Ingentia prima, the first giant

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

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

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

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

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

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



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.


Mary Anning, ‘the greatest fossilist the world ever knew’.

Duria Antiquior famous watercolor by the geologist Henry de la Beche based on fossils found by Mary Anning. From Wikimedia Commons.

By the 19th century, the study of the Earth became central to the economic and cultural life of Great Britain. Women were free to take part in collecting fossils and mineral specimens, and they were allowed to attend lectures but they were barred from membership in scientific societies. England was ruled by an elite, and of course, these scholarly activities only occurred within the upper echelon of British society. Notwithstanding, the most famous fossilist of the 19th century was a women of a low social station: Mary Anning.

Mary Anning was born on Lyme Regis on May 21, 1799. Her father was a carpenter and an amateur fossil collector who died when Mary was eleven. He trained Mary and her brother Joseph in how to look and clean fossils. After the death of her father, Mary and Joseph used those skills to search fossils that they sold as “curiosities”. The source of those fossils was the coastal cliffs around Lyme Regis, part of a geological formation known as the Blue Lias.

The shore of Lyme Bay where Mary Anning did most of her collecting.

Invertebrate fossils, like ammonoids or belemnites, were the most common findings. But when Mary was 12, her brother Joseph found a skull protruding from a cliff and few month later, Mary found the rest of the skeleton. They sold it for £23. Later, in 1819, the skeleton was purchased by Charles Koenig of the British Museum of London who suggested the name “Ichthyosaur” for the fossil.

In 1819 the Annings were in considerable financial difficulties. They were rescued by the generosity of Thomas James Birch (1768–1829), who arranged for the sale of his personal collection, largely purchased from the Annings, in Bullock’s Museum in London.  The auction took place in May 1820, during which George Cuvier bought several pieces for the Muséum National d’Histoire Naturelle.

Mary Anning’s sketch of belemnites. From original manuscripts held at the Natural History Museum, London. © The Natural History Museum, London

On December 10, 1823, she discovered the first complete Plesiosaur skeleton at Lyme Regis in Dorset. The fossil was acquired by the Duke of Buckingham. Noticed about the discovery, George Cuvier wrote to William Conybeare suggesting that the find was a fake produced by combining fossil bones from different animals. William Buckland and Conybeare sent a letter to Cuvier including anatomical details, an engraving of the specimen and a sketch made by Mary Morland (Buckland’s wife) based on Mary Anning’s own drawings and they convinced Cuvier that this specimen was a genuine find. From that moment, Cuvier treated Mary Anning as a legitimate and respectable fossil collector and cited her name in his publications.

Autograph letter about the discovery of plesiosaurus, by Mary Anning. From original manuscripts held at the Natural History Museum, London. © The Natural History Museum, London

By the age of 27, Mary was the owner of a little shop: Anning’s Fossil Depot. Many scientist and fossil collectors from around the globe went to Mary´s shop. She was friend of Henry De la Beche, the first director of the Geological Survey of Great Britain, who knew Mary since they were both children and lived in Lyme Regis. De la Beche was a great supporter of Mary’s work. She also corresponded with Charles Lyell, William Buckland and Mary Morland, Adam Sedgwick and Sir Roderick Murchison.
It’s fairly to say that Mary felt secure in the world of men, and a despite her religious beliefs, she was an early feminist. In an essay in her notebook, titled Woman!, Mary writes:  “And what is a woman? Was she not made of the same flesh and blood as lordly Man? Yes, and was destined doubtless, to become his friend, his helpmate on his pilgrimage but surely not his slave…”

A) Mary Anning (1799- 1847) B) William Buckland (1784- 1856)

On December of 1828, Mary found the first pterosaur skeleton outside Germany. William Buckland made the announcement of Mary’s discovery in the Geological Society of London and named Pterodactylus macronyx in allusion to its large claws. The skull of Anning’s specimen had not been discovered, but Buckland thought that the fragment of jaw in the collection of the Philpot sisters of Lyme belonged to a pterosaur.
In 1829,  Mary Anning discovered Squaloraja polyspondyle, a fish. Unfortunately, the specimen was lost in the destruction of the Bristol Museum by a German bombing raid in November, 1940.
From her correspondence is clear that Mary learned anatomy by dissecting modern organisms. In a letter to J.S. Miller of the Bristol Museum, dated 20 January 1830, she wrote: “…I have dissected a Ray since I received your letter, and I do not think it the same genus, the Vertebrae alone would constitute it a different genus being so unlike any fish vertebrae they are so closely anchylosed that they look like one bone but being dislocated at two places show that each thin line is a separate vertebrae with the ends flat…”. 

