; with estimated body masses of approximately 6 tonnes) and the largest sauropods known to date. They had their major radiation during the middle Early Cretaceous. The evolution of body mass in this clade is key element to understand sauropod evolution.

 

Cranial elements of MAU-Pv-CM-522/1. From Filippi et al., 2019.

Kaijutitan maui, is the first basal sauropod titanosaur from the Sierra Barrosa Formation (Upper Coniacian, Upper Cretaceous). The holotype (MAU-Pv-CM-522) consists of cranial, axial, and appendicular elements presenting an unique combination of plesiomorphic and apomorphic characters. The generic name Kaijutitan is derived from Kaiju, Japanese word that means “strange beast” or “monster”, and titan, from the Greek “giant”.  The species name refers to the acronym of the Museo Municipal Argentino Urquiza, Rincon de los Sauces, Neuquén, Argentina.

The cranial elements of this specimen include the complete neurocranium (the supraoccipital, exoccipital, left paraoccipital process, left exoccipital-opisthotic-prootic complex, left laterosphenoid and orbitosphoid, and basioccipital-basisphenoid complex). The impossibility of recognizing clear sutures suggest an ontogenetic adult stage of the specimen. One of the most notable autapomorphies exhibited by Kaijutitan is the anterior cervical vertebra with bifid neural spine, a feature usually found in diplodocids and dicraeosaurids. Unfortunately, the femur and humerus of Kaijutitan maui are incomplete, therefore the body mass of this titanosaur can only be estimated by comparison with other titanosauriforms. Kaijutitan would have had a body mass similar or intermediate to that of Giraffatitan (38.000 kg) and Notocolossus (60.398 kg).

 

References:

Filippi, L.S., Salgado, L., Garrido, A.C., A new giant basal titanosaur sauropod in the Upper Cretaceous (Coniacian) of the Neuquén Basin, Argentina, Cretaceous Research, https://doi.org/10.1016/j.cretres.2019.03.008.

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The mounting of the cast of Diplodocus carnegii at the Museo de La Plata.

Diplodocus carnegii at the Museo de La Plata, 1912 (From Otero and Gasparini, 2014)

Diplodocus carnegii at the Museo de La Plata, 1912 (From Otero and Gasparini, 2014).

Diplodocus is one of the most popular dinosaurs of all time. The first remains of a Diplodocus were found by Benjamin Mudge and Samuel Wendell Williston, in the Upper Jurassic outcrops of Cañon City, Colorado, United States, in 1877. One year later, Othniel Charles Marsh named the species Diplodocus longus on the basis of remains of the hind limb and tail. The name Diplodocus means ‘double beam’ in reference to the particular two-pronged morphology of the posterior hemal arches. D. carnegii, was discovered in 1899 during an expedition carried out by the Carnegie Museum to the Upper Jurassic Morrison Formation of Wyoming. John Bell Hatcher dedicated the new species of to Andrew Carnegie.

A sketch from the of Diplodocus carnegii, which Carnegie had framed and mounted on a wall at his castle in Scotland.

William Jacob Holland, director of the Carnegie Museum, sent a sketch of the skeleton of Diplodocus to Andrew Carnegie. At the time, the steel tycoon was at his Castle, Skibo, in Sutherland County, Scotland. The King Edward VII of England, saw the sketch and asked Carnegie to give him a specimen for the British Museum of Natural History in London. Holland proposed to Carnegie to make a life-sized replica of D. carnegii to be given to the British Museum of Natural History. On May 12, 1905, the long skeleton was unveiled to a crowd of 300 people, and became an instant star.

Mounting of the cast of Diplodocus carnegii at the Museo de La Plata, Argentina. Arthur Coggeshall and William Holland are second and third from left (Adapted from ‘Caras y Caretas’ magazine, 1912).

