A brief history of the Spinosaurus.

One of the photographs donate by W. Stromer. Image from the Washington University in St. Louis

Despite its low fossil record, Spinosaurus is one of the most famous dinosaur of all time. This gigantic theropod possessed highly derived cranial and vertebral features sufficiently distinct for it to be designated as the nominal genus of the clade Spinosauridae. In 1910, E. Stromer went to his third paleontological expedition to Egypt. He arrived to Alexandria on November 7. He was initially looking for early mammals and planned visit the area of Bahariya, in the Western Desert, which has sediments from the Cretaceous era. But an expedition to the Western Desert needed the permission by the English and French colonial authorities and of course the Egyptian authorities. Although diplomatic relations with Germany were rapidly deteriorating, Stromer managed to get the permissions. He arrived to the Bahariya Oasis on January 11, 1911. After facing some difficulties during the journey, on January 17 he began to explore the area of Gebel el Dist, and at the bottom of the Bahariya Depression, Stromer found  the remains of four immense and entirely new dinosaurs (Aegyptosaurus, Bahariasaurus, Carcharodontosaurus and Spinosaurus aegyptiacus), along with dozens of other unique specimens. Stromer and Markgraf recovered the right and left dentaries and splenials from the lower jaw; a straight piece of the left maxilla that was described but not drawn; 20 teeth; 2 cervical vertebrae; 7 dorsal (trunk) vertebrae; 3 sacral vertebrae; 1 caudal vertebra; 4 thoracic ribs; and gastralia. This gigantic predator is estimated to have been about 14 m, with unusually long spines on its back that probably formed a large, sail-like structure.

1) Photograph of the right mandibular ramus of the holotype of Spinosaurus aegyptiacus Stromer, 1915 (BSP 1912 VIII 19), in lateral view. 2) Reproduction of Stromer’s (1915, pl. I, fig. 12a) illustration of the right mandibular ramus.

Due to political tensions before and after World War I, many of this fossils were damaged after being inspected by colonial authorities and not arrived to Munich until 1922. The shipping from El Cairo was paid by the Swiss paleontologist Bernhard Peyer (1885-1963), a former student and friend of Stromer. During the World War II, E. Stromer tried to convince Karl Beurlen -a young nazi paleontologist who was in charge of the collection- that he had to move the fossils to a safer place, but Beurlen refused to do it. Unfortunately, on April 24, 1944, a British Royal Air Force raid bombed the museum and incinerated its collections. Only two photographs of the holotype of Spinosaurus aegyptiacus were recovered in in the archives of the Paläontologische Museum in June 2000, after they were donated to the museum by Ernst Stromer’s son, Wolfgang Stromer, in 1995. These photographs provide additional insight into the anatomy of the holotype specimen of Spinosaurus aegyptiacus.

“Illustrations of the vertebrate “sail” bones of Spinosaurus that appeared in one of Stromer’s monographs. From Wikimedia Commons.

In his original monograph, Stromer emphasized the peculiar character of the teeth of this unusual theropod. Because of their morphological convergence with those of crocodilians and other fish-eating reptiles, isolated spinosaurid teeth have frequently been misinterpreted. It appears that Baryonyx-like teeth were collected by Gideon Mantell in Sussex around 1820. Georges Cuvier was the first to publish an illustration of the four teeth from Tilgate Forest. These teeth, however, were generally considered as belonging to crocodilians, and when Richard Owen erected the taxon Suchosaurus cultridens to designate them he placed it among the crocodiles. Even when Owen realized that these teeth were peculiar in many respects and hinted at possible affinities with dinosaurs, he persistently classified Suchosaurus as a crocodilian, an interpretation that was accepted by most subsequent authors.

Although Stromer’s original description of Spinosaurus aegyptiacus was published in 1915, a more complete detailed picture of its anatomy, evolution, and biogeography only begun to emerge in recent decades.



HONE, D. W. E. and HOLTZ, T. R. (2017), A Century of Spinosaurs – A Review and Revision of the Spinosauridae with Comments on Their Ecology. Acta Geologica Sinica, 91: 1120–1132. doi: 10.1111/1755-6724.13328



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).


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).