Sketch of Mary Anning by Henry De la Beche.

Mary Anning, ‘the greatest fossilist the world ever knew’, died of breast cancer on 9 March, 1847, at the age of 47. She was buried in the cemetery of St. Michaels. In the last decade of her life, Mary received  three accolades. The first was an annuity of £25, in return for her many contributions to the science of geology. The second was in 1846, when the geologists of the Geological Society of London organized a further subscription for her. The third accolade was her election, in July 1846, as the first Honorary Member of the new Dorset County Museum in Dorchester.

After her death, Henry de la Beche, Director of the Geological Survey and President of the Geological Society of London, wrote a very affectionate obituary published in the Quarterly Journal of the Geological Society on February 14, 1848, the only case of a non Fellow who received that honour.

Mary Anning’s Window, St. Michael’s Church. From Wikimedia Commons.

In February 1850 Mary was honoured by the unveiling of a new window in the parish church at Lyme, funded through another subscription among the Fellows of the Geological Society of London, with the following inscription: “This window is sacred to the memory of Mary Anning of this parish, who died 9 March AD 1847 and is erected by the vicar and some members of the Geological Society of London in commemoration of her usefulness in furthering the science of geology, as also of her benevolence of heart and integrity of life.”

In 1865, Charles Dickens wrote an article about Mary Anning’s life in his literary magazine “All the Year Round”, where emphasised the difficulties she had overcome: “Her history shows what humble people may do, if they have just purpose and courage enough, toward promoting the cause of science. The inscription under her memorial window commemorates “her usefulness in furthering the science of geology” (it was not a science when she began to discover, and so helped to make it one), “and also her benevolence of heart and integrity of life.” The carpenter’s daughter has won a name for herself, and has deserved to win it.”


Buckland, Adelene: Novel Science : Fiction and the Invention of Nineteenth-Century Geology, University of Chicago Press, 2013.

BUREK, C. V. & HIGGS, B. (eds) The Role of Women in the History of Geology. Geological Society, London, Special Publications, 281, 1–8. DOI: 10.1144/SP281.1.

Davis, Larry E. (2012) “Mary Anning: Princess of Palaeontology and Geological Lioness,”The Compass: Earth Science Journal of Sigma Gamma Epsilon: Vol. 84: Iss. 1, Article 8.

Hugh Torrens, Mary Anning (1799-1847) of Lyme; ‘The Greatest Fossilist the World Ever Knew’, The British Journal for the History of Science Vol. 28, No. 3 (Sep., 1995), pp. 257-284. Published by: Cambridge University Press.

De la Beche, H., 1848a. Obituary notices. Quarterly Journal of the Geological Society of London, v. 4: xxiv–xxv.

Dickens, C., 1865. Mary Anning, the fossil finder. All the Year Round, 13 (Feb 11): 60–63.



Ichthyornis and the evolution of the avian skull.


Ichthyornis skull

Birds originated from a theropod lineage more than 150 million years ago. By the Early Cretaceous, they diversified, evolving into a number of groups of varying anatomy and ecology. Much of birds anatomical variety is related to their skulls and in particulary with their beaks.

Discovered in 1870 by Benjamin Franklin Mudge, a professor from Kansas State Agricultural College and good friend of Othniel Charles Marsh, Ichthyornis, which means‭ ‘‬fish bird‭’‭, was a small early ornithuromorph from the Late Cretaceous of North America. Ornithuromorphs, include Gansus, Patagopteryx, Yixianornis, and Apsaravis, which form a grade on the line to Ornithurae, a derived subgroup that includes modern birds and their closest fossil relatives.