Nine replicas of D. carnegii were made and donated to kings and presidents of Europe and Latin America. On November of 1911, Argentinean president Dr. Roque Saenz Peña communicated to Andrew Carnegie his request to have a replica of D. carnegii. His request was accepted, and on July 1, 1912, 34 boxes containing the cast of the animal were sent to Argentina on the S.S. ‘Sallust’. William Holland and Mr. Arthur Coggeshall were in charge of mounting the replica. The site where the replica would be mounted in the Museo de La Plata, would be the Sala III, which was dedicated to invertebrates and plants. Holland insisted that the plans used for the mounting of D. carnegii at Vienna were followed in mounting the skeleton in La Plata. After the skeleton was mounted, the Director of the Museum, Dr. Samuel Lafone-Quevedo, gave a speech expressing his gratitude to Andrew Carnegie and his representatives, in which William Holland was designated an Honorary Member of La Plata University.

Reference:

Alejandro Otero and Zulma Gasparini “The History of the Cast Skeleton of Diplodocus carnegii Hatcher, 1901, at the Museo De La Plata, Argentina,” Annals of Carnegie Museum 82(3), (2014). doi: http://dx.doi.org/10.2992/007.082.0301

BARRETT, P., P. PARRY, AND S. CHAPMAN. 2010. Dippy: The Tale of a Museum Icon. Natural History Museum, London. 48 pp.

 

Introducing Bajadasaurus pronuspinax.

Bajadasaurus reconstruction (Museo Municipal Ernesto Bachmann, Villa El Chocón, Neuquén).

Dicraeosauridae is a family of mid-sized sauropod dinosaurs characterized by a distinctive vertebral column with paired, long, neural spines. The group was first described in 1914 by Werner Janensch with the discovery of the nearly complete skeletons of Dicraeosaurus in the expeditions to the upper Jurassic beds of Tendaguru, Tanzania. Dicraeosauridae includes  Amargasaurus, Pilmatueia, Suuwassea, and Brachytrachelopan. Now, the description of Bajadasaurus pronuspinax gen. et sp. nov., from the Early Lower Cretaceous of Bajada Colorada Formation in Northern Patagonia, Argentina), shed new light on the function of its spines and the defense behavior in sauropod dinosaurs.

Bajadasaurus was discovered in 2013, by a team of paleontologists from CONICET, Fundación Félix de Azara, Universidad Maimónides, and Museo Paleontológico Ernesto Bachmann. The generic name derived from Bajada (Spanish, in reference to the locality Bajada Colorada) and saurus (Greek, lizard). The specific name derived from pronus (Latin, bent over forward) and spinax (Greek, spine), referring to the anteriorly pointed, curved, neural spines of the cervical vertebrae.

Skeletal elements of Bajadasaurus pronuspinax. From Gallina et al., 2019.

The holotype, MMCh-PV 75, includes a nearly complete skull (left maxilla, left lacrimal, both prefrontals, both frontals, both parietals, both postorbitals, both squamosals, left quadratojugal, both pterygoids, both quadrates, supraoccipital, exoccipital-opisthotic complex, basioccipital, basisphenoid, both prootics, both laterosphenoids, both orbitosphenoids, both dentaries, left surangular, both angulars, both splenials, left prearticular, left articular, isolated upper tooth row), both proatlases, atlantal neurapophyses, axis and the fifth cervical vertebra.

The skull of Bajadasaurus is gracile, with dorsally exposed orbits, dorsoventrally compressed occipital condyle, extremely narrow basipterygoid processes, elongate and slender anterior processes of the squamosals, medially extended post-temporal fenestrae, short lateral temporal fenestrae and a reduced dentition in the maxilla and dentary, that largely differs from other known taxa within Dicraeosauridae. But the most striking feature of Bajadasaurus is the presence of extremely long cervical neural spines that curve anteriorly. Amargasaurus exhibit the same development of cervical neural spine elongation as Bajadasaurus, but the spines of the former point backwards rather than forwards. Dicraeosaurus and Brachytrachelopan show anteriorly inclined neural spines in the cervical vertebrae, but the spines are much shorter than in Bajadasaurus.

A group of Bajadasaurus. Illustration: Jorge A. González.

The discovery of Amargasaurus cazaui in 1991, from the Early Cretaceous beds of La Amarga Formation of Northern Patagonia, renewed the discussion on the peculiar vertebral anatomy of these sauropod dinosaurs, including interpretations as a support structure for a thermoregulatory sail, a padded crest as a display and/or clattering structure, a dorsal hump, or as internal cores of dorsal horn. Those explanation, except the last one, require that these long and extremely gracile bone projections, now recognized in Bajadasaurus as well, can support enough physical stress to avoid fracturing. Bone is stronger and stiffer in passive situations, however, horns and other keratin-based materials are tougher and highly resistant to impact-related fractures. Therefore, the keratinous sheath in Amargasaurus and perhaps Bajadasaurus provides a better mechanical solution against a potential fracture.

