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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Introducing Moros intrepidus, the harbinger of doom.

Moros intrepidus. Credit: Jorge Gonzalez

Tyrannosauroidea, the superfamily of carnivorous dinosaurs that includes the iconic Tyrannosaurus rex, originated in the Middle Jurassic, approximately 165 million years ago, and was a dominant component of the dinosaur faunas of the Northern Hemisphere. All tyrannosaurs were bipedal predators characterized by premaxillary teeth with a D-shaped cross section, fused nasals, extreme pneumaticity in the skull roof and lower jaws, a pronounced muscle attachment ridge on the ilium, and an elevated femoral head. But for most of their evolutionary history, tyrannosauroids were mostly small-bodied animals and only reached gigantic size during the final 20 million years of the Cretaceous. Now, the discovery of a new, diminutive tyrannosauroid, Moros intrepidus gen. et sp. nov., shed lights on the successful radiation of Campanian tyrannosauroids.

The holotype (NCSM 33392), preserves a partial right hind limb including portions of the femur, tibia, second and fourth metatarsals, and phalanges of the fourth pedal digit. It was recovered from the lower Mussentuchit Member (6–7 m above the Ruby Ranch contact), upper Cedar Mountain Formation, Emery County, Utah, USA. This small-bodied, gracile-limbed tyrannosauroid lived about 96 million years ago. The name derived from Greek word Moros (an embodiment of impending doom) in reference to the establishment of the Cretaceous tyrannosauroid lineage in NA, and the Latin word intrepidus (intrepid), in reference to the hypothesized intracontinental dispersal of tyrannosaurs during this interval.

Bone microstructure of M. intrepidus (NCSM 33392). From Zanno et al., 2019.

NCSM 33392 derives from a skeletally immature individual (6-7 years) nearing adult size . According to the histological analysis, M. intrepidus exhibits a moderate growth rate, similar to Guanlong, a more primitive tyrannosauroid from the Late Jurassic of China. By contrast, large-bodied, tyrannosaurines from the last stages of the Cretaceous, like Gorgosaurus, were already triple their masses at similar ages. M. intrepidus suggests that North American tyrannosauroids were restricted to small sizes for a protracted period of ~15 million years and at some point at the Turonian, they embarked on a trend of rapid body size increases, to became the top predators of the Cretaceous.

 

References:

Zanno, L.E, Tucker, R.T., Canoville, A., Avrahami, H.M., Gates, T.A., Makovicky, P.J. (2019), Diminutive fleet-footed tyrannosauroid narrows the 70-million-year gap in the North American fossil record, Communications Biology, DOI: 10.1038/s42003-019-0308-7

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

Introducing Macrocollum itaquii.

M. itaquii, the oldest long-necked dino ever found, dating back 225 million years. (Credit: Müller et al 2018)

Sauropodomorphs were the largest land animals ever recorded in the history of life. Additionally to their colossal size, the sauropodomorph bauplan is also characterised by a small head, long neck, barrel-shaped body and columnar limbs. The group was successful and diverse, achieving a worldwide geographical distribution. Nevertheless, the rise of sauropodomorphs is still poorly understood due to the scarcity of well-preserved fossils in early Norian rocks. The Wachholz site (Caturrita Formation), in southern Brazil, is an important window to early Norian land ecosystems. This unit has yielded several sauropodomorphs, including Unaysaurus tolentinoi and the recently described Macrocollum itaquii, the oldest long-necked dinosaur known, that shed light on the rise of the group.

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.

 

Skull of Macrocollum itaquii (From Müller et al 2018)

The generic name combines the Greek word macro (long) and the Latin word collum (neck), referring to the animal’s elongated neck. The specific epithet honours José Jerundino Machado Itaqui, one of the main actors behind the creation of CAPPA/UFSM (Centro de Apoio à Pesquisa Paleontológica da Quarta Colônia/Universidade Federal de Santa Maria).

M. itaquii was only 3.5 meters long and weighed about 101.6 kilograms, and differs from all other known sauropodomorphs in possessing the following characters: antorbital fossa perforated by a promaxillary fenestra; medial margin of the supratemporal fossa with a simple smooth curve at the frontal/parietal suture; proximal articular surface of metacarpal I transversely narrow; acetabulum not fully open; ischiadic longitudinal groove not reaching the caudal half of the ischium; absence of trochanteric shelf on the femur; medial condyle of distal femoral articulation subrectangular in distal view; proximal end of metatarsal II with a straight medial margin.

An artist’s impression of M. itaquii.

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.