3D reconstruction of the skull of I. dispar (From Field et al., 2018)

The skull of I. dispar shows a transitional point in the evolutionary history of birds. The upper margin of the beak is concave in profile, a derived condition shared with living birds. The fused, toothless premaxillae have a terminal hook, and occupy the anterior quarter of the rostrum. Neurovascular foramina indicate the presence of a highly keratinized region of rhamphotheca called the premaxillary nail. The maxilla is plesiomorphically long. The dentition is extensive in both upper and lower jaws. A sulcus on the rostral half of the maxilla suggests a broad naso-maxillary contact and a correspondingly broad postnarial bar. The palatine is narrow and elongate, unlike that of Archaeopteryx and more stemward theropods. The quadrate exhibits two rounded capitular condyles that fit into cotyles on the prootic and squamosal bones to form a mobile joint with the cranium. The arrangement of the rostrum, jugal, and quadratojugal, the mobile suspensorium and the narrow, linear palatine all indicate that I. dispar possessed a fully functional avian cranial kinetic system.

The endocranial cavity appears essentially modern in sagittal section. The forebrain was enlarged and posteroventrally rotated while the optic lobes were inflated and laterally shifted, as in living birds. The squamosal exhibits an archaic, deinonychosaur-like morphology. The zygomatic process is deep and triangular in lateral view. The nuchal crest extends from the midline of the skull onto the zygomatic process, forming the upper edge of the squamosal bone, as in non-avialan theropods.

Darwin’s letter to Marsh (Yale Peabody Museum Archives)

Since its discovery, Ichthyornis has been viewed as a classical example of evolution, due to the combination of an advanced postcranial morphology and retention of toothed jaws. In a letter, dated August 31, 1880, Charles Darwin thanks Marsh for a copy of his monograph Odontornithes, which reported two contrasting bird genera: Hesperornis, which was about 1.8 metres tall, and Ichthyornis, which had an average wingspan of about 60 centimetres. In his letter, Darwin wrote: “I received some time ago your very kind note of July 28th, & yesterday the magnificent volume. I have looked with renewed admiration at the plates, & will soon read the text. Your work on these old birds & on the many fossil animals of N. America has afforded the best support to the theory of evolution, which has appeared within the last 20 years.”



Daniel J. Field, Michael Hanson, David Burnham, Laura E. Wilson, Kristopher Super, Dana Ehret, Jun A. Ebersole & Bhart-Anjan S. Bhullar, Complete Ichthyornis skull illuminates mosaic assembly of the avian head, Nature (2018).
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.


An early juvenile enantiornithine specimen from the Early Cretaceous of Spain

The slab and counterslab of MPCM-LH-26189

Mesozoic remains of juvenile birds are rare. To date, the only records are from the Early Cretaceous of China and Spain, from the mid-Cretaceous of  Myanmar, and from the Late Cretaceous of Argentina and Mongolia. The most recent finding from the Early Cretaceous of Las Hoyas, Spain, provide an insight into the osteogenesis of the Enantiornithes, the most abundant clade of Mesozoic birds. Previous records of Enantiornithes from the Las Hoyas fossil site include: Eoalulavis hoyasi, Concornis lacustris, and Iberomesornis romerali.

The latest specimen, MPCM-LH-26189, a nearly complete and largely articulated skeleton (only the feet, most of its hands, and the tip of the tail are missing), is very small. The specimen died around the time of birth, a crucial moment to study the osteogenesis in birds. The skull, is partially crushed, and is large compared to the body size. The braincase is fractured. The frontals and the parietals form a uniformly curved cranial vault. The cerebrocast shows a very slight inflation, suggesting that the cerebral anatomy of MPCM-LH-26189 falls in between that of the Archaeopteryx, and the putative basal ornithurine Cerebavis, whose telencephalic expansion is close to most extant birds. The cervical series is composed of 9 vertebrae. There are 10  thoracic vertebrae, and the sacrum appears to be composed of 5–6 vertebrae. The prezygapophyses of the mid-thoracic vertebrae extend beyond the cranial articular surface. The thoracic ribs are joint to the thoracic vertebrae. The two coracoids, the furcula, and three sternal ossifications are preserved. The furcula is Y-shapped. Both humeri, ulnae, and radii are also preserved.

Reconstruction of MPCM-LH-26189 by Raúl Martín

The osteohistological analysis of the left humerus shows a dense pattern of longitudinal grooves. Those grooves correspond to primary cavities, which open onto the surface of the cortex in young and fast-growing bone. The shaft of the tibia and radius show very-thin cortices. In addition,  the primary nature of the vascularisation, the round shape of the osteocytes lacunae and the uneven peripheral margin of the medullary cavity (with no endosteal bone), strongly suggests that the bone was actively growing when the bird died.