 

References:

Gallina, Pablo A., Apesteguía, Sebastián, Canale, Juan I., Haluza, Alejandro (2019), A new long-spined dinosaur from Patagonia sheds light on sauropod defense system, Scientific Reports volume 9, Article number: 1392 DOI: https://doi.org/10.1038/s41598-018-37943-3

Janensch, W. Die Wirbelsäule der Gattung Dicraeosaurus. Palaeontographica Supplement 7, 37–133 (1929).

Salgado, L. & Bonaparte, J. F. Un nuevo saurópodo Dicraeosauridae, Amargasaurus cazaui gen et sp. nov., de la Formación La Amarga, Neocomiano de la provincia del Neuquén, Argentina. Ameghiniana 28, 333–346 (1991).

Top fossils discoveries of 2018.

Ingentia prima outcropping from the soil.

Paraphrasing Dickens, 2018 was the best of years, and it was the worst of years. Marked by extreme weather, earthquakes, and an intense volcanic activity, 2018 is also noted by amazing fossil discoveries. My top list include:

  • The oldest Archaeopteryx

Articulated dorsal vertebral column of the new Archaeopteryx, including dorsal ribs and gastralia. Scale bar is 10 mm. (From Rauhut et al., 2018)

The Archaeopteryx story began in  the summer of 1861, two years after the publication of the first edition of Darwin’s Origin of Species, when workers in a limestone quarry in Germany discovered the impression of a single 145-million-year-old feather. Over the years, eleven Archaeopteryx specimens has being recovered. The new specimen from the village of Schamhaupten, east-central Bavaria is the oldest representative of the genus (earliest Tithonian). The shoulder girdles and arms, as well as the skull have been slightly dislocated from their original positions, but the forelimbs remain in articulation. The skull is triangular in lateral outline and has approximately 56 mm long. The orbit is the largest cranial opening (approximately 16 mm long), and the lateral temporal fenestra is collapsed. There are probably four tooth positions in the premaxilla, nine in the maxilla and 13 in the dentary. The postcranial skeleton was affected by breakage and loss of elements prior to or at the time of discovery.

  • Tratayenia rosalesi

Fossilized vertebrae and right hip bone of Tratayenia rosalesi. From Porfiri et al., 2018

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 Fukuiraptor, Aerosteon, Australovenator, Megaraptor, Murusraptor, and Orkoraptor, 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. Tratayenia rosalesi is the first megaraptoran theropod described from the Santonian Bajo de la Carpa Formation of the Neuquén Group. The genus name is for Tratayén, the locality where the holotype was collected. The specific name honors Diego Rosales, who discovered the specimen in 2006. Tratayenia is also the largest carnivorous taxon known from Bajo de la Carpa Formation, reinforcing the hypothesis that megaraptorids were apex predators in South America from the Turonian through the Santonian or early Campanian, following the extinction of carcharodontosaurids.

  • Lingwulong shenqi

Skeletal reconstruction and exemplar skeletal remains of Lingwulong shenqi. Scale bars = 100 cm for a and 5 cm for b–o. From Xi et al., 2018

Sauropods were the largest terrestrial vertebrates. Their morphology is easy recognizable: a long, slender neck and a tail at the end of a large body supported by four columnar limbs. Sauropods dominated many Jurassic and Cretaceous terrestrial faunas. Although they were globally distributed, the absence of Diplodocoidea from East Asia has been interpreted as a biogeographic pattern caused by the Mesozoic fragmentation of Pangea. Lingwulong shenqi — literally the “amazing dragon from Lingwu” — is the first well-preserved confirmed diplodocoid from East Asia (23 synapomorphies support the placement of Lingwulong within Diplodocoidea with 10 of these being unequivocal). The holotype, (LM) V001a, is a partial skull comprising the braincase, skull roof, and occiput, and an associated set of dentary teeth. The paratype, (LGP) V001b, comprises a semi-articulated partial skeleton including a series of posterior dorsal vertebrae, complete sacrum, the first caudal vertebra, partial pelvis, and incomplete right hind limb. The Lingwulong specimens were found in the Yanan Formation at Ciyaopu, in northwest China. The presence of a conchostracans assemblage (including Palaeoleptoestheria, Triglypta, and Euestheria) is indicative of a Middle Jurassic age. The discovery of Lingwulong undermines the EAIH (East Asian Isolation Hypothesis), forcing a significant revision of hypotheses concerning the origins and early radiation of Neosauropoda.