 

References:

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

 

 

Introducing Dynamoterror dynastes, the powerful terror ruler.

Frontals of Dynamoterror dynastes in rostral view. From McDonald et al., 2018. (Scale bars = 5 cm)

Tyrannosauroidea is a relatively derived group of theropod dinosaurs more closely related to birds than to other large theropods such as allosauroids and spinosaurids. The clade originated in the Middle Jurassic, approximately 165 million years ago, and for most of their evolutionary history, tyrannosauroids were mostly small-bodied animals that only reached gigantic size during the final 20 million years of the Cretaceous. Until recently, all tyrannosaurs fossils were limited to Asia and North America, but the latest discoveries suggest a more cosmopolitan distribution during their early evolution.

All tyrannosaurs were bipedal predators characterized by premaxillary teeth with a D-shaped cross section, fused nasals, extreme pneumaticity in the skull roof and lower jaws, a pronounced muscle attachment ridge on the ilium, and an elevated femoral head. The clade was a dominant component of the dinosaur faunas of the American West shortly after the emplacement of the Western Interior Seaway (about 99.5 Mya).

Paleogeography of North America during the late Campanian Stage of the Late Cretaceous (∼75 Ma). From Sampson et al., 2010

Dynamoterror dynastes, the most recent taxon described from the lower Campanian of northwestern New Mexico, provides additional data on the morphology and diversity of early tyrannosaurines in Laramidia. The new specimen lived during the Late Cretaceous period, approximately 78 million years ago. The name derived from Greek word dynamis (“power”) and the Latin word terror. The specific name is a Latin word meaning “ruler. Dynamoterror was collected in San Juan County, New Mexico, and is the first associated tyrannosaurid skeleton reported from the Menefee Formation.

The holotype (UMNH VP 28348) is an incomplete associated skeleton including the left and right frontals, four fragmentary vertebral centra, fragments of dorsal ribs, right metacarpal II, supraacetabular crest of the right ilium, unidentifiable fragments of long bones, phalanx 2 of left pedal digit IV, and phalanx 4 of left pedal digit IV. The right and left frontals both are incomplete; the dimensions of the right frontal are similar to a subadult specimen of Tyrannosaurus rex, suggesting that UMNH VP 28348 represents a subadult or adult individual. The reconstructed skull roof of Dynamoterror present several tyrannosaurine features, such as large supratemporal fossae and a tall sagittal crest on the frontals, providing an expanded attachment area for enormous jaw-closing muscles.

 

References:

McDonald AT, Wolfe DG, Dooley AC Jr. (2018) A new tyrannosaurid (Dinosauria: Theropoda) from the Upper Cretaceous Menefee Formation of New Mexico. PeerJ 6:e5749 https://doi.org/10.7717/peerj.5749

Brusatte SL, Norell MA, Carr TD, Erickson GM, Hutchinson JR, et al. (2010) Tyrannosaur paleobiology: new research on ancient exemplar organisms. Science 329: 1481–1485. doi: 10.1126/science.1193304

Sampson SD, Loewen MA, Farke AA, Roberts EM, Forster CA, Smith JA, et al. (2010) New Horned Dinosaurs from Utah Provide Evidence for Intracontinental Dinosaur Endemism. PLoS ONE 5(9): e12292. https://doi.org/10.1371/journal.pone.0012292

 

 

 

 

 

 

 

 

 

 

 

 

The Tyrannosauroids from the Southern Hemisphere.

Santanaraptor lived in South America during the Early Cretaceous about 112 million years ago (From Wikimedia Commons).

Tyrannosauroidea, the superfamily of carnivorous dinosaurs that includes the iconic Tyrannosaurus rex, was mainly distributed in the Northern Hemisphere. However, a few specimens from Australia (Timimus hermani and the articulated pubes NMV P186046) and Brazil (Santanaraptor placidus), have been referred to this clade.

Santanaraptor was unearthed in 1996 in the Romualdo Group (Santana Formation). The holotype is a juvenile partial skeleton that may have reached 1.25 metres (4.1 ft) in length, and it was presumed to be similar to Dilong and Guanlong. It was first described as a coelurosaurian theropod by Alexander Kellner in 1999, but in 2014 Thomas Holtz classified Santanaraptor as the first tyrannosauroid known from Gondwana. Delcourt and Grillo (2018), also placed Santanaraptor within Tyrannosauroidea based on the following features: the absence of an accessory ridge on the lateral surface of the cnemial crest; the absence of a horizontal groove across the astragalar condyles anteriorly; a deep fossa on the medial surface of the femoral head, lateral to the trochanteric fossa; an ischial medial apron positioned along the anterior margin of its shaft in medial view; the lesser trochanter and the greater trochanter extending to approximately the same level proximally; the proximal margin of the femur is concave in posterior view due to a greater trochanter that is elevated substantially relative to the lateral portion of the proximal surface of the head; and a shallow femoral extensor groove on the anterior surface of the distal end that is expressed as a broad concave anterior margin in distal view but present as an extensive depression on the anterior surface of the femur.