Enantiornithines show a mosaic of characters, reflecting their intermediate phylogenetic position between the basal-pygostylians and modern bird. In this clade, the sternum adopts an elaborate morphology, and in adult Enantiornithes, no more than eight free caudal vertebrae precede the pygostyle. The differences observed in the ossification of the sternum and the number of free caudal vertebrae in MPCM-LH-26189, when it compared to other juvenile enantiornithines, reveal a clade-wide asynchrony in the sequence of ossification of the sternum and tail, suggesting that the developmental strategies of these basal birds may have been more diverse than previously thought.


Fabien Knoll, et al., “A diminutive perinate European Enantiornithes reveals an asynchronous ossification pattern in early birds,” Nature Communications, volume 9, Article number: 937 (2018) doi:10.1038/s41467-018-03295-9

Chiappe, L. M., Ji, S. & Ji, Q. Juvenile birds from the Early Cretaceous of China: implications for enantiornithine ontogeny. Am. Mus. Novit. 3594, 1–46 (2007).



The oldest Archaeopteryx

Overview of the skeleton of the new Archaeopteryx specimen (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. On August 15, 1861, German paleontologist Hermann von Meyer wrote a letter to the editor of the journal Neues Jahrbuch für Mineralogie, Geologie und Palaeontologie, where he made the first description of the fossil. On on September 30, 1861, he wrote a new letter:  “I have inspected the feather from Solenhofen closely from all directions, and that I have come to the conclusion that this is a veritable fossilisation in the lithographic stone that fully corresponds with a birds’ feather. I heard from Mr. Obergerichtsrath Witte, that the almost complete skeleton of a feather-clad animals had been found in the lithographic stone. It is reported to show many differences with living birds. I will publish a report of the feather I inspected, along with a detailed illustration. As a denomination for the animal I consider Archaeopteryx lithographica to be a fitting name”. 

The near complete fossil skeleton found in a Langenaltheim quarry near Solnhofen – with clear impressions of wing and tail feathers –  was examined by Andreas Wagner, director of the Paleontology Collection of the State of Bavaria in Germany. He reached the conclusion that the fossil was a reptile, and gave it the name Griphosaurus. He wrote: “Darwin and his adherents will probably employ the new discovery as an exceedingly welcome occurrence for the justification of their strange views upon the transformations of animals.” In later editions of The Origin of Species, Darwin indeed mention the Archaeopteryx“That strange bird, Archaeopteryx, with a long lizardlike tail, bearing a pair of feathers on each joint, and with its wings furnished with two free claws . . . Hardly any recent discovery shows more forcibly than this, how little we as yet know of the former inhabitants of the world.”

Archaeopteryx lithographica, Archaeopterygidae, Replica of the London specimen; Staatliches Museum für Naturkunde Karlsruhe, Germany. From Wikimedia Commons

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 new specimen was discovered by a private collector. Although it was registered as German national cultural heritage, which guarantees its permanent availability, the specimen remains in private hands (Datenbank National Wertvollen Kulturgutes number DNWK 02924).

The new specimen is preserved as a largely articulated skeleton. However, 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.

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

The postcranial skeleton was affected by breakage and loss of elements prior to or at the time of discovery. The sacral region, and the caudal vertebrae are very poorly preserved. Several dorsal ribs are preserved, and gastralia ribs are present in the thoracic region and the abdominal region. The shoulder girdle comprises the scapula, coracoid and furcula. Both humeri are poorly preserved. Parts of the phalanges are, largely poorly, preserved, and they do not show much detail. The pubic shafts are slender and very slightly flexed posteroventrally, an as in all specimens of Archaeopteryx, the distal end of the ischium is bifurcated. The femora are also poorly preserved and largely collapsed. Both tibiae are preserved in articulation with the fibulae and the proximal tarsals. The digits of the feet are complete on both sides, but partially overlapping.

Until recently, the referral of new specimens from the Solnhofen Archipelago to the genus Archaeopteryx, seems unproblematic, but the re-examination of the fourth (Haarlem) specimen of Archaeopteryx, and the discovery in the last years of specimens from the Late Jurassic of China that are similar to Archaeopteryx, raises the question if the specimens referred to Archaeopteryx represent a monophyletic taxon.