  • Ingentia prima

Skeletal anatomy of Ingentia prima (From Apaldetti et al., 2018)

Ingentia prima — literally the “first giant” in Latin — from the Late Triassic of Argentina shed new lights on the origin of gigantism in this group. The holotype, PVSJ 1086, composed of six articulated posterior cervical vertebrae, glenoid region of right scapula and right forelimb lacking all phalanges, has been recovered from the southern outcrops of the Quebrada del Barro Formation, northwestern Argentina. Discovered in 2015 by Diego Abelín and a team led by Cecilia Apaldetti of CONICET-Universidad Nacional de San Juan, Argentina, this new fossil weighed up to 11 tons and measured up to 32 feet (10 meters) long. Ingentia was unearthed with three new specimens of Lessemsaurus sauropoides. The four dinosaurs belongs to the clade Lessemsauridae, that differs from all other Sauropodomorpha dinosaurs in possessing robust scapulae with dorsal and ventral ends equally expanded; slit-shaped neural canal of posterior dorsal vertebrae; anterior dorsal neural spines transversely expanded towards the dorsal end; a minimum transverse shaft width of the first metacarpal greater than twice the minimum transverse shaft of the second metacarpal; and bone growth characterized by the presence of thick zones of highly vascularized fibrolamellar bone, within a cyclical growth pattern.

  • Caelestiventus hanseni

A 3D printed model of the C. hanseni skull discovered in Utah

Caelestiventus hanseni, from the Upper Triassic of North America, is the oldest pterosaur ever discovered, and it predates all known desert pterosaurs by more than65 million years. The holotype, BYU 20707, includes the left maxilla fused with the jugal, the right maxilla, the right nasal, the fused frontoparietals, the right and left mandibular rami, the right terminal wing phalanx and three fragments of indeterminate bones. The maxilla, jugal, frontoparietal, and mandibular rami of the specimen are pneumatic. The unfused skull and mandibular elements suggest that BYU 20707 was skeletally immature or had indeterminate growth. Based on the relationship between the length of the terminal wing phalanges and wing span in other non-pterodactyloid pterosaurs the new taxon would have a wing span greater than 1.5 m. The significance of C. hanseni lies in its exceptional state of preservation, and its close phylogenetic relationship with Dimorphodon macronyx, indicating that dimorphodontids originated by the Late Triassic and survived the end-Triassic extinction event.

  • Macrocollum itaquii

Skull of Macrocollum itaquii (From Müller et al 2018)

Macrocollum itaquii is the oldest long-necked dinosaur known. Discovered in 2012, from rocks belonging to the upper part of the Candelaria Sequence constrained as about 225 Ma, the three individuals described as M. itaquii are relatively well preserved. The holotype specimen (CAPPA/UFSM 0001a) consists of an almost complete and articulated skeleton. The two paratype specimens (CAPPA/UFSM 0001b and CAPPA/UFSM 0001c) are both articulated skeletons with one missing a skull and its cervical series. The clustered preservation of the three skeletons also represents the oldest evidence of gregarious behaviour in sauropodomorphs, a pattern seen in other Triassic associations, such as the ‘Plateosaurus bonebed’ from Central Europe, and the Mussaurus remains from the Laguna Colorada Formation, Argentina. M. itaquii was only 3.5 meters long and weighed about 101.6 kilograms. In contrast to most Carnian members of the group, the teeth of M. itaquii and other Norian taxa are fully adapted to an omnivore/herbivore diet. The neck elongation may also have provided a competitive advantage for gathering food resources, allowing members of the group to reach higher vegetation. The modifications of the hindlimb of M. itaquii could be related to the progressive loss of cursorial habits.

