Historical perspective on the dinosaur family tree

Megalosaurus at Crystal Palace Park, London. From Wikimedia Commons.

In the 19th century, the famous Victorian anatomist Richard Owen diagnosed Dinosauria using three taxa: Megalosaurus, Iguanodon and Hylaeosaurus, on the basis of three main features: large size and terrestrial habits, upright posture and sacrum with five vertebrae (because the specimens were from all Late Jurassic and Cretaceous, he didn’t know that the first dinosaurs had three or fewer sacrals). These characteristics were more mammalian than reptilian. But new fossil findings from Europe and particularly North America forced to a new interpretation about those gigantic animals.

In 1887, Harry Govier Seeley summarised the works of Cope, Huxley and Marsh who already subdivided the group Dinosauria into various orders and suborders. However, he was the first to subdivide dinosaurs into Saurischians and the Ornithischians, based on the nature of their pelvic bones and joints. He wrote: The characters on which these animals should be classified are, I submit, those which pervade the several parts of the skeleton, and exhibit some diversity among the associated animal types. The pelvis is perhaps more typical of these animals than any other part of the skeleton and should be a prime element in classification. The presence or absence of the pneumatic condition of the vertebrae is an important structural difference…” Based on these features, Seeley denied the monophyly of dinosaurs.

Seeley’s (1901) diagram of the relationships of Archosauria. From Padian 2013

At the mid 20th century, the consensual views about Dinosauria were: first, the group was not monophyletic; second almost no Triassic ornithischians were recognised, so they were considered derived morphologically, which leads to the third point, the problem of the ‘‘origin of dinosaurs’’ usually was reduced to the problem of the ‘‘origin of Saurischia,’’ because theropods were regarded as the most primitive saurischians. A great influence on the views about the dinosaur origins was Alan Charig. He was Curator of Amphibians, Reptiles and Birds at the British Museum (Natural History), now the Natural History Museum, in London for almost thirty years. Charig thought that the first dinosaurs were quadrupedal, not bipedal. He based this on the kinds of animals that he and his colleagues found in the early Triassic localities of eastern and South Africa. He thought that forms such as ‘‘Mandasuchus’’ were related to dinosaurs, but that they had a posture intermediate between a sprawling and upright gait that he called ‘‘semi-improved” or ‘‘semi-erect’’.

Herrerasaurus skull. From Wikimedia Commons.

The discovery of Lagosuchus and Lagerpeton from the Middle Triassic of Argentina induced a change in the views of dinosaurs origins. Also from South America came Herrerasaurus from the Ischigualasto Formation, the basal sauropodomorphs Saturnalia, Panphagia, Chromogisaurus, and the theropods Guibasaurus and Zupaysaurus, but no ornithischians except a possible heterodontosaurid jaw fragment from Patagonia. The 70s marked the beginning of a profound shift in thinking on nearly all aspects of dinosaur evolution, biology and ecology. Robert Bakker and Peter Galton, based on John Ostrom’s vision about Dinosauria, proposed, for perhaps the first time since 1842, that Dinosauria was indeed a monophyletic group and that it should be separated (along with birds) from other reptiles as a distinct ‘‘Class”. In 1986, the palaeontologist Jacques Gauthier showed that dinosaurs form a single group, which collectively has specific diagnostic traits that set them apart from all other animals.

From Baron et al., 2017.

Phylogenetic analyses of early dinosaurs have  supported the traditional scheme. But back in March of this year, a paper, authored by Matthew Baron, David Norman and Paul Barrett, challenged this paradigm with a new phylogenetic analysis that places theropods and ornithischians together in a group called Ornithoscelida. The team analysed a wide range of dinosaurs and dinosauromorphs (74 taxa were scored for 457 characters), and they arrived at a dinosaur evolutionary tree containing one main branch that subdivides into the groupings of Ornithischia and Theropoda, and a second main branch that contains the Sauropoda and Herrerasauridae (usually positioned as either basal theropods or basal Saurischia, or outside Dinosauria but close to it). The term Ornithoscelida was coined in 1870 by Thomas Huxley for a group containing the historically recognized groupings of Compsognatha, Iguanodontidae, Megalosauridae and Scelidosauridae. The synapomorphies that support the formation of the clade Ornithoscelida includes: an anterior premaxillary foramen located on the inside of the narial fossa; a sharp longitudinal ridge on the lateral surface of the maxilla; short and deep paroccipital processes; a post-temporal foramen enclosed within the paroccipital process; a straight femur, without a sigmoidal profile; absence of a medioventral acetabular flange; a straight femur, without a sigmoidal profile; and fusion of the distal tarsals to the proximal ends of the metatarsals.