Holotypic left femur of Timimus hermani (From Wikimedia Commons)

Timimus was unearthed in 1994 from Eumeralla Formation and shares similar features with Tyrannosauroidea, but due to the incompleteness of the Timimus holotype, is difficult to properly evaluate its phylogenetic position. The same applies to NMV P186046. It was suggested (Benson et al., 2012) that NMV P186046 and the Timimus holotype may represent a single taxon given their similar phylogenetic positions and congruent sizes, although they did not come from the same site.

Time-calibrated phylogeny of Tyrannosauroidea. From Delcourt and Grillo, 2018.

Tyrannosauroidea had a Eurasian distribution, but basal lineages of the newly proposed clade, Pantyrannosauria (the most inclusive clade, containing Tyrannosaurus rex and Dilong paradoxus, but not Proceratosaurus bradleyi), were distributed across Europe (Juratyrant, Eotyrannus and Aviatyrannis), North America (Stokesosaurus), South America (Santanaraptor), Australia (Timimus), and Asia (Dilong and Xiongguanlong). It was hypothesized that all basal lineages of Pantyrannosauria were already established in the Late Jurassic before the complete separation of Gondwana and Laurasia.

The fact that Santanaraptor and Timimus were relatively small suggests that Gondwanan tyrannosauroids remained small in comparison to northern species. The presence of Santanaraptor in a semi-arid environment with marine incursion also suggests that tyrannosauroids were not only found in humid paleoenvironments.

References:

Rafael Delcourt, Orlando Nelson Grillo , Tyrannosauroids from the Southern Hemisphere: Implications for biogeography, evolution, and taxonomy. Palaeo (2018), doi: 10.1016/j.palaeo.2018.09.003

Apesteguía, S., Smith, N.D., Juárez Valieri, R., Makovicky, P.J., 2016. An Unusual New Theropod with a Didactyl Manus from the Upper Cretaceous of Patagonia, Argentina. PLoS One 11, 1–41. doi:10.1371/journal.pone.0157793

Benson, R.B.J., Rich, T.H., Vickers-Rich, P., Hall, M., 2012. Theropod fauna from southern Australia indicates high polar diversity and climate-driven dinosaur provinciality. PLoS One 7, e37122. doi:10.1371/journal.pone.0037122

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. https://doi.org/10.1016/j.cretres.2014.04.007

Holtz Jr, T.R., 2004. Tyrannosauroidea, in: Weishampel, D.B., Dodson, P., Osm (Eds.), The Dinosauria. University of California Press, Berkeley, pp. 111–136.

A very short history of Dinosaurs.

Evolutionary relationships of dinosaurs. From Benton 2018.

On 20 February 1824, William Buckland published the first report of a large carnivore animal: the Megalosaurus. The description was based on specimens in the Ashmolean Museum, in the collection of Gideon Algernon Mantell of Lewes in Sussex, and a sacrum donated by Henry Warburton (1784–1858). One year later, the Iguanodon entered in the books of History followed by the description of Hylaeosaurus in 1833. After examined the anatomy of these three genera, Richard Owen erected the clade Dinosauria in 1842.

Dinosaurs likely originated in the Early to Middle Triassic. The closest evolutionary relatives of dinosaurs include flying pterosaurs and herbivorous silesaurids. Early ecological divergences in dinosaur evolution are signaled by disparity in dental morphology, which indicates carnivory in early theropods, herbivory in ornithischians, and omnivory in sauropodomorph (subsequently sauropodomorphs underwent a transition to herbivory).

Eoraptor lunensis, outcropping from the soil. Valle de la Luna (Moon Valley), Parque Provincial Ischigualasto, Provincia de San Juan, Argentina.