A Brief Introduction to the Osteology of Viavenator exxoni

Viavenator exxoni, Museo Municipal Argentino Urquiza

The Abelisauridae is the best-known carnivorous dinosaur group from Gondwana. Their fossil remains have been recovered in Argentina, Brazil, Morocco, Niger, Libya, Madagascar, India, and France. These theropods exhibit spectacular cranial ornamentation in the form of horns and spikes and strongly reduced forelimbs and hands. The group was erected by Jose Bonaparte with the description of  Abelisaurus comahuensis, and includes: Carnotaurus sastrei, 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. Viavenator series of autapomorphies are: transversely compressed parietal depressions on both sides of the supraoccipital crest; ventral edges of the paraoccipital processes located above the level of the dorsal edge of the occipital condyle; basioccipital-opisthotic complex about two and a half times the width and almost twice the height of the occipital condyle, in posterior view; well-developed crest below the occipital condyle; deeply excavated and sub-circular basisphenoidal recess; basipterygoid processes horizontally placed with respect to the cranial roof and located slightly dorsally to the basal tubera; mid and posterior cervical centra with slightly convex lateral and ventral surfaces; presence of an interspinous accessory articular system in middle and posterior dorsal vertebrae; presence of a pair of pneumatic foramina within the prespinal fossa in anterior caudal vertebrae; distal end of the scapular blade posteriorly curved.

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.

Viavenator presents highly-derived postcranial characters, and a relatively plesiomorphic skull in comparison with Carnotaurus and Aucasaurus. Cranial elements of this specimen include the complete neurocranium: frontals, parietals, sphenethmoids, orbitosphenoids, laterosphenoids, prootics, opisthotics, supraoccipital, exoccipitals, basioccipital, parasphenoids and basisphenoids. The plesiomorphic traits of the skull of Viavenator are mainly related with the anatomy of frontals, wich lack osseous prominences such as domes or horns. The dorsal surface of the frontals exhibits an ornamentation that consists of pits and sinuous furrows and ridges, although it is not well-preserved. The  exoccipitals form the lateral and possibly the laterodorsal margins of the foramen magnum, as apparently occurs in Carnotaurus. 

Vertebrae of Viavenator exxoni. Scale bar: 5 cm. From Filippi et al., 2017),

The postcranial skeleton of Viavenator is represented by eight cervical vertebrae (the atlas; seven dorsal vertebrate, four of them articulated; twelve caudal vertebrae); ribs; gastralias; one chevron; scapulocoracoid; ischium foot; and fibulae. The atlas is similar to that of Carnotaurus, though less robust and anteroposteriorly shorter; and there  are not observed prezygapophyseal facets in the neurapophyses, so it is inferred that the proatlas was absent, as also occurs in Carnotaurus and Majungasaurus. The shape of the epipophyses of the cervical region, which are
characterized by anterior and posterior projections, is shared by Viavenator and Carnotaurus, but it is not present in pre-Santonian forms such as Ilokelesia and Skorpiovenator. The derived vertebral characters of Viavenator are linked with an increase in the structural rigidity of the vertebral column, and with an increase in the cursorial abilities of these abelisaurids. This combination of plesiomorphic and derived traits suggests that Viavenator is a transitional form.



Filippi, L.S., Méndez, A.H., Gianechini, F.A., Juárez Valieri, Rubé.D., Garrido, A.C., Osteology of Viavenator exxoni (Abelisauridae; Furileusauria) from the Bajo de la Carpa Formation, NW Patagonia, Argentina, Cretaceous Research (2017), doi: 10.1016/j.cretres.2017.07.019.

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

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


Geomythology: On Cyclops and Lestrigons

Pellegrino Tibaldi, The Blinding of Polyphemus, c. 1550-1

In Greek mythology giants are connected to the origin of the cosmos and represent the primordial chaos which contrasts with the rationality of the Gods. They were the sons of the earth (Gea) fertilized by the blood of the castrated Uranus (Heaven). In that chaotic, primal era, strange creatures proliferated, such as the Cyclopes, and the Centaurs. Lestrigons, a tribe of man-eating giants, appears in Homer’s Odyssey. Polyphemus, is one of the Cyclopes also described in Homer’s Odyssey. Greeks believed that the Laestrygonians, as well as the Cyclopes, had once inhabited Sicily.

But the ancient myth of giants is a common element in almost all cosmogonies. In Scandinavians legends, the blood of the giant Ymo formed the seas of th Earth, and his bones formed the mountains. In Peru, Brazil, and Mexico, the giants are part of the folk tradition. Judaism, more precisely, the Talmud and the Torah, converges with Genesis on the origin of the giants.