Of course, those results have great implications for the very origin of dinosaurs. Ornithischia don’t begin to diversify substantially until the Early Jurassic. By contrast, the other dinosaurian groups already existed by at least the early Late Triassic. If the impoverished Triassic record of ornithischians reflects a true absence, ornithischians might have evolved from theropods in the Late Triassic (Padian, 2017). The study also suggest that dinosaurs might have originated in the Northern Hemisphere, because most of their basal members, as well as their close relatives, are found there. Furthermore, their analyses places the origin of dinosaurs at the boundary of the Olenekian and Anisian stages (around 247 Ma), slightly earlier than has been suggested previously.


The dinosaur family tree Credit: Max Langer

More recently, an international team of early dinosaur evolution specialists, led by Max Langer, highlighted that the lack of some important taxa (for example, the early thyreophoran Scutellosaurus, the possible theropod Daemonosaurus, and the newly described Ixalerpeton and Buriolestes) may have a substantial effect on character optimizations near the base of the dinosaur tree, and thus on the interrelationships of early dinosaurs. The study did not find strong evidence to discard the traditional Ornithischia–Saurischia division. But they reintroduced a third possibility that was articulated in the 1980s but rarely discussed since: that sauropodomorphs and ornithischians may form their own herbivorous group, separate from the ancestrally meat-eating theropods. The Phytodinosauria hypothesis was coined by Robert T. Bakker in his book The Dinosaur Heresies: “Therefore all the plant-eating dinosaurs of every sort really constitute one, single natural group branching out from one ancestor, a primitive anchisaurlike dinosaur. And a new name is required for this grand family of vegetarians. So I hereby christen them the Phytodinosauria, the “plant dinosaurs”‘


Max C. Langer, Martín D. Ezcurra, Oliver W. M. Rauhut, Michael J. Benton, Fabien Knoll, Blair W. McPhee, Fernando E. Novas, Diego Pol & Stephen L. Brusatte, Untangling the dinosaur family tree, Nature 551 (2017) doi; oi:10.1038/nature24012

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. Dividing the dinosaurs. Nature 543, 494–495 (2017) doi:10.1038/543494a

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).

Seeley, H. G. On the classification of the fossil animals commonly named DinosauriaProc. R. Soc. Lond. 43165171 (1887).

Huxley, T. H. On the classification of the Dinosauria, with observations on the Dinosauria of the Trias. Quarterly Journal of the Geological Society, London 26, 32-51. (1870).


The Winds of Winter

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

Almost 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, including non-avian dinosaurs, other vertebrates, marine reptiles and invertebrates, planktonic foraminifera and ammonites. Marine ecosystems lost about half of their species while freshwater environments shows low extinction rates, about 10% to 22% of genera.

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

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. This phenomenon is called “impact winter”.

Recent drilling of the peak ring of the Chicxulub impact crater has been used to create 3-D numerical simulations of the crater formation. It was estimate that the angle of impact at Chicxulub was ~60° with a downrange direction to the southwest. The new study indicates that the impact may have released around three times as much sulfur and much less carbon dioxide compared with previous calculations, suggesting that surface temperatures were likely to have been significantly reduced for several years and ocean temperatures affected for hundreds of years after the Chicxulub impact.



Artemieva, N., Morgan, J., & Expedition 364 Science Party (2017). Quantifying the release of climate-active gases by large meteorite impacts with a case study of Chicxulub. Geophysical Research DOI: 10.1002/2017GL074879


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.


Halloween special V: Lovecraft’s paleontological Journey

H.P. Lovecraft’s love for astronomy is well known. As an amateur astronomer, Lovecraft attended several lectures from leading astronomers and physicists of his time. In 1906 he wrote a letter to the Scientific American on the subject of  finding planets in the solar system beyond Neptune. Around this time he began to write two astronomy columns for the Pawtuket Valley Gleaner and the Providence Tribune. He also wrote a treatise, A Brief Course in Astronomy – Descriptive, Practical, and Observational; for Beginners and General Readers. In several of his astronomical articles he describes meteors as  “the only celestial bodies which may be actually touched by human hands”. But Lovecraft was also obsessed with the concept of deep time, so geology and paleontology were also present in his writings.