  • Soft-tissue evidence in a Jurassic ichthyosaur.

Stenopterygius specimen from the Holzmaden quarry. Credit: Johan Lindgren

During the Norian, the evolution of ichthyosaurs took a major turn, with the appearance of the clade Parvipelvia (ichthyosaurs with a small pelvic girdle). They were notably similar in appearance to extant pelagic cruisers such as odontocete whales. An exquisitely fossilized parvipelvian Stenopterygius from the Early Jurassic (Toarcian) of the Holzmaden quarry in southern Germany, indicates that their resemblance with dolphin and whales is more than skin deep. The specimen (MH 432; Urweltmuseum Hauff, Holzmaden, Germany) reveals endogenous cellular, sub-cellular and biomolecular constituents within relict skin and subcutaneous tissue. The external surface of the body is smooth, and was presumably comparable in life to the skin of extant cetaceans. The histological and microscopic examination of the fossil, evinced a multi-layered subsurface architecture. The approximately 100-μm-thick epidermis retains cell-like structures that are likely to represent preserved melanophores. The subcutaneous layer is over 500 μm thick, and comprises a glossy black material superimposed over a fibrous mat. The anatomical localization, chemical composition and fabric of the subcutaneous material is interpreted as fossilized blubber, a hallmark of warm-blooded marine amniotes.

  • Pterosaurs and feathers

 

Type 3 filaments (arrows) and similar structures (triangles). Scale bars: 10 mm in a, c and d; 1 mm in b. From Yang et al., 2018

Feathers were once considered to be unique avialan structures. Recent studies indicated that non avian dinosaurs, as part of Archosauria, possessed the entirety of the known non keratin protein-coding toolkit for making feathers. Primitive theropods, such as Sinosauropteryx and the tyrannosaurs Dilong and Yutyrannus, and some plant-eating ornithischian dinosaurs, such as Tianyulong and Kulindadromeus, are known from their spectacularly preserved fossils covered in simple, hair-like filaments called ‘protofeathers’. Other integumentary filaments, termed pycnofibres, has been reported in several pterosaur specimens, but there is still a substantial disagreement regarding their interpretation. J. Yang and colleagues described two specimens of short-tailed pterosaurs (NJU–57003 and CAGS–Z070) from the Middle-Late Jurassic Yanliao Biota, in northeast China (around 165-160 million years ago) with preserved structural fibres (actinofibrils) and four different types of pycnofibres. The specimens resemble Jeholopterus and Dendrorhynchoides, but they are relatively small. Pterosaurs were winged cousins of the dinosaurs and lived from around 200 million years ago to 66 million years ago. In the early 1800’s, a fuzzy integument was first reported from the holotype of Scaphognathus crassirostris. A recent study on this specimen shows a subset of pycnofibers and actinofibrils. The discovery of integumentary structures in other pterosaurs, such as Pterorhynchus wellnhoferi(another rhamphorhynchoid pterosaur), and these exquisitely preserved pterosaurs from China, suggest that all Avemetatarsalia (the wide clade that includes dinosaurs, pterosaurs and close relatives) were ancestrally feathered.

References:

Rauhut OWM, Foth C, Tischlinger H. (2018The oldest Archaeopteryx (Theropoda: Avialiae): a new specimen from the Kimmeridgian/Tithonian boundary of Schamhaupten, BavariaPeerJ 6:e4191 https://doi.org/10.7717/peerj.4191

Porfiri, J.D., Juárez Valieri, Rubé.D., Santos, D.D.D., Lamanna, M.C., A new megaraptoran theropod dinosaur from the Upper Cretaceous Bajo de la Carpa Formation of northwestern Patagonia, Cretaceous Research (2018), doi: 10.1016/j.cretres.2018.03.014.