Laestrygonians Hurling Rocks at the Fleet of Odysseus

The discovery of huge fossil bones has always stimulated the imagination of local people, giving rise to legends. We found direct reference in the works of Herodotus which mentions the large bones of the giant Orestes recovered in Acadia, or even Virgil in his Georgics speaks of gigantic bones. In the sixteenth century, Italian historians, such as the Sicilian Tommaso Fazello, used the sacred texts to demonstrate that the first populations of many islands of the Mediterranean (among them Sicily and Sardinia), were of giants. At the same time, the first notices of South American fossils were reported by early Spanish explorers. These fossils were interpreted as the remains of an ancestral race of giant humans erased from the face of the Earth by a divine intervention. Fray Reginaldo de Lizarraga (1540-1609) also wrote about those “graves of giants” found in Córdoba, Argentina.

The case of Filippo Bonanni, an Italian Jesuit scholar, is very curious. He used the topic of the giants as an element in support of his theory of the inorganic origin of fossils. He properly rejects the myth of giants, but wrongly identify the nature of fossils. The most strong supporter for the organic origin of fossils was the italian painter Agostino Scilla. He published only one scientific treatise: La vana speculazione disingannata dal senso, lettera risponsiva Circa i Corpi Marini, che Petrificati si trouano in vari luoghi terrestri (The vain speculation disillusioned by the sense, response letter concerning the marine remains, which are found petrified in various terrestrial places). The aim of the work was the demonstration that fossils, which are found embedded in sediments on mountains and hills, represent the remains of lithified organisms, which at one time lived in the marine environment. The text was later translated to Latin and it was written as a response to a letter sent to him by Giovanni Francesco Buonamico, a doctor from Malta.

Femur of Mammuth interpreted as a bone of a giant and preserved as a relic in St. Stephen’s Cathedral in Vienna.

Madrisio (1718) is one of the first authors in Italy to suggest that much of this giant bones may be referred, without problem, to elephants from the past. But te real interpretative turning point takes place with the influential work of the Hans Sloane, who stressed the importance of a comparative study of the bones in various vertebrates. Applying this method, he demonstrated how the big bones and teeth found in sediments or in caves are nothing more than remains of cetaceans and large quadrupeds, remarking on the major anatomical differences between humans and other known vertebrates. Among the few precursors of Sloan, the Italian naturalist Giovanni Ciampini in 1688, using direct comparisons with the famous elephant exhibited in Florence in the Medicean Museum, was able to correctly interpret the bones found at Vitorchiano near Viterbo, initially attributed to gigantic men.


Marco Romano & Marco Avanzini (2017): The skeletons of Cyclops and Lestrigons: misinterpretation of Quaternary vertebrates as remains of the mythological giants, Historical Biology, DOI: 10.1080/08912963.2017.1342640

Dark skies at the end of the Cretaceous

A time-lapse animation showing severe cooling due to sulfate aerosols from the Chicxulub asteroid impact 66 million years ago (Credit: PKI)

Thirty years ago, the discovery of anomalously high abundance of iridium and other platinum group elements in the Cretaceous/Palaeogene (K-Pg) boundary led to the hypothesis that an asteroid collided with the Earth and caused one of the most devastating events in the history of life. The impact created the 180-kilometre wide Chicxulub crater causing widespread tsunamis along the coastal zones of the surrounding oceans and released an estimated energy equivalent of 100 teratons of TNT and produced high concentrations of dust, soot, and sulfate aerosols in the atmosphere. Three-quarters of the plant and animal species on Earth disappeared. Marine ecosystems lost about half of their species while freshwater environments shows low extinction rates, about 10% to 22% of genera.

Recent studies suggest that the amount of sunlight that reached Earth’s surface was reduced by approximately 20%. Photosynthesis stopped and the food chain collapsed. The decrease of sunlight caused a drastic short-term global reduction in temperature (15 °C on a global average, 11 °C over the ocean, and 28 °C over land). While the surface and lower atmosphere cooled, the tropopause became much warmer, eliminate the tropical cold trap and allow water vapor mixing ratios to increase to well over 1,000 ppmv in the stratosphere. Those events accelerated the destruction of the ozone layer. During this period, UV light was able to reach the surface at highly elevated and harmful levels.

