Marine ecosystems have entered the Anthropocene

Sampling of foraminifera found in a sediment core from the Caribbean, dating back to before the Industrial Revolution. CREDIT MICHAL KUCERA

Anthropogenic climate change and ocean acidification resulting from the emission of vast quantities of CO2 and other greenhouse gases pose a considerable threat to ecosystems and modern society. Planktonic foraminifera are a group of marine zooplankton that made their first appearance in the Late Triassic. Although, identifying the first occurrence of planktonic foraminifera is complex, with many suggested planktonic forms later being reinterpreted as benthic. They are present in different types of marine sediments, such as carbonates or limestones, and are excellent biostratigraphic markers. Their test are made of  globular chambers composed of secrete calcite or aragonite, with no internal structures and different patterns of chamber disposition: trochospiral, involute trochospiral and planispiral growth. During the Cenozoic, some forms exhibited supplementary apertures or areal apertures. The tests also show perforations and a variety of surface ornamentations like cones, short ridges or spines. The phylogenetic evolution of planktonic foraminifera are closely associated with global and regional changes in climate and oceanography.

John Murray, naturalist of the CHALLENGER Expedition (1872-1876) found that differences in species composition of planktonic foraminifera from ocean sediments contain clues about the temperatures in which they lived. The ratio of heavy and light Oxygen in foraminifera shells can reveal how cold the ocean was and how much ice existed at the time the shell formed. Another tool to reconstruct paleotemperatures is the ratio of magnesium to calcium (Mg/Ca) in foraminiferal shells. Mg2+ incorporation into foraminiferal calcite  is influenced by the temperature of the surrounding seawater, and the Mg/Ca ratios increase with increasing temperature.

Planktonic foraminifera from the Sargasso Sea in the North Atlantic Ocean. (Photograph courtesy Colomban de Vargas, EPPO/SBRoscoff.)

Analyzing previously collected sediment samples from over 3,500 sites around the world’s ocean, researchers found that the composition of the planktonic foraminifera has changed significantly since the pre-industrial period. The shifts in planktonic foraminifera are indicative of a more-general phenomenon across marine ecosystem, with zooplankton communities shifting poleward by an average 374 miles as a result of warming ocean temperatures.

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. Ocean warming reduces the solubility of oxygen, and raises metabolic rates accelerating the thermal stratification.


Jonkers, L., Hillebrand, H., & Kucera, M. (2019). Global change drives modern plankton communities away from the pre-industrial state. Nature. doi:10.1038/s41586-019-1230-3


Forgotten women of paleontology: Hildegarde Howard

Hildegard Howard with fossil bird from the Rancho La Brea.

The birth of modern science was hostile to women’s participation. The world’s major academies of science were founded in the 17th century: the Royal Society of London (1662), the Paris Académie Royale des Sciences (1666), and the Berlin Akademie der Wissenschaften (1700). Unfortunately, women were not become members of these societies for over 300 years. Yvonne Choquet-Bruhat became the first woman to be elected to the Paris Academy of Science in 1979. Although the Royal Society was less rigid in terms of memberships than the Paris Academy of Science, it was not until 1945 that the first women were admitted as fellows of the Royal Society: the X-ray crystallographer Kathleen Yardley Lonsdale (1903–1971), and biochemist and microbiologist Marjory Stephenson (1885-1948).

Despite the barriers, between 1880 and 1914, some 60 women contributed papers to Royal Society publications. Meanwhile, in the United States, geology was a marginal subject in the curricula of the early women’s colleges until an intense programme was started at Bryn Mawr College in the 1890s.

Hildegard Howard measures specimens from the Rancho La Brea Collection. Image from The Natural History Museum of Los Angeles County Archives.

Florence Bascom was one of the pioneers when geological education at universities became available to women. She received her PhD degree from Johns Hopkins University in 1893 by special dispensation, as women were not admitted officially until 1907; while Carlotta Joaquina Maury attended Cornell University, where she became one of the first women to receive her PhD in paleontology in 1902.

When Hildegarde Howard began attending the Southern Branch of the University of California (now known as the University of California at Los Angeles), women were still barred from scientific societies. She was born on April 3, 1901 in Washington D.C., but moved to Los Angeles at the age of 5. Her main interest was journalism, until she met her first biology instructor, Miss Pirie Davidson. In 1921, Hildegarde obtained a part-time job working for Dr. Chester Stock, sorting bones from Rancho La Brea in the basement of the Los Angeles Museum of History, Science and Art (now known as the Natural History Museum of Los Angeles County). One year later, she went to Berkeley to finish her degree.

Dr. Hildegarde Howard, in her office in 1961.Copyright Natural History Museum of Los Angeles County

In 1928, she obtained her Ph.D. degree. Her dissertation, entitled “The Avifauna of Emeryville Shellmound”, became one of her most popular works, and remained as the principal reference of its kind until the appearance of the first edition of Nomina Anatomica Avium in 1979. She obtained a permanent position with the museum in 1929. Although she was a curator, she did not receive that official title until 1938. Through that decade, she wrote twenty-four papers on fossil birds in the American Southwest. She was promoted to the curator of Avian Paleontology in 1944, and she would serve in that role until 1951, when she was promoted to Chief Curator of Science, She became the first woman to receive the Brewster Medal for outstanding research in ornithology in 1953.

Hildegarde Howard officially retired in 1961, although continued research on fossil birds, publishing her last paper in 1992. During her extraordinary career, Dr. Howard described 3 families, 13 genera, 57 species, and 2 subspecies, and remains highly regarded as one of the foremost experts in her field. She died on February 28, 1998.



Campbell Jr., Kenneth. 2000c. “In Memoriam, Hildegarde Howard 1901-1998.” The Auk, vol.117, no.3, 775-779.


