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.

 

References:

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

Foth C, Rauhut OWM. 2017. Re-evaluation of the Haarlem Archaeopteryx and the radiation of maniraptoran theropod dinosaurs. BMC Evolutionary Biology 17:236 https://doi.org/10.1186/s12862-017-1076-y

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

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Our once and future oceans

Earth is the only planet in our Solar System with high concentrations of gaseous diatomic oxygen. Simultaneously, this unique feature of Earth’s atmosphere has allowed the presence of an ozone layer that absorbed UV radiations. The progressive oxygenation of the atmosphere and oceans was sustained by an event of high organic carbon burial, called the Paleoproterozoic Lomagundi Event (ca. 2.3-2.1 billion years ago), which lasted well over 100 million years.

Oxygen is fundamental to life, and influences biogeochemical processes at their most fundamental level. But the oxygen content of Earth has varied greatly through time. In Earth history there have been relatively brief intervals when a very significant expansion of low-oxygen regions occurred throughout the world’s oceans. The discovery of black shales at many drill sites from the Atlantic, Indian, and the Pacific Ocean led to the recognition of widespread anoxic conditions in the global ocean spanning limited stratigraphic horizons. In 1976, S. O. Schlanger and H. C. Jenkyns termed these widespread depositional black shale intervals as “Oceanic Anoxic Events”. This was one of the greatest achievement of the DSDP (Deep Sea Drilling Project).

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

Human activity is a major driver of the dynamics of Earth system. After the World War II, the impact of human activity on the global environment dramatically increased. Over the past 50 years, open ocean lost an estimated 2%, or 4.8 ±2.1 petamoles (77 billion metric tons), of its oxygen, and ocean oxygen minimum zones (OMZs) have expanded by an area about the size of the European Union. Deoxygenation is linked to other ocean stressors, including warming and acidification.

Ocean warming reduces the solubility of oxygen, and raises metabolic rates accelerating the rate of oxygen consumption. Warming also influence on thermal stratification and indirectly enhances salinity driven stratification through its effects on ice melt and precipitation. The increased stratification alters the mainly wind-driven circulation in the upper few hundred meters of the ocean and slows the deep overturning circulation. Intensified stratification may account for the remaining 85% of global ocean oxygen loss by reducing ventilation nd by affecting the supply of nutrients controlling production of organic matter and its subsequent sinking out of the surface ocean. Warming is predicted to exacerbate oxygen depletion in coastal systems through mechanisms similar to those of the open ocean.

Time scale [Gradstein et al., 2005] illustrating the stratigraphic position and nomenclature of OAEs (From Jenkyns, 2010).

The geological records show that large and rapid global warming events occurred repeatedly during the course of Earth history. The growing concern about modern climate change has accentuated interest in understanding the causes and consequences of these ancient abrupt warming events. The early Toarcian Oceanic Anoxic Event  (T-OAE; ∼183 mya) in the Jurassic Period is associated with a major negative carbon isotope excursion, mass extinction, marine transgression and global warming. Besides, the marked expansion of the oxygen minimum zone over the last decades, is quite similar to the model originally invoked for the genesis of Cretaceous OAEs. The better understanding of the Mesozoic ocean-climate system and the formation of OAEs would help us to predict environmental and biotic changes in a future greenhouse world.

References:

DENISE BREITBURG, LISA A. LEVIN, ANDREAS OSCHLIES, MARILAURE GRÉGOIRE, FRANCISCO P. CHAVEZ, DANIEL J. CONLEY, VÉRONIQUE GARÇON, DENIS GILBERT, DIMITRI GUTIÉRREZ, KIRSTEN ISENSEE, GIL S. JACINTO, KARIN E. LIMBURG, IVONNE MONTES, S. W. A. NAQVI, GRANT C. PITCHER, NANCY N. RABALAIS, MICHAEL R. ROMAN, KENNETH A. ROSE, BRAD A. SEIBEL, MACIEJ TELSZEWSKI, MORIAKI YASUHARA, JING ZHANG (2018), Declining oxygen in the global ocean and coastal waters, Science, Vol. 359, Issue 6371, DOI: 10.1126/science.aam7240

Jenkyns, H. C. (2010), Geochemistry of oceanic anoxic eventsGeochem. Geophys. Geosyst.11, Q03004, doi:10.1029/2009GC002788.

Holz, M., Mesozoic paleogeography and paleoclimates – a discussion of the diverse greenhouse and hothouse conditions of an alien world, Journal of South American Earth Sciences (2015), doi: 10.1016/j.jsames.2015.01.001

Tennant, J. P., Mannion, P. D., Upchurch, P., Sutton, M. D. and Price, G. D. (2016), Biotic and environmental dynamics through the Late Jurassic–Early Cretaceous transition: evidence for protracted faunal and ecological turnover. Biol Rev. doi:10.1111/brv.12255 

 

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.

References:

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. 

 

References:

Jones, W., Rinderknecht, A., Alvarenga, H. et al. PalZ (2017), The last terror birds (Aves, Phorusrhacidae): new evidence from the late Pleistocene of Uruguay, https://doi.org/10.1007/s12542-017-0388-y

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.