Gravity anomaly map of the Chicxulub impact structure (From Wikimedia Commons)

In 1980, Walter Alvarez and his father, Luis Alvarez ignited a huge controversy when they concluded that the anomalous iridium concentration at the K-Pg boundary is best interpreted as the result of an asteroid impact. They even calculated the size of the asteroid (about 7 km in diameter) and the crater that this body might have caused (about 100–200 km across). In 1981, Pemex (a Mexican oil company) identified Chicxulub as the site of a this massive asteroid impact. The crater is more than 180 km (110 miles) in diameter and 20 km (10 miles) in depth, making the feature one of the largest confirmed impact structures on Earth.



Charles G. Bardeen, Rolando R. Garcia, Owen B. Toon, and Andrew J. Conley, On transient climate change at the Cretaceous−Paleogene boundary due to atmospheric soot injections, PNAS 2017 ; published ahead of print August 21, 2017 DOI: 10.1073/pnas.1708980114

Brugger J.G. Feulner, and S. Petri (2016), Baby, it’s cold outside: Climate model simulations of the effects of the asteroid impact at the end of the CretaceousGeophys. Res. Lett.43,  doi:10.1002/2016GL072241.

Introducing Shringasaurus indicus

Cranial anatomy of Shringasaurus indicus (From Sengupta et al., 2017)

In the aftermath of the Permo-Triassic mass extinction (~252 Ma), well diversified archosauromorph groups appear for the first time in the fossil record, including aquatic or semi aquatic forms, highly specialized herbivores, and massive predators. Allokotosaurians, meaning “strange reptiles” in Greek, comprise a bizarre suite of herbivorous archosauromorphs with a high disparity of craniodental features.

Shringasaurus indicus, from the early Middle Triassic of India, is a new representative of the Allokotosauria. The generic name is derived from ‘Śṛṅga’ (Shringa), horn (ancient Sanskrit), and ‘sauros’ (σαῦρος), lizard (ancient Greek), referring to the horned skull.  The species name ‘indicus’, refers to the country where it was discovered. The holotype ISIR (Indian Statistical Institute, Reptile, India) 780, consist of a partial skull roof (prefrontal, frontal, postfrontal, and parietal) with a pair of large supraorbital horns. The fossil bones have been collected from the Denwa Formation of the Satpura Gondwana Basin. At least seven individuals of different ontogenetic stages were excavated in the same area. Most of them were disarticulated, with exception of a partially articulated skeleton.

Skeletal anatomy of Shringasaurus indicus (From Sengupta et al., 2017)

Shringasaurus reached a relatively large size (3–4 m of total length) that distinctly exceeds the size range of other Early-Middle Triassic archosauromorphs. This new species shows convergences with sauropodomorph dinosaurs, including the shape of marginal teeth, and a relative long neck.  

Shringasaurus has a proportionally small skull with a short, rounded snout and confluent external nares. The premaxilla lacks a prenarial process and has four tooth positions. The prefrontal, nasal, frontal, and postfrontal of each side of the skull are fused to each other in large individuals. But the most striking feature of Shringasaurus indicus is the presence of a pair of large supraorbital horns, ornamented by tangential rugosities and grooves. Individuals of Shringasaurus of different ontogenetic stages indicate the size and robustness of the horns were exacerbated towards the adulthood, with a distinct variability in their orientation and anterior curvature in large individuals. Several amniotes have horns very similar to those of Shringasaurus (e.g. bovid mammals, chamaeleonid lepidosaurs). The independent evolution of similar horn shapes and robustness among different groups can be explained as the result of sexual selection.


Saradee Sengupta, Martín D. Ezcurra and Saswati Bandyopadhyay. 2017. A New Horned and Long-necked Herbivorous Stem-Archosaur from the Middle Triassic of India. Scientific Reports. 7, Article number: 8366. DOI: s41598-017-08658-8

Ezcurra MD. (2016The phylogenetic relationships of basal archosauromorphs, with an emphasis on the systematics of proterosuchian archosauriformsPeerJ 4:e1778 https://doi.org/10.7717/peerj.1778


The Enigmatic Chilesaurus and the evolution of ornithischian dinosaurs

Chilesaurus diegosuarezi (MACN)

Chilesaurus diegosuarezi is a bizarre dinosaur from the Upper Jurassic of southern Chile. Holotype specimen (SNGM-1935) consists of a nearly complete, articulated skeleton, approximately 1.6 m long. Four other partial skeletons (specimens SNGM-1936, SNGM-1937, SNGM-1938, SNGM-1888) were collected in the lower beds of Toqui Formation. All the preserved specimens of Chilesaurus show ventrally flexed arms with the hands oriented backwards, an arrangement that closely resembles the resting posture similar described in Mei long, Sinornithoides youngi, and Albinykus baatar. 