The oldest dinosaurs remains are from the late Carnian (230 Ma) of the lower Ischigualasto Formation in northwestern Argentina. Similarly, the Santa Maria and Caturrita formations in southern Brazil preserve basal dinosauromorphs, basal saurischians, and early sauropodomorphs. In North America, the oldest dated occurrences of vertebrate assemblages with dinosaurs are from the Chinle Formation. Two further early dinosaur-bearing formations, are the lower (and upper) Maleri Formation of India and the Pebbly Arkose Formation of Zimbabwe. These skeletal records of early dinosaurs document a time when they were not numerically abundant, and they were still of modest size.

During the Late Triassic period numerous extinctions, diversifications and faunal radiations changed the ecosystems dynamics throughout the world. Nevertheless, dinosaurs exhibited high rates of survival. According to the competitive model, the success of dinosaurs was explained in terms of their upright posture, predatory skills, or warm-bloodedness. In the opportunistic model, dinosaurs emerged in the late Carnian or early Norian, and then diversified explosively. The current model contains some aspects of both the classic competition model and the opportunistic model. In this model, the crurotarsan-dominated faunas were replaced by a gradual process probably accelerated by the ecological perturbation of the CPE (Carnian Pluvial Episode).

Ingentia prima outcropping from the soil.

In the Jurassic and Cretaceous dinosaurs achieved enormous disparity. Sauropodomorphs achieved a worldwide distribution and become more graviportal and increased their body size. Gigantism in this group 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, the discovery of Ingentia prima, from the Late Triassic of Argentina, indicates that this feature was not strictly necessary for the acquisition of gigantic body size.

Ornithischian were primitively bipedal, but reverted to quadrupedality on at least three occasions: in Ceratopsia, Thyreophora and Hadrosauriformes. The presence of early armored dinosaurs (thyreophorans) in North America, Asia, and Europe, but their absent from the southern African record, suggests some degree of provinciality in early ornithischian faunas.

Archaeopteryx lithographica, specimen displayed at the Museum für Naturkunde in Berlin. (From Wikimedia Commons)

Theropod dinosaurs also increased their diversity and exhibit a greater range of morphological disparity. The group underwent multiple parallel increases in brain size. The volumetric expansion of the avian endocranium began relatively early in theropod evolution. For instance, the endocranium of Archaeopteryx lithographica is volumetrically intermediate between those of more basal theropods and crown birds. The digital brain cast of Archaeopteryx also present an indentation that could be from the wulst, a neurological structure present in living birds used in information processing and motor control with two primary inputs: somatosensory and visual. The extensive skeletal pneumaticity in theropods such as Majungasaurus demonstrates that a complex air-sac system and birdlike respiration evolved in birds’ theropod ancestors. Anatomical features like aspects of egg shape, ornamentation, microstructure, and porosity of living birds trace their origin to the maniraptoran theropods, such as oviraptorosaurs and troodontids. In addition, some preserving brooding postures, are known for four oviraptorosaurs, two troodontids, a dromaeosaur, and one basal bird providing clear evidence for parental care of eggs.

Nonavian dinosaurs disappeared more or less abruptly at the end of the Cretaceous (66 mya). Birds, the only living dinosaurs, with more than 10,500 living species are the most species-rich class of tetrapod vertebrates.

 

References:

Benson, R. B. J. (2018). Dinosaur Macroevolution and Macroecology. Annual Review of Ecology, Evolution, and Systematics, 49(1).  doi:10.1146/annurev-ecolsys-110617-062231

Michael J. Benton et al. The Carnian Pluvial Episode and the origin of dinosaurs, Journal of the Geological Society (2018). DOI: 10.1144/jgs2018-049

Xing Xu, Zhonghe Zhou, Robert Dudley, Susan Mackem, Cheng-Ming Chuong, Gregory M. Erickson, David J. Varricchio, An integrative approach to understanding bird origins, Science, Vol. 346 no. 6215, DOI: 10.1126/science.1253293.

 

Lingwulong shenqi, the “Amazing Dragon”, and the dispersal of Sauropods.

Skeletal reconstruction and exemplar skeletal remains of Lingwulong shenqi. Scale bars = 100 cm for a and 5 cm for b–o. From Xu 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. However, a newly discovered dinosaur from the Middle Jurassic of northern China suggests that Sauropods dispersed and diversified earlier than previously thought.

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.

An artist’s interpretation of what Lingwulong shenqi (Image: Zhang Zongda)

The Lingwulong specimens were found in the Yanan Formation at Ciyaopu, in northwest China. This formation has been divided in four or five members. Although, no radiometric constraints have been obtained for the Yanan Formation, its age has been estimated on the basis of biostratigraphy. The presence of a conchostracans assemblage (including Palaeoleptoestheria, Triglypta, and Euestheria) is indicative of a Middle Jurassic age.