Introducing Kaijutitan, the strange beast.

The entrance to the town of Rincón de los Sauces.

Since the discovery of dinosaur remains in the Neuquen basin in 1882, Argentina has gained the title of Land of the Giants. The tittle was reinforced by the discoveries of titanosaurs like Argentinosaurus, Dreadnoughtus, Notocolossus, Puertasaurus, and Patagotitan. The study of this diverse group of sauropod dinosaurs embrace an extensive list of important contributions, which started with Richard Lydekker’s pioneering work on Patagonian dinosaurs, and by the classic Friedrich von Huene monograph on Argentinean saurischians and ornithischians.

Titanosaurus were a diverse group of sauropod dinosaurs represented by more than 30 genera, which included all descendants of the more recent common ancestor of Andesaurus and Saltasaurus. The group includes the smallest (e.g. Rinconsaurus, Saltasaurus; with estimated body masses of approximately 6 tonnes) and the largest sauropods known to date. They had their major radiation during the middle Early Cretaceous. The evolution of body mass in this clade is key element to understand sauropod evolution.


Cranial elements of MAU-Pv-CM-522/1. From Filippi et al., 2019.

Kaijutitan maui, is the first basal sauropod titanosaur from the Sierra Barrosa Formation (Upper Coniacian, Upper Cretaceous). The holotype (MAU-Pv-CM-522) consists of cranial, axial, and appendicular elements presenting an unique combination of plesiomorphic and apomorphic characters. The generic name Kaijutitan is derived from Kaiju, Japanese word that means “strange beast” or “monster”, and titan, from the Greek “giant”.  The species name refers to the acronym of the Museo Municipal Argentino Urquiza, Rincon de los Sauces, Neuquén, Argentina.

The cranial elements of this specimen include the complete neurocranium (the supraoccipital, exoccipital, left paraoccipital process, left exoccipital-opisthotic-prootic complex, left laterosphenoid and orbitosphoid, and basioccipital-basisphenoid complex). The impossibility of recognizing clear sutures suggest an ontogenetic adult stage of the specimen. One of the most notable autapomorphies exhibited by Kaijutitan is the anterior cervical vertebra with bifid neural spine, a feature usually found in diplodocids and dicraeosaurids. Unfortunately, the femur and humerus of Kaijutitan maui are incomplete, therefore the body mass of this titanosaur can only be estimated by comparison with other titanosauriforms. Kaijutitan would have had a body mass similar or intermediate to that of Giraffatitan (38.000 kg) and Notocolossus (60.398 kg).



Filippi, L.S., Salgado, L., Garrido, A.C., A new giant basal titanosaur sauropod in the Upper Cretaceous (Coniacian) of the Neuquén Basin, Argentina, Cretaceous Research,

Meet Iberodactylus.

Partial rostrum of Iberodactylus andreui. From Holgado et. al, 2019

Pterosaurs were the first flying vertebrates. The group achieved high levels of morphologic and taxonomic diversity during the Mesozoic, with more than 200 species recognized so far. From the Late Triassic to the end of the Cretaceous, the evolution of pterosaurs resulted in a variety of eco-morphological adaptations, as evidenced by differences in skull shape, dentition, neck length, tail length and wing span. Because of the fragile nature of their skeletons the fossil record of pterosaurs is strongly biased towards marine and lacustrine depositional environments.

Pterosaurs have been divided into two major groups: “rhamphorhynchoids” and “pterodactyloids”. Rhamphorhynchoids are characterized by a long tail, and short neck and metacarpus. Pterodactyloids have a much larger body size range, an elongated neck and metacarpus, and a relatively short tail, and ruled the sky from the Late Jurassic to the End Cretaceous.

Comparison of the rostrum of Iberodactylus andreui with a cast of a skull of Hamipterus tianshanensis. From Holgado et al., 2019

The record of Iberian pterosaurs is scarce, but a new taxa from the Lower Cretaceous of Spain reveals an unexpected relationship with Hamipterus tianshanensis from the Lower Cretaceous of China. Iberodactylus andreui gen. et sp. nov., was recovered at Los Quiñones site, close to the village of Obón (Teruel, Spain), at the end of the 1980s by Javier Andreau. The holotype (MPZ-2014/1) consists of the anterior portion of the rostrum (~198 mm in length), and includes a partially preserved premaxillary crest, and a fragment of the maxillary bone with several fragmentary teeth. The specimen preserved its original 3D shape, although exhibits frequent fractured bones, that added to the eroded bone surfaces, reveal an external thing layer of cortical bone of 1.5 mm. The robustness and height of the premaxillary crest, suggest that MPZ-2014/1 may represent a male specimen.

The most striking feature of MPZ-2014/1 is the premaxillary crest. This crest exhibits well-developed elongated, sub-vertical striae and sulci, anteriorly curved, a combination that is quite similar to Hamipterus tianshanensis from the Berriasian-Albian of China. It was suggested that the sulci could be interpreted as a trait related to the attachment of the rhamphotheca, as in the case of some extant birds.

Origin and radiation of the clade Anhangueria during the Early Cretaceous. From Holgado et al., 2019

Phylogenetic analyses indicate that Hamipterus tianshanensis and Iberodactylus andreui gen. et sp. nov. form a monophyletic group, the Hamipteridae fam. nov., that falls within the Anhangueria, sharing with other anhanguerians the presence of a lateral expansion on the rostral tips. Anhanguerians has been recorded elsewhere in the Early Cretaceous of Europe, however Iberodactylus is not closely related to any known European anhanguerian, suggesting that the clade Anhangueria could have ancestral connections to eastern Laurasia.

Other tetrapod lineages are recorded in the Iberian Peninsula with close affinities