Chilesaurus possesses a number of surprisingly plesiomorphic traits on the hindlimbs, especially in the ankle and foot, which resemble basal sauropodomorphs; but the pubis closely resembles that of basal ornithischians. The bizarre anatomy of Chilesaurus raises interesting questions about its phylogenetic relationships. The features supporting the basal position of Chilesaurus within Tetanurae are: scapular blade elongate and strap-like; distal carpal semilunate; and manual digit III reduced.

Chilesaurus holotype cast (MACN)

But the position of Chilesaurus within within Tetanurae conflicts with the presence of several highly derived coelurosaurian features (e.g., opisthopubic pelvis, large supratrochanteric process on ilium, reduced supracetabular crest) which are present in combination with a number of surprisingly plesiomorphic traits present in basal sauropodomorphs.

Ornithischian features of Chilesaurus (From Baron and Barret, 2017)

Chilesaurus also shows several characters typical of ornithischians. The features include a premaxilla with an edentulous anterior region;  loss of recurvature in maxillary and dentary teeth; a postacetabular process that is 25–35% of the total anteroposterior length of the ilium; possession of a retroverted pubis; a pubis with a rod-like pubic shaft; a pubic symphysis that is restricted to the distal end of the pubis; and a femur that is straightened in anterior view.

The unique combination of ‘primitive’ and ‘derived’ characters for Chilesaurus has the potential to illuminate the order in which traditional ornithischian synapomorphies were acquired. For instance, Chilesaurus lacks a predentary bone, one of the features previously regarded as a fundamental ornithischian feature, although it possesses a retroverted pubis, suggesting that opisthopuby preceded the evolution of some craniodental modifications. Opisthopuby has also been related to herbivory, as it has been suggested that pubic retroversion might be related to the evolution of a more complex, longer digestive tract (Baron and Barret, 2017).


Baron MG, Barrett PM. 2017, A dinosaur missing-link? Chilesaurus and the early evolution of ornithischian dinosaurs. Biol. Lett. 13: 20170220. http://dx.doi.org/10.1098/rsbl.2017.0220

Nicolás R. Chimento, Federico L. Agnolin, Fernando E. Novas, Martín D. Ezcurra, Leonardo Salgado, Marcelo P. Isasi, Manuel Suárez, Rita De La Cruz, David Rubilar-Rogers & Alexander O. Vargas (2017) Forelimb posture in Chilesaurus diegosuarezi (Dinosauria, Theropoda) and its behavioral and phylogenetic implications. Ameghiniana doi: 10.5710/AMGH.11.06.2017.3088

Novas, F.E., Salgado, L., Suarez, M., Agnolín, F.L., Ezcurra, M.D., Chimento, N.R., de la Cruz, R., Isasi, M.P., Vargas, A.O., and Rubilar-Rogers, D. 2015. An enigmatic plant-eating theropod from the Late Jurassic period of Chile. Nature 522: 331-334. doi:10.1038/nature14307

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.



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.


Pisanosaurus revisited

Reconstructed skeleton of Pisanosaurus (Royal Ontario Museum)

Pisanosaurus mertii was originally described by Argentinian paleontologist Rodolfo Casamiquela in 1967, based on a poorly preserved but articulated skeleton from the upper levels of the Ischigualasto Formation (Late Triassic). The holotype and only known specimen (PVL 2577) is a fragmentary skeleton including partial upper and lower jaws, seven articulated dorsal vertebrae, four fragmentary vertebrae of uncertain position in the column, the impression of the central portion of the pelvis and sacrum, an articulated partial hind limb including the right tibia, fibula, proximal tarsals and pedal digits III and IV, the distal ends of the right and left femora, a left scapular blade (currently lost), a probable metacarpal III, and the impressions of some metacarpals (currently lost).

Pisanosaurus mertii holotype. Right lower mandible in medial (A) and lateral (B) views. Scale bar: 5 cm. From Agnolín and Rozadilla, 2017.

In the original description, Casamiquela considered that Pisanosaurus was a very distinct ornithischian, and even proposed a family: Pisanosauridae. The dentition and tooth-bearing bones of Pisanosaurus possess a large number of ornithischian traits, like its barricade-like dentition. But Pisanosaurus shows some features that strongly differ from those of ornithischians. For instance, vertebral centra are very elongated and transversely compressed, differing from the short and stout dorsal vertebrae of known ornithischians, including heterodontosaurids. The pelvis is another portion of the skeleton of Pisanosaurus strongly different from that of ornithischians.