Lovecraft’s monsters are certainly titanic, biologically impossible beings, from dimensions outside our own. He began conjuring monsters almost from the start of his career. In “The Nameless City”, published in the November 1921 issue of the amateur press journal The Wolverine, and often considered the first Cthulhu Mythos story, he describes an ancient race of reptiles that built the city: “They were of the reptile kind, with body lines suggesting sometimes the crocodile, sometimes the seal, but more often nothing of which either the naturalist or the palaeontologist ever heard.”  

Panorama of Ross Island showing Hut Point Peninsula (foreground), Mount Erebus (left) and Mount Terror (right), Antarctica. Photo: John Bortniak, NOAA

According to his biographer S. T. Joshi, Lovecraft was fascinated by Antarctica since an early age. Much of this fascination is recognizable in his famous novel “At the Mountains of Madness”, written in 1931. The novel was rejected by Weird Tales and finally was published by Astounding Stories in a serial form in 1936. “At the Mountains of Madness” is told from the perspective of William Dyer, a geologist from Miskatonic University who flies into an unexplored region of Antarctica. He’s accompanied by Professor Lake, a biologist, Professor Pabodie, an engineer, and some graduate students. The basic plot of the novel is the discovery of the frozen remains of bizarre entities from the deep space and their even more terrifying “slaves”:  the  shoggoths. The story could be divided in two parts. The first one is particularly rich, detailed and shows an impressive scientific erudition. This is clear in the following paragraph when he describes something that Professor Lake found: “He  was strangely convinced that the marking was the print of some bulky, unknown, and radically unclassifiable organism of considerably advanced evolution, notwithstanding that the rock which bore it was of so vastly ancient a date—Cambrian if not actually pre-Cambrian— as to preclude the probable existence not only of all highly evolved life, but of any life at all above the unicellular or at most the trilobite stage. These fragments, with their odd marking, must have been 500 million to a thousand million years old”. 

Of course, one of the most fascinating parts of the novel is the description of the Elder Things: “Cannot yet assign positively to animal or vegetable kingdom, but odds now favour animal. Probably represents incredibly advanced evolution of radiata without loss of certain primitive features. Echinoderm resemblances unmistakable despite local contradictory evidences. Wing structure puzzles in view of probable marine habitat, but may have use in water navigation. Symmetry is curiously vegetable-like, suggesting vegetable’s essentially up-and-down structure rather than animal’s fore-and-aft structure. Fabulously early date of evolution, preceding even simplest Archaean protozoa hitherto known, baffles all conjecture as to origin.” According with  S.T. Joshi, Lovecraft based his description of the Elder Thing in the fossil crinoids drawn by E. Haeckel in  Kunstformen der Natur.

E. Haeckel’s Kunstformen der Natur (1904), plate 90: Cystoidea. From Wikimedia Commons

“The Shadow Out of Time” (1935) was H. P. Lovecraft’s last major story. It’s told from the perspective of Nathaniel Wingate Peaslee, a professor of political economy at Miskatonic University. During five years, this man suffers a bizarre form of amnesia  followed by vivid dreams of aliens cities in ancient landscapes.  Later, Peaslee discovered that a small number of people throughout history suffered the same type of amnesia. They were possessed by the Great Race, a group of cone shaped creatures who developed the technique of swapping minds with creatures of another era with the purpose of learn the secrets of the Universe. Peaslee describes the gardens that surround the cities of his visions with detail. There was calamites, cycads, trees of coniferous aspect, and small, colourless flowers: “The far horizon was always steamy and indistinct, but I could see that great jungles of unknown tree-ferns, calamites, lepidodendra, and sigillaria lay outside the city, their fantastic frondage waving mockingly in the shifting vapours.”

Calamites was a genus of tree-sized, spore-bearing plants that lived during the Carboniferous and Permian periods (about 360 to 250 million years ago), closely related to modern horsetails. They reached their peak diversity in the Pennsylvanian and were major constituents of the lowland equatorial swamp forest ecosystems. The Cycadales are an ancient group of seed plants that can be traced back to the Pennsylvanian. Cycads have a stem or trunk that commonly looks like a large pineapple and composed of the coalesced bases of large leaves.  Today’s cycads are found in the tropical, subtropical and warm temperate regions of both the north and south hemispheres.