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

Foth C, Rauhut OWM. 2017. Re-evaluation of the Haarlem Archaeopteryx and the radiation of maniraptoran theropod dinosaurs. BMC Evolutionary Biology 17:236

MEYER v., H. (1861): Archaeopterix lithographica (Vogel-Feder) und Pterodactylus von Solenhofen. Neues Jahrbuch fur Mineralogie, Geognosie, Geologie und Petrefakten-Kunde. 6: 678-679

Wellnhofer Peter, A short history of research on Archaeopteryx and its relationship with dinosaurs, Geological Society, London, Special Publications, 343:237-250, doi:10.1144/SP343.14, 2010

Introducing Caihong juji

Caihong juji holotype specimen (Hu, et al., 2018)

Over the last 10 years, theropod dinosaurs from the Middle-Late Jurassic Yanliao Biota have offered rare glimpses of the early paravian evolution and particularly the origin of birds. The first discovered Yanliao non-scansoriopterygid theropod was Anchiornis huxleyi, and since then several other extremely similar species have also been reported. Caihong juji, a newly discovered Yanliao specimen, exhibits an array of osteological features, plumage characteristics, and putative melanosome morphologies not previously seen in other Paraves. The name Caihong is from the Mandarin ‘Caihong’ (rainbow). The specific name, juji is from the Mandarin ‘ju’ (big) and ‘ji’ (crest), referring to the animal’s prominent lacrimal crests.

The holotype (PMoL-B00175) is a small, articulated skeleton with fossilized soft tissues, preserved in slab and counter slab, collected by a local farmer from Qinglong County, Hebei Province, China, and acquired by the Paleontological Museum of Liaoning in February, 2014. The specimen (estimated to be ~400 mm in total skeletal body length with a body mass of ~475 g) exhibits the following autapomorphies within Paraves: accessory fenestra posteroventral to promaxillary fenestra, lacrimal with prominent dorsolaterally oriented crests, robust dentary with anterior tip dorsoventrally deeper than its midsection and short ilium.

Caihong juji differs from Anchiornis huxleyi in having a shallow skull with a long snout, forelimb proportionally short, and forearm proportionally long. Caihong also resembles basal troodontids and to a lesser degree basal dromaeosaurids in dental features (anterior teeth are slender and closely packed, but middle and posterior teeth are more stout and sparsely spaced; and serrations are absent in the premaxilla and anterior maxilla).

Platelet-like nanostructures in Caihong juji and melanosomes in iridescent extant feathers (Hu, et al., 2018)

Feathers are well preserved over the body, but in some cases, they are too densely preserved to display both gross and fine morphological features. The contour feathers are proportionally longer than those of other known non-avialan theropods. The tail feathers resemble those of Archaeopteryx, and the troodontid Jinfengopteryx in having large rectrices attaching to either side of the caudal series forming a frond-shaped tail, a feature that has been suggested to represent a synapomorphy for the Avialae.

But, the most remarkable feature observed in Caihong, is the presence of some nanostructures preserved in the head, chest, and parts of its tail, that have been identified as melanosomes. They are long, flat, and organized into sheets, with a pattern similar of those of the iridescent throat feathers of hummingbirds.

Recovered as a basal deinonychosaur, Caihong shows the earliest asymmetrical feathers and proportionally long forearms in the theropod fossil record wich indicates locomotor differences among closely related Jurassic paravians and has implications for understanding the evolution of flight-related features.


Hu, et al. A bony-crested Jurassic dinosaur with evidence of iridescent plumage highlights complexity in early paravian evolution. Nature (2018) doi:10.1038/s41467-017-02515-y

Godefroit, P. et al. A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds. Nature 498, 359–362 (2013).

The last terror birds

Skeleton of the terror bird Titanis walleri at the Florida Museum of Natural History.

In 1887, Florentino Ameghino, the “father of Argentinian Palaeontology”, described a large, toothless jaw from the Miocene of the Province of Santa Cruz, naming it Phorusrhacos longissimus and assigning it to a new family of edentulous mammal. He used this finding as a critical evidence for his contention that modern mammalian lineages originated in Argentina and later spread across the globe. Four years later, Moreno and Mercerat recognized for the first time that the mandible described by Ameghino was really that of a bird.