Pisanosaurus mertii holotype. Dorsal vertebrae in left lateral (A) and right lateral (B) views. Scale bar: 5 cm. From Agnolín and Rozadilla, 2017.

On the other hand, Pisanosaurus shows some derived traits that resulted as unambiguous synapomorphies of the Silesauridae clade, and include: reduced to absent denticles on maxillary and dentary teeth; sacral ribs shared between two sacral vertebrae; lateral side of proximal tibia with a fibular flange (present also in heterodontosaurids and several saurischians); dorsoventrally flattened ungual phalanges; and ankylothecodonty, teeth partially fused to maxilla and dentary bone. The first and last characters are lacking in ornithischians. Of course, the inclusion of Pisanosaurus within Silesauridae implies that this taxon does not constitute the oldest ornithischian. This also suggests a significant gap between Pisanosaurus and the oldest unambiguous records of ornithischians: Laquintasaura and Lesothosaurus, which may be dated as Hettangian in age. This is consistent with previous interpretations proposing that ornithischian radiation occurred after the Triassic–Jurassic boundary.


Federico L. Agnolín & Sebastián Rozadilla (2017): Phylogenetic reassessment of Pisanosaurus mertii Casamiquela, 1967, a basal dinosauriform from the Late Triassic of Argentina, Journal of Systematic Palaeontology DOI: 10.1080/14772019.2017.1352623

Baron, M. G., Norman, D. B. & Barrett, P. M. A new hypothesis of dinosaur relationships and early dinosaur evolution.  Nature 543, 501–506  (2017).  doi:10.1038/nature21700

Padian K. The problem of dinosaur origins: integrating three approaches to the rise of Dinosauria. Earth and Environmental Science Transactions of the Royal Society of Edinburgh, Available on CJO 2013 doi:10.1017/S1755691013000431 (2013).

Meet Borealopelta markmitchelli

Holotype of Borealopelta markmitchelli (From Brown et al., 2017)

The Ankylosauria is a group of herbivorous, quadrupedal, armoured dinosaurs subdivided in two major clades, the Ankylosauridae and the Nodosauridae. The most derived members of this clade are characterized by shortened skulls, pyramidal squamosal horns, and tail clubs, among other features. Nodosauridae have a kinked ischium and more massive osteoderms, but lack a tail club. Ankylosaurs were present primarily in Asia and North America,  but the early origins of this clade are ambiguous. A three-dimensionally preserved ankylosaurian discovered in the Suncor Millennium Mine in northeastern Alberta, Canada, offers new evidence for understanding the anatomy of this group.

The new specimen, Borealopelta markmitchelli, from the Early Cretaceous of Alberta, preserves integumentary structures as organic layers, including continuous fields of epidermal scales and intact horn sheaths capping the body armor. The generic name Borealopelta is derived from “borealis” (Latin, “northern”) and “pelta” (Greek, “shield”). The specific epithet markmitchelli honors Mark Mitchell for his preparation of the holotype.

Schematic drawing of TMP 2011.033.0001 in dorsal view (From Brown et al., 2017)

The holotype (TMP 2011.033.0001), with an estimated living mass of 1,300 kg, is an articulated specimen preserving the head, neck, most of the trunk and sacrum, a complete right and a partial left forelimb and manus, and partial pes. The skull is covered in dermal plates, which are overlain by their associated epidermal scales. Cervical and thoracic osteoderms form continuous transverse rows completely separated by transverse rows of polygonal basement scale. Osteoderms are covered by a thick, dark gray to black organic layer, representing the original, diagenetically altered, keratinous epidermal scales. The distribution of the film correlates well to the expected distribution of melanin, a pigment present in some vertebrate integumentary structures. The keratinized tissues in this nodosaur are heavily pigmented. The possible presence of eumelanin and pheomelanin, suggested it had reddish-brown camouflage. The evidence of countershading in a large, heavily armored herbivorous dinosaur also provides a unique insight into the predator-prey dynamic of the Cretaceous Period.



Brown, C.M.; Henderson, D.M.; Vinther, J.; Fletcher, I.; Sistiaga, A.; Herrera, J.; Summons, R.E. “An Exceptionally Preserved Three-Dimensional Armored Dinosaur Reveals Insights into Coloration and Cretaceous Predator-Prey Dynamics”. Current Biology. doi:10.1016/j.cub.2017.06.071

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