 

References:

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. https://doi.org/10.1371/journal.pone.0184637

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

Pisanosaurus and the Triassic ornithischian crisis

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

In 1887, Harry Govier Seeley was the first to subdivide dinosaurs into Saurischians and the Ornithischians based on the nature of their pelvic bones and joints. While the clade Saurischia is well represented in the Late Triassic, the record of the Ornithischia is certainly more problematic. Only a single Triassic ornithischian taxon was generally considered to still be valid: Pisanosaurus mertii, originally described by Argentinian paleontologist Rodolfo Casamiquela in 1967, based on a poorly preserved but articulated skeleton from the upper levels of the Ischigualasto Formation (Late Triassic).

The holotype and only known specimen (PVL 2577) is a fragmentary skeleton including partial upper and lower jaws, seven articulated dorsal vertebrae, four fragmentary vertebrae of uncertain position in the column; the impression of the central portion of the pelvis and sacrum; an articulated partial hind limb including the right tibia; fibula; proximal tarsals and pedal digits III and IV; the distal ends of the right and left femora; a left scapular blade (currently lost); a probable metacarpal III;  and the impressions of some metacarpals (currently lost).

Reconstructed skeleton reflecting the traditional interpretation of Pisanosaurus (Royal Ontario Museum)

But Pisanosaurus shows some derived traits that resulted as unambiguous synapomorphies of the Silesauridae clade, and include: reduced to absent denticles on maxillary and dentary teeth; sacral ribs shared between two sacral vertebrae; lateral side of proximal tibia with a fibular flange; dorsoventrally flattened ungual phalanges; and ankylothecodonty, teeth partially fused to maxilla and dentary bone. The inclusion of Pisanosaurus within Silesauridae implies that this taxon does not constitute the oldest ornithischian. This is consistent with previous interpretations proposing that ornithischian radiation occurred after the Triassic–Jurassic boundary.

To explain the relatively low diversity exhibited by Ornithischia in the Late Triassic-Early Jurassic, several hypotheses have been proposed. One, suggests that Ornithischia is the sister-taxon of Neotheropoda (the least inclusive clade that includes Coelophysis and modern birds) within the clade of ‘traditional theropod taxa’. In this model, a ‘transitional’ ornithischian may possess some anatomical features of theropods that appear to be more like those in more derived than Eodromaeus murphi and Tawa hallae.

Hypothesis 4, in which Ornithischia forms the sister-taxon of Averostra (From Baron 2017)

In a second hypothesis, Ornithischia is positioned as the sister-taxon to the coelophysids. In this model, Neotheropoda and Ornithoscelida would encompass the same set of taxa, but Ornithoscelida would, theoretically, take priority of Neotheropoda as it is the older name. In a third hypothesis, Ornithischia is positioned as the sister-taxon to the ‘other neotheropods’ not contained in the coelophysid clade.

Another hypothesis proposes that Ornithischia forms the sister-taxon of Averostra. Like Ornithischia, Averostra is only known from the Jurassic and Cretaceous Periods, and both share a number of anatomical features, such as fusion of the sacral neural. Another anatomical traits that could unite such a group include the possession of six or more sacral vertebrae; and the fusion of the sacral neural spines into a broad and continuous sheet, as in early ornithischians like Lesothosaurus diagnosticus and tetanuran theropods like Megalosaurus bucklandii. It’s worth mentioning the fact the earliest known unambiguous members of both Ornithischia and Averostra, are found in the same formation in South America: Laquintasaura venezuelae and Tachiraptor admirabilis.

Laquintasaura venezuelae gen. et sp. nov (From Barret et al., 2014)

 

It was suggested (Baron and Barrett 2017) that Chilesaurus diegosaurezi from the Late Jurassic, might represent the earliest diverging member of Ornithischia. Chilesaurus 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.

The Phytodinosauria hypothesis suggest that Ornithischia is nested among the taxa traditionally termed as sauropodomorphs could also offer a solution to the problem of the lack of unambiguous ornithischians in the Carnian and Late Triassic in general.

 

References:

Baron, M. G. (2017): Pisanosaurus mertii and the Triassic ornithischian crisis: could phylogeny offer a solution?, Historical Biology, DOI: 10.1080/08912963.2017.1410705

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

Baron M. G, Barrett P. M. 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

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

Barrett, Paul M.; Butler, Richard J.; Mundil, Roland; Scheyer, Torsten M.; Irmis, Randall B.; Sánchez-Villagra, Marcelo R. (2014). A palaeoequatorial ornithischian and new constraints on early dinosaur diversification, Proceedings of the Royal Society B, DOI: 10.1098/rspb.2014.1147

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

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

 

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.

References:

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.

References:

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 https://doi.org/10.5710/102.054.0501

Novas, F.E., et al., Evolution of the carnivorous dinosaurs during the Cretaceous: The evidence from Patagonia, Cretaceous Research (2013), http://dx.doi.org/10.1016/j.cretres.2013.04.001 

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

 

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”‘

References:

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.

 

References:

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.