Xing Xu, Paul Upchurch, Philip D. Mannion, Paul M. Barrett, Omar R. Regalado-Fernandez, Jinyou Mo, Jinfu Ma and Hongan Liu. 2018. A New Middle Jurassic Diplodocoid Suggests An Earlier Dispersal and Diversification of Sauropod Dinosaurs. Nature Communications.9, 2700.  DOI:  10.1038/s41467-018-05128-1 

Cecilia Apaldetti, Ricardo N. Martínez, Ignacio A. Cerda, Diego Pol and Oscar Alcober (2018). An early trend towards gigantism in Triassic sauropodomorph dinosaurs. Nature Ecology & Evolution. https://doi.org/10.1038/s41559-018-0599-y

Brooks B. Britt et al. Caelestiventus hanseni gen. et sp. nov. extends the desert-dwelling pterosaur record back 65 million years, Nature Ecology & Evolution (2018). DOI: 10.1038/s41559-018-0627-y

Müller RT, Langer MC, Dias-da-Silva S. 2018, An exceptionally preserved association of complete dinosaur skeletons reveals the oldest long-necked sauropodomorphs. Biol. Lett. 14: 20180633. http://dx.doi.org/10.1098/rsbl.2018.0633

Lindgren, J., Sjövall, P., Thiel, V., Zheng, W., Ito, S., Wakamatsu, K., … Schweitzer, M. H. (2018). Soft-tissue evidence for homeothermy and crypsis in a Jurassic ichthyosaur. Nature. doi:10.1038/s41586-018-0775-x

Yang Z. et al., 2018. Pterosaur integumentary structure with complex feather-like branching. Nature Ecology and Evolution https://doi.org/10.1038/s41559-018-0728-7

Ingentia prima, the first giant

Skeletal anatomy of Ingentia prima (From Apaldetti et al., 2018)

During the Late Triassic period numerous extinctions, diversifications and faunal radiations changed the ecosystem dynamics throughout the world. Followed the extinction of rhynchosaurs in most, or all, parts of the world, there was a burst of dinosaurian diversity in the late Carnian, represented by the upper Ischigualasto Formation and coeval units, with mostly carnivorous small- to medium-sized dinosaurs. Then, the long span of the early Norian, from 228.5–218 Ma, during which dicynodonts and sauropodomorph dinosaurs were the major herbivores.

Sauropods evolved from small, gracile, bipedal forms, and it was long thought that acquisition of giant body size in this clade occurred during the Jurassic and was linked to several skeletal modifications. Ingentia prima — literally the “first giant” in Latin — from the Late Triassic of Argentina shed new lights on the origin of gigantism in this group. The holotype, PVSJ 1086, composed of six articulated posterior cervical vertebrae, glenoid region of right scapula and right forelimb lacking all phalanges, has been recovered from the southern outcrops of the Quebrada del Barro Formation, northwestern Argentina. Discovered in 2015 by Diego Abelín and a team led by Cecilia Apaldetti of CONICET-Universidad Nacional de San Juan, Argentina, this new fossil weighed up to 11 tons and measured up to 32 feet (10 meters) long.

Bones of Ingentia prima (Image credit: Cecilia Apaldetti, CONICET-Universidad Nacional de San Juan, Argentina)

Ingentia was unearthed with three new specimens of Lessemsaurus sauropoides. The four dinosaurs belongs to the clade Lessemsauridae, that differs from all other Sauropodomorpha dinosaurs in possessing robust scapulae with dorsal and ventral ends equally expanded; slit-shaped neural canal of posterior dorsal vertebrae; anterior dorsal neural spines transversely expanded towards the dorsal end; a minimum transverse shaft width of the first metacarpal greater than twice the minimum transverse shaft of the second metacarpal; and bone growth characterized by the presence of thick zones of highly vascularized fibrolamellar bone, within a cyclical growth pattern.

The age of the oldest lessemsaurid (mid-Norian) indicates the appearance of an early trend towards large body size at least 15 Myr earlier than previously thought. For a long time, gigantism in eusauropods has been proposed as the result of a complex interplay of anatomical, physiological and reproductive intrinsic traits. For example, the upright position of the limbs has been highlighted as a major feature of the sauropodomorph bauplan considered an adaptation to gigantism. However Lessemsaurids lacked the purported adaptations related to a fully erect forelimb and the marked modifications of the hindlimb lever arms in eusauropods, showing that these features were not strictly necessary for the acquisition of gigantic body size. Another feature interpreted as a key acquisition was the elongated neck. However, lessemsaurids also lacked an elongated neck as they had proportionately short cervical vertebrae, indicating that the neck elongation was not a prerequisite for achieving body sizes comparable to those of basal eusauropods or gravisaurians.