Lepidodendron (fossil tree) on display at the State Museum of Pennsylvania, From Wikimedia Commons

Lepidodendron was a tree-like (‘arborescent’) tropical plant, related to the lycopsids. The name of the genus comes from the Greek lepido, scale, and dendron, tree, because of the distinctive diamond shaped pattern of the bark. The name Lepidodendron is a generic name given to several fossil that clearly come from arborescent lycophytes but are difficult to assign to one species. Fossil remains indicate that some trees attained heights in excess of 40 m and were at least 2m in diameter at the base. They were dominant components of swamp ecosystems in the Carboniferous and frequently associated with Sigillaria, another extinct genus of tree-sized lycopsids from the Carboniferous Period. The absence of extensive branching and the structure of the leaf bases are the principal feature that distinguish Sigillaria from other lycopsids (Taylor et al, 2009). Sigillariostrobus is the name assigned to the reproductive organs or cones of Sigillaria. Unlike Lepidodendron cones, which were attached attached individually near the tip of the branches, Sigillaria cones occurred in clusters attached in certain places along the upper stem.

Tunguska forest (Photograph taken by Evgeny Krinov near the Hushmo river, 1929).

“The Colour Out of Space” is a short story written by  H. P. Lovecraft in 1927.  The story is set in the fictional town of Arkham, Massachusetts, where an unnamed narrator investigates a local area known as the “blasted heath”. Ammi Pierce, a local man, relates him the tragic story of a man named Nahum Gardner and how his life crumbled when a great rock fell out of the sky onto his farm. Within the meteorite there was a coloured globule impossible to describe that infected Gardner’s family, and spread across the property, killing all living things. It’s the first of Lovecraft’s major tales that combines horror and science fiction. The key question of the story of course is the meteorite. Although “the coloured globule” inside the meteorite has mutagenic properties we cannot define their nature. But as Lovecraft stated once, the things we fear most are those that we are unable to picture.

“The Colour Out of Space” was published nineteen year after the Tunguska Event. On the morning of June 30, 1908, eyewitnesses reported a large fireball crossing the sky above Tunguska in Siberia. The object entered Earth’s atmosphere traveling at a speed of about 33,500 miles per hour and released the energy equal to 185 Hiroshima bombs. The night skies glowed and the resulting seismic shockwave was registered with sensitive barometers as far away as England. In 1921, Leonid Kulik, the chief curator for the meteorite collection of the St. Petersburg museum led an expedition to Tunguska, but failed in the attempt to reach the area of the blast. Later, in 1927, a new expedition, again led by Kulik, discovered the huge area of leveled forest that marked the place of the Tunguska “meteorite” fall. At the time, Kulik mistook shallow depressions called thermokarst holes for many meteorites craters. However, he didn’t find remnants of the meteorite, and continued to explore the area until World War II. In the early 1930s, British astronomer Francis Whipple suggested that the Tunguska Event was caused by the core of a small comet, while Vladimir Vernadsky, suggested the cause was a lump of cosmic matter. (Rubtsov, 2009). More than a century later the cause of the Tunguska Event remains a mystery.



Lovecraft, H. P, “At the Mountains of Madness”, Random House, 2005.

Lovecraft, H. P, “The Dreams in the Witch House and Other Weird Stories”, Penguin Books, 2004.

Joshi, S. T. (2001). A dreamer and a visionary: H.P. Lovecraft in his time. Liverpool University Press, 302.

LONG, J. (2003): Mountains of Madness – A Scientist’s Odyssey in Antarctica. Jospeh Henry Press, Washington: 252

N. Taylor, Edith L. Taylor, Michael Krings: “Paleobotany: The Biology and Evolution of Fossil Plants”. 2nd ed., Academic Press 2009.

Kathy Willis, Jennifer McElwain, The Evolution of Plants, Oxford University Press, 2013



Brief history of the Ocean Acidification through time: an update

Corals one of the most vulnerable creatures in the ocean. Photo Credit: Katharina Fabricius/Australian Institute of Marine Science

About one third of the carbon dioxide released by anthropogenic activity is absorbed by the oceans. Once dissolved in seawater, most of the CO2 is transported into deep waters via thermohaline circulation and the biological pump. But a smaller fraction of the CO2, forms carbonic acid and causes a decline in pH in the surface ocean. This phenomenon is called ocean acidification, and is occurring at a rate faster than at any time in the last 300 million years.