The Phorusrhacidae, the so-called “terror birds”, were a group of medium-to large sized extinct ground-dwelling birds, which lived during the Cenozoic, and became the dominant carnivores of South America while it was an isolated continent. They are characterized by their elongated hindlimbs, narrow pelvis, reduced forelimbs, and their huge skull with their tall, long, narrow, and hollow beaks ended in a hook. Kelenken guillermoi, is the largest known phorusrhacid and lived in the Miocene of Argentina. The skull reaches a length of 71.6 cm and the whole animal would reach 3 m high. Kelenken is also represented by a tarsometatarsus and a broken phalanx and proceeds from the locality of Comallo (Río Negro Province, Argentina). Titanis walleri, lived during the Pliocene and Pleistocene of North America. It was 2.5 metres tall and weighed approximately 150 kilograms. This giant bird is interpreted as an early immigrant during the Great American Interchange.

Proximal portion of the left humerus of Psilopterus sp. Caudal, b ventral, c cranial and d dorsal views (From Jones et. al., 2017)

At the end of the Pliocene, Phorusrhacids decline in diversity. Two new specimens support the hypothesis that the latest geologic occurrence of the Phorusrhacidae comes from late Pleistocene sediments of Uruguay. The remains comprise the distal portion of right tarsometatarsus and a left humerus; the latter is assigned to the genus Psilopterus. The first material (MPAB-520) comes from Soriano, Uruguay, and the sediments belong to the Dolores Formation (Lujanian Stage-Age, late Pleistocene/early Holocene). The following features identify the specimen as a phorusrhacid bird. (1) a large and distally expanded trochlea metatarsi III; (2) a very narrow trochlea metatarsi II with the articular surface transversally convex and without any longitudinal sulcus (in dorsal and distal views); (3) in dorsal view the trochlea metatarsi II is almost parallel and much shorter than the middle trochlea, and forming a narrow notch between trochleae II and III. The second material consists of a left humerus without distal epiphysis belonged to Museo Paleontológico Alejandro Berro (MPAB-2024).

There are two explanatory hypotheses proposed for the decline of the terror birds: environmental reasons or direct competition (at least for the larger specimens) with placental carnivore’s immigrants to South America after the setting of the Panamanian bridge. 



Jones, W., Rinderknecht, A., Alvarenga, H. et al. PalZ (2017), The last terror birds (Aves, Phorusrhacidae): new evidence from the late Pleistocene of Uruguay,

ALVARENGA, Herculano M.F.  and  HOFLING, Elizabeth. Systematic revision of the Phorusrhacidae (Aves: Ralliformes). Pap. Avulsos Zool. (São Paulo) [online]. 2003, vol.43, n.4 [cited  2015-03-24], pp. 55-91 .

Top fossils discoveries of 2017


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

On April 22, 2017, the March for Science reunited more than a million persons worldwide to protest against the attack on science, budget cuts and censorship. A fight that doesn’t over yet. But despite all that, 2017 was also an extraordinary year for palaeontological discoveries. My top list includes:

  • Borealopelta

Holotype of Borealopelta markmitchelli

Borealopelta markmitchelli, from the Early Cretaceous of Alberta, is a three-dimensionally preserved ankylosaurian,  discovered in the Suncor Millennium Mine in Canada. The generic name Borealopelta is derived from “borealis” (Latin, “northern”) and “pelta” (Greek, “shield”). 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.

  • Isaberrysaura


Isaberrysaura skull in lateral view and maxillary teeth (Adapted from Salgado et al., 2017)


Isaberrysaura mollensis gen. et sp. nov. is the first dinosaur recovered in the marine-deltaic deposits of the Los Molles Formation (Neuquén Province, Argentina), and the first neornithischian dinosaur known from the Jurassic of South America. The holotype of Isaberrysaura is an incomplete articulated skeleton with an almost complete skull, and a partial postcranium consisting of 6 cervical vertebrae, 15 dorsal vertebrae, a sacrum with a partial ilium and an apparently complete pubis, 9 caudal vertebrae, part of a scapula, ribs, and unidentifiable fragments. One of the most notable features of the discovery is the presence of permineralized seeds in the middle-posterior part of the thoracic cavity. The seeds were assigned to the Cycadales (Zamiineae) on the basis of a well-defined coronula in the micropylar region. The findings suggest the hypothesis of interactions (endozoochory) between cycads and dinosaurs, especially in the dispersion of seeds.