The East Asian Isolation Hypothesis (EAIH) has become a well-established explanation of profound differences between Jurassic (and sometimes Early Cretaceous) Asian terrestrial faunas, that resulted in the evolution of endemic groups such as mamenchisaurid sauropods, and the early diverging lineage of tetanurans, oviraptorosaurs, therizinosaurs. In this model, the isolation ended in the Early Cretaceous when marine regressions allowed the invasion of groups from elsewhere in Pangaea, and the dispersal of Asian endemics (e.g., oviraptorosaurs, marginocephalians) into Europe and North America. However, it was claimed that diplodocoids never took part in these dispersals because the end-Jurassic extinction that greatly reduced their diversity and geographic range in the Early Cretaceous. The discovery of Lingwulong undermines the EAIH, forcing a significant revision of hypotheses concerning the origins and early radiation of Neosauropoda.

 

References:

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 

 

 

 

Introducing Akainacephalus johnsoni

Skeletal reconstructions of Akainacephalus johnsoni. From Wiersma and Irmis, 2018

The Ankylosauria is a group of herbivorous, quadrupedal, armoured dinosaurs subdivided in two major clades, the Ankylosauridae and the Nodosauridae. The group is predominantly recorded from the Late Cretaceous (Turonian—late Maastrichtian) of Asia and the last Cretaceous (early Campanian—late Maastrichtian) of western North America (Laramidia). Ankylosauridae were present primarily in Asia and North America, and the most derived members of this clade are characterized by shortened skulls, pyramidal squamosal horns, and tail clubs.

Akainacephalus johnsoni, a new genus and species of an ankylosaurid dinosaur from the upper Campanian Kaiparowits Formation of southern Utah, represents the most complete ankylosaurid specimen from southern Laramidia to date, and reveals new details about the diversity and evolution of this clade. The genus name is derived from the Greek akaina, meaning “thorn” or “spine,” referring to the thorn-like cranial caputegulae of the holotype; and “cephalus,” the Greek meaning for head. The specific epithet honors Randy Johnson, volunteer preparator at the Natural History Museum of Utah.

Skull of Akainacephalus johnsoni. From Wiersma and Irmis, 2018

The holotype (UMNH VP 20202) is a partial skeleton comprising a complete skull, both mandibles, predentary, four dorsal, four dorsosacral, three sacral, one caudosacral, and eight caudal vertebrae, dorsal ribs, a complete tail club, both scapulae, left coracoid, right humerus, right ulna, partial left ilium, left femur, left tibia, left fibula, phalanx, two partial cervical osteoderm half rings, and 17 dorsal and lateral osteoderms of various sizes and morphologies.

The most striking feature of Akainacephalus johnsoni is the skull ornamentation comprising several symmetrical rows of small pyramidal and conical caputegulae along the dorsolateral surface of the skull. The postorbital horns are dorsoventrally tall, backswept, and project laterally in dorsal view. The quadratojugal horns display an  asymmetrical triangular morphology with a vertically positioned apex. Only a partial squamosal horn is preserved, but is largely broken.

Life reconstruction of Akainacephalus johnsoni (Image credit: Andrey Atuchin and the Denver Museum of Nature & Science)

The unique anatomical features of Akainacephalus johnsoni indicate a close taxonomic relationship with Nodocephalosaurus kirtlandensis, that clearly distinguish them from other Late Cretaceous Laramidian (although both taxa are temporally separated by nearly three million years). Because both taxa a more closely related to Asian ankylosaurids, the geographic distribution of Late Cretaceous ankylosaurids throughout the Western Interior could be the result of several geologically brief intervals of lowered sea level that allowed Asian ankylosaurid dinosaurs to immigrate to North America several times during the Late Cretaceous. The dispersal of ankylosaurids into Laramidia is coeval with the dispersal of other dinosaur clades, like tyrannosaurids and ceratopsians. The climate gradients and the fluctuations in sea level, may have helped reinforced Campanian provincialism.

 

References:

Wiersma JP, Irmis RB. (2018) A new southern Laramidian ankylosaurid, Akainacephalus johnsoni gen. et sp. nov., from the upper Campanian Kaiparowits Formation of southern Utah, USA. PeerJ 6:e5016 https://doi.org/10.7717/peerj.5016

Arbour, V. M.; Currie, P. J. (2015). “Systematics, phylogeny and palaeobiogeography of the ankylosaurid dinosaurs”. Journal of Systematic Palaeontology: 1–60. doi: 10.1080/14772019.2015.1059985