Acidification affects the biogeochemical dynamics of calcium carbonate, organic carbon, nitrogen, and phosphorus in the ocean and interferes with a range of processes, including growth, calcification, development, reproduction and behaviour in a wide range of marine organisms like planktonic coccolithophores, foraminifera, pteropods and other molluscs,  echinoderms, corals, and coralline algae.

The pH within the ocean surface has decreased ~0.1 pH units since the industrial revolution and is predicted to decrease an additional 0.2 – 0.3 units by the end of the century. An eight-year study carried out by the Biological Impacts of Ocean Acidification group (Bioacid), with the support of the German government, has contributed to quantifying the effects of ocean acidification on marine organisms and their habitats. Among the many effects of ocean acidification on marine organisms are including: decreased rate of skeletal growth in reef-building corals, reduced ability to maintain a protective shell among free-swimming zooplankton, and reduced survival of larval marine species, including commercial fish and shellfish. Even worse, the effects of acidification can intensify the effects of global warming, in a dangerous feedback loop.

Coccolithophores exposed to differing levels of acidity. Adapted by Macmillan Publishers Ltd: Nature Publishing Group, Riebesell, U., et al., Nature 407, 2000.

The geologic record of ocean acidification provide valuable insights into potential biotic impacts and time scales of recovery.  Rapid additions of carbon dioxide during extreme events in Earth history, including the end-Permian mass extinction (252 million years ago) and the Paleocene-Eocene Thermal Maximum (PETM, 56 million years ago) may have driven surface waters to undersaturation. But, there’s  no perfect analog for our present crisis, because we are living in an “ice house” that started 34 million years ago  with the growth of ice sheets on Antarctica, and this cases corresponded to events initiated during “hot house” (greenhouse) intervals of Earth history.

The end-Permian extinction is the most severe biotic crisis in the fossil record, with as much as 95% of the marine animal species and a similarly high proportion of terrestrial plants and animals going extinct . This great crisis ocurred about 252 million years ago (Ma) during an episode of global warming. The cause or causes of the Permian extinction remain a mystery but new data indicates that the extinction had a duration of 60,000 years and may be linked to massive volcanic eruptions from the Siberian Traps. The same study found evidence that 10,000 years before the die-off, the ocean experienced a pulse of light carbon that most likely led to a spike of carbon dioxide in the atmosphere. This could have led to ocean acidification, warmer water temperatures that effectively killed marine life.

Taxonomic variation in effects of ocean acidification (From Kroeker et al. 2010)

The early Aptian Oceanic Anoxic Event (120 million years ago) was an interval of dramatic change in climate and ocean circulation. The cause of this event was the eruption of the Ontong Java Plateau in the western Pacific, wich led to a major increase in atmospheric pCO2 and ocean acidification. This event was characterized by the occurrence of organic-carbon-rich sediments on a global basis along with evidence for warming and dramatic change in nanoplankton assemblages. Several oceanic anoxic events (OAEs) are documented in Cretaceous strata in the Canadian Western Interior Sea.

The Paleocene-Eocene Thermal Maximum (PETM; 55.8 million years ago) was a short-lived (~ 200,000 years) global warming event. Temperatures increased by 5-9°C. It was marked by the largest deep-sea mass extinction among calcareous benthic foraminifera in the last 93 million years. Similarly, planktonic foraminifer communities at low and high latitudes show reductions in diversity. The PETM is also associated with dramatic changes among the calcareous plankton,characterized by the appearance of transient nanoplankton taxa of heavily calcified forms of Rhomboaster spp., Discoaster araneus, and D. anartios as well as Coccolithus bownii, a more delicate form.

The current rate of the anthropogenic carbon input  is probably greater than during the PETM, causing a more severe decline in ocean pH and saturation state. Also the biotic consequences of the PETM were fairly minor, while the current rate of species extinction is already 100–1000 times higher than would be considered natural. This underlines the urgency for immediate action on global carbon emission reductions.