 

References:

Cecilia Apaldetti, Ricardo N. Martínez, Ignacio A. Cerda, Diego Pol and Oscar Alcober (2018). An early trend towards gigantism in Triassic sauropodomorph dinosaurs. Nature Ecology & Evolution. https://doi.org/10.1038/s41559-018-0599-y

New tetrapod assemblage from the Chañares Formation

Skeletal anatomy of the erpetosuchid pseudosuchian Tarjadia ruthae. From Ezcurra et al., 2017

In the aftermath of the Permo-Triassic mass extinction (~252 Ma), several typical Palaeozoic synapsids and parareptiles were replaced by stem and crown archosaurs (archosauromorphs) and eucynodonts, and the Late Triassic fossil record of South America has been crucial to shed light on their evolutionary histories.

The Chañares Formation is part of the Ischigualasto-Villa Unión Basin, and represents one of the most continuous continental Triassic succesions in South America. Located in Talampaya National Park (La Rioja Province), the Chañares Formation is characterized at its base by a sandstone–siltstone fluvial facies with distinct lower and upper levels. The lower levels are composed of light olive grey fine-grained sandstones with abundant small brown carbonate concretions. The upper levels include fine-grained sandstones and siltstones that yielded a rich tetrapod assemblage composed of kannemeyeriiform dicynodonts, traversodontid and probainognathian cynodonts, proterochampsid stem-archosaurs, stem-crocodylians, and dinosaur precursors.

Volcanism played an important role in the generation and preservation of the Chañares Formation’s exceptional tetrapod fossil record. Recent radioisotopic datings temporally constrained most of the lower half of this unit to the earliest Carnian (236–231 Ma), showing that this assemblage preceded the oldest members of typical Late Triassic archosaur clades that are found in the Ischigualasto Formation. The new assemblage is called here as the Tarjadia Assemblage Zone, while the upper, historically known assemblage is called the Massetognathus–Chanaresuchus Assemblage Zone. This new assemblage sheds light on the link between the Early–Middle Triassic tetrapod assemblages of Africa (for example, Karoo, Ruhuhu and Otiwarongo basins) and those from the Middle–Late Triassic of South America.

The Chañares Formation (© 2012 Idean)

Tarjadia ruthae is characterized by a dorsoventrally thick skull roof ornamented by deep pits and grooves of random arrangement; Y-shaped tuberosity on the dorsal surface of the anterior end of the parietals; marginal dentition with serrations; spine table of the presacral and anterior caudal vertebrae with a transversely concave dorsal surface; a femur with a poorly developed fourth trochanter and a hook-shaped tibial condyle; and thick dorsal osteoderms with a coarse pitted ornamentation. The abundance of the erpetosuchid Tarjadia in the lowermost levels of the Chañares Formation indicates that this pseudosuchian was an important secondary consumer in its ecosystem

The Tarjadia and Massetognathus–Chanaresuchusassemblage zones currently do not share species or low level taxa, indicating a profound faunal replacement involving both primary and secondary consumers. Therefore, the rise of dinosaurs and other archosauromorph clades that diversified worldwide in the Late Triassic was preceded by a phase of relatively rapid changing ecosystems in southwestern Pangaea, including two (Tarjadia and Massetognathus–Chanaresuchus assemblage zones) profound faunal replacements in a time span shorter than 6 Myr (around 236–231 Ma).

References:

Martín D. Ezcurra, Lucas E. Fiorelli, Agustín G. Martinelli, Sebastián Rocher, M. Belén von Baczko, Miguel Ezpeleta, Jeremías R. A. Taborda, E. Martín Hechenleitner, M. Jimena Trotteyn & Julia B. Desojo; Deep faunistic turnovers preceded the rise of dinosaurs in southwestern Pangaea, Nature Ecology & Evolution (2017) doi:10.1038/s41559-017-0305-5

Benton, M. J., Tverdokhlebov, V. P. & Surkov, M. V. Ecosystem remodelling among vertebrates at the Permian–Triassic boundary in Russia. Nature 432, 97–100 (2004).

Patagotitan and the problem of body mass estimation

Image: A. Otero.