  •  Junornis 


Holotype of Junornis houi. From Chiappe et. al; 2017)

Junornis houi, from the Yixian Formation of eastern Inner Mongolia, represents a new addition to the enantiornithine diversity of the Jehol BiotaThe holotype (BMNHC-PH 919; Beijing Museum of Natural History), from the Early Cretaceous (~ 126±4 mya) of Yixian Formation,  is a nearly complete and articulated skeleton contained in two slabs, and surrounded by feather impressions defining the surface of its wings and body outline. Based on the well-preserved skeleton and exquisite plumage of Junornis, it was possible  make some estimation of its flight capacity. The body and wings of this bird were similar to those of modern passeriforms such as Alauda arvensis. 

  • Shringasaurus

Cranial anatomy of Shringasaurus indicus

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.

  • Patagotitan 

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.



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

Salgado, L. et al. A new primitive Neornithischian dinosaur from the Jurassic of Patagonia with gut contents. Sci. Rep. 7, 42778; doi: 10.1038/srep42778 (2017)

Liu D, Chiappe LM, Serrano F, Habib M, Zhang Y, Meng Q (2017) Flight aerodynamics in enantiornithines: Information from a new Chinese Early Cretaceous bird. PLoS ONE12(10): e0184637.

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

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

The bizarre Halszkaraptor escuilliei

H. escuilliei MPC D-102/109. From Cau et al., 2017.

Maniraptoran lineages evolved novel ecomorphologies during the Cretaceous period, including active flight, gigantism, cursoriality and herbivory. This group share the following characteristics: large brain but a reduced skull in comparison to their body size, beaks, and smaller teeth. Now, a well-preserved maniraptoran from Mongolia, revealed a mosaic of features, most of them absent among non-avian maniraptorans but shared by reptilian and avian groups with aquatic or semiaquatic ecologies. This new theropod, Halszkaraptor escuilliei gen. et sp. nov., adds an amphibious ecomorphology to those evolved by maniraptorans.

The holotype, MPC (Institute of Paleontology and Geology, Mongolian Academy of Sciences, Ulaanbaatar, Mongolia) D-102/109, is an articulated and almost complete skeleton preserved three-dimensionally. The generic name, honours Halszka Osmólska (1930–2008) for her contributions to theropod palaeontology. The species name, ‘escuilliei’ refers to François Escuillié, who returned the holotype to Mongolia.

Reconstruction of Halszkaraptor escuilliei. Photograph: Lukas Panzarin/Andrea Cau

Halszkaraptor is related to other enigmatic Late Cretaceous maniraptorans from Mongolia in a novel clade at the root of Dromaeosauridae. It was the size of a mallard. Originally poached from Ukhaa Tolgod, Mongolia, the fossil was in private collections in Japan and England for an unknown amount of time, and later it  was transferred to the Royal Belgian Institute of Natural Sciences (RBINS). Thanks to a cooperation agreement between the Ministry of Education, Culture and Science of Mongolia, the Belgian Science Policy Office and the RBINS, the specimen returned to the Institute of Paleontology and Geology, Mongolian Academy of Science.

The skeleton is almost complete. The skull is lightly built, and is still articulated with the first cervical vertebra. The preorbital region forms 60% of basicranial length, and each premaxilla is elongate, bearing eleven teeth, the highest number among dinosaurs. The presacral vertebrae include 10 cervicals and 12 dorsals. The neck forms 50% of snout–sacrum length.

Skull of H. escuilliei. From Cau et al., 2017

The forelimb is relatively shorter than in most dromaeosaurids. The ulna is flattened and possesses an acute posterior margin. The hand has a morphology that is unique among theropods, with a progressive elongation of the lateral fingers, with the third being the longest and most robust. The 76 mm long femur has a robust greater trochanter. The metatarsus lacks cursorial adaptations and measures 80% of femoral length. The feet are complete and articulated, although some elements are poorly visible.

Based on the neck hyperelongation for food procurement, the forelimb proportions that may support a swimming function, and postural adaptations convergent with short-tailed birds, Halszkaraptor may represent the first case among non-avian dinosaurs of a double locomotory module.