David A. Hutchins & Feixue Fu, Microorganisms and ocean global change, Nature Microbiology 2, Article number: 17058 (2017) doi:10.1038/nmicrobiol.2017.58 

Kump, L.R., T.J. Bralower, and A. Ridgwell. 2009. Ocean acidification in deep time. Oceanography 22(4):94–107, http://dx.doi.org/10.5670/oceanog.2009.100

Parker, L. M. et al. Adult exposure to ocean acidification is maladaptive for larvae of the Sydney rock oyster Saccostrea glomerata in the presence of multiple stressors. Biology Letters 13 (2017). DOI: 10.1098/rsbl.2016.0798

Kristy J. Kroeker, Rebecca L. Kordas, Ryan N. Crim, Gerald G. Singh, Meta-analysis reveals negative yet variable effects of ocean acidification on marine organisms, Ecology Letters (2010) 13: 1419–1434
DOI: 10.1111/j.1461-0248.2010.01518.x


Vegaviidae, a new clade of southern diving birds

Vegavis iaai by Gabriel Lio. / Photo: CONICET

The fossil record of Late Cretaceous–Paleogene modern birds in the Southern Hemisphere is fragmentary.  It includes Neogaeornis wetzeli from Maastrichtian beds of Chile, Polarornis gregorii and Vegavis iaai from the Maastrichtian of Antarctica, and Australornis lovei from the Paleogene of New Zealand. The phylogenetic relationships of these taxa have been variously interpreted by different authors. In a more recent analysis, Polarornis, Vegavis, Neogaeornis, and Australornis, are including in a new clade: Vegaviidae.

Vegaviids share a combination of characters related to diving adaptations, including compact and thickened cortex of hindlimb bones, femur with anteroposteriorly compressed and bowed shaft, deep and wide popliteal fossa delimited by a medial ridge, tibiotarsus showing notably proximally expanded cnemial crests, expanded fibular crest, anteroposterior compression of the tibial shaft, and a tarsometatarsus with a strong transverse compression of the shaft.

Histological sections of Vegavis iaai (MACN-PV 19.748) humerus (a), femur (b), polarized detail of humerus (c). Scale bar equals 10 mm for (a), (b) and 5 mm for (c). From Agnolín et al., 2017

The recognition of Polarornis, Vegavis, Neogaeornis, Australornis, and a wide array of isolated specimens as belonging to the new clade Vegaviidae reinforces the hypothesis that southern landmasses constituted a center for neornithine diversification, and emphasizes the role of Gondwana for the evolutionary history of Anseriformes and Neornithes.

The most informative source for anatomical comparison among Australornis, Polarornis, Vegavis as well as other southern avian is a recently published Vegavis skeleton (MACN-PV 19.748). Vegavis overlaps with Australornis in the proximal portion of the humerus, proximal part of the coracoid, scapula, and ulna; with Polarornis in the humerus, femur, and proximal end of the tibia; and with Neogaeornis in the tarsometatarsus.

Phylogeny with geographical distribution of Vegaviidae. From Agnolín et al., 2017.

The humerus is probably the most diagnostic element among anseriforms. In Vegavis and Australornis the humerus is notably narrow and medially tilted on its proximal half, and the deltopectoral crest extends for more than one third of the humeral length. The femur is well known both in Vegavis and Polarornis, and share a combination of characters absent in other Mesozoic or Paleogene birds, including strongly anteriorly bowed and anteroposteriorly compressed shaft (especially near its distal end)

Osteohistological analysis of the femur and humerus of V. iaai. shows a highly vascularized fibrolamellar matrix lacking lines of arrested growths, features widespread among modern birds. The femur has some secondary osteons, and shows several porosities, one especially large, posterior to the medullar cavity. The humerus exhibits a predominant fibrolamellar matrix, but in a portion of the anterior and medial sides of the shaft there are a few secondary osteons, some of them connected with Volkman’s canals, and near to these canals, there are a compact coarse cancellous bone (CCCB) with trabeculae. This tissue disposition and morphology suggests that Vegavis had remarkably high growth rates, a physiological adaptation that may be critical for surviving in seasonal climates at high latitudes, and  may also constitute the key adaptation that allowed vegaviids to survive the K/T mass extinction event.



Agnolín, F.L., Egli, F.B., Chatterjee, S. et al. Sci Nat (2017) 104: 87. https://doi.org/10.1007/s00114-017-1508-y

Jordi Alexis Garcia Marsà, Federico L. Agnolín & Fernando Novas (2017): Bone microstructure of Vegavis iaai (Aves, Anseriformes) from the Upper Cretaceous of Vega Island, Antarctic Peninsula, Historical Biology, DOI: 10.1080/08912963.2017.1348503

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


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

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