Since the discovery of dinosaur remains in the Neuquen basin in 1882, Argentina has gained the title of Land of the Giants. The tittle was reinforced by recent discoveries of more remains of giant titanosaurs like Argentinosaurus, Dreadnoughtus, Notocolossus, Puertasaurus.

Titanosaurus were a diverse group of sauropod dinosaurs represented by more than 30 genera, which included all descendants of the more recent common ancestor of Andesaurus and Saltasaurus. The group includes the smallest (e.g. Rinconsaurus, Saltasaurus; with estimated body masses of approximately 6 tonnes) and largest sauropods known to date. They had their major radiation during the middle Early Cretaceous. The evolution of body mass in this clade is key element to understand sauropod evolution.

Patagotitan reconstruction (Image: Diego Pol)

Patagotitan mayorum, originally discovered in 2010 by the rural farmer Aurelio Hernandez  is the largest and the most complete titanosaur taxa recovered to date. The generic name Patagotitan is derived from Patago (in reference to the geographic origin of the fossils, Patagonia), and titan (symbolic of its large size). The species name honours the Mayo family (owner of La Flecha Farm, the place where the fossils were found). The holotype (MPEF-PV 3400), includes an anterior and two middle cervical vertebrae, three anterior, two middle and two posterior dorsal vertebrae, six anterior caudal vertebrae, three chevrons, dorsal ribs, both sternal plates, right scapulocoracoid, both pubes and both femora. Six individuals were found in the same quarry, distributed in three distinct but closely spaced horizons, corresponding to  three different burial events. The first estimations of Patagotitan body mass suggest that it would weigh around 70 tons. The dorsal vertebrae preserved in Patagotitan, Argentinosaurus and Puertasaurus allows distinguishing the new taxon from previously known giant titanosaurs from the ‘mid-Cretaceous’ of Patagonia.

(a) Middle cervical vertebra in right lateral view; (b) anterior dorsal vertebra in anterior view (From Carballido et al., 2017)

During the last decades Argentinosaurus hiunculensis has been considered the largest dinosaur that ever walked the Earth. But because of the fragmentary nature of the type specimen, quantitative methods for body mass estimation cannot be directly applied. Two previous studies (Mazzetta et al., 2004; Benson et al., 2014) estimated the body mass of Argentinosaurus by applying scaling equations and measurements taken from two isolated femoral shafts found in deposits of the Huincul Formation. Calculations based in one of these fragmentary femora, housed at the Museo de La Plata collection and at the Museo Municipal “Carmen Funes”, estimates a body mass of 73 tons, but for the moment none of the femora can be confidentially referred to Argentinosaurus given the complete absence of femoral remains in the type material.

The team lead by Dr. José Luis Carballido from the Egidio Feruglio Paleontology Museum (Mef), used the anterior dorsal vertebrae (preserved in Argentinosaurus, Puertasaurus, Notocolossus) for a size comparison between Patagotitan and other giant titanosaurs from Patagonia. The direct comparison of these elements indicate that the dorsal vertebrae of Patagotitan are 8%–18% larger than that of Argentinosaurus and Puertasaurus, and even larger when compared to Notocolossus. Unfortunatelly, as the team remarks, this cannot be extrapolate to determine the body mass for Argentinosaurus and Puertasaurus and the only way to obtain a reliable body mass estimation is contingent on finding new associated material that can be referred to these taxa.

 

References:

Carballido JL, Pol D, Otero A, Cerda IA, Salgado L, Garrido AC, Ramezani J, Cúneo NR, Krause JM. 2017 A new giant titanosaur sheds light on body mass evolution among sauropod dinosaurs. Proc. R. Soc. B 284: 20171219.
DOI: 10.1098/rspb.2017.1219

Mazzetta, G. V., Christiansen, P., & Fariña, R. a. (2004). Giants and Bizarres: Body Size of Some Southern South American Cretaceous Dinosaurs. Historical Biology: A Journal of Paleobiology, 16(2–4), 71–83. http://dx.doi.org/10.1080/08912960410001715132

Benson, R. B. J., Campione, N. E., Carrano, M. T., Mannion, P. D., Sullivan, C., Upchurch, P., & Evans, D. C. (2014). Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage. PLoS Biology, 12(5), http://doi.org/10.1371/journal.pbio.1001853.

 

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)