Cau, A.; Beyrand, V.; Voeten, D.; Fernandez, V.; Tafforeau, P.; Stein, K.; Barsbold, R.; Tsogtbaatar, K.; Currie, P.; Godrfroit, P.; “Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs”. Nature. doi:10.1038/nature2467

A brief history of Mesozoic theropods research in Gondwana

Snout of the ceratosaurian Genyodectes serus

In the last decades, the study of Gondwanan non-avian theropods has been highly prolific, showing that the group reached a great taxonomic and morphological diversity comparable to that of Laurasia. The Mesozoic Gondwanan neotheropod record includes: coelophysoids, basal averostrans, ceratosaurids, abelisauroids, megalosauroids, carcharodontosaurids, megaraptorans, basal coelurosaurs, compsognathids, alvarezsauroids, unenlagiids, and basal avialans, as well as putative tyrannosauroids, ornithomimosaur-like forms, and troodontid. Therefore, the Gondwanan fossil record has been crucial to understand the evolution and global biogeography of dinosaurs during the Mesozoic.

The first probable theropod remains from Gondwana were discovered in Colombia by Carl Degenhard, a German engineer, in 1839. At that time the word “dinosaur” did not even exist yet. Although Degenhard identified them as bird footprints, his brief description suggests that they were tracks of bipedal dinosaurs. But it was not until 1896 that the first Gondwanan theropod was named by the French palaeontologist Charles Depéret as “Megalosaurus” crenatissimus from the Upper Cretaceous of Madagascar. Several theropod remains were described from India, Africa, and South America during the 19th century. These early fragmentary discoveries lead the authors of the late XIX and early XX centuries to interpret them as belonging the same lineages present in Europe and North America.

Elaphrosaurus bambergi (Museum für Naturkunde 4960, holotype) from the Upper Jurassic of Tanzania (Janensch, 1920)

In 1901, A. Smith Woodward described Genyodectes, based on fragmentary skull bones, including portions of both maxillas, premaxillae,  parts of the supradentaries, and some teeth, discovered by Santiago Roth in Chubut, at the end of the 1880s. Genyodectes remained as the most completely known  theropod from South American until the 1970s. In 2004, O. Rauhut concluded that Genyodectes is more closely related to Ceratosaurus than the more derived abelisaurs.

Between 1915 and 1933, the most relevant Gondwanan theropod discoveries were produced by the work of the German palaeontologists Frederich von Huene, Ernst Stromer, and Werner Janensch, including for the first time the publication of very informative partial skeletons, such as those of Spinosaurus aegyptiacus and Elaphrosaurus bambergi (Stromer, 1915; Janensch, 1920). 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. But during and after the Second World War the influence of the German palaeontology in the research of Gondwanan theropods abruptly declined.

Skull and neck of Carnotaurus sastrei

By the 1960s, the Argentine biologist Osvaldo Reig, together with Rodolfo Casamiquela and José Bonaparte, began to explore the Mesozoic rocks of Argentina looking for fossil tetrapods. In 1985, Bonaparte published a note presenting Carnotaurus sastrei as a new genus and species and briefly describing the skull and lower jaw. It was collected in the lower section of La Colonia Formation, Chubut Province. The discoveries of Bonaparte and his collaborators resulted in the recognition of the Patagonian theropod record as the most relevant and informative among Gondwanan continents. Some of the theropod species discovered in Patagonia are known on the basis of skulls and fairly complete skeletons offering insights into the anatomy and phylogeny of abelisaurids, carcharodontosaurids, and maniraptorans.


Martín D. Ezcurra, and Federico L. Agnolín (2017). Gondwanan perspectives: Theropod dinosaurs from western Gondwana. A brief historical overview on the research of Mesozoic theropods in Gondwana. Ameghiniana 54: 483–487. Published By: Asociación Paleontológica Argentina

Novas, F.E., et al., Evolution of the carnivorous dinosaurs during the Cretaceous: The evidence from Patagonia, Cretaceous Research (2013), 

Buffetaut, E. 2000A forgotten episode in the history of dinosaur ichnology; Carl Degenhardt’s report on the first discovery of fossil footprints in South America (Colombia, 1839). Bulletin de la Société Géologique de France 171: 137140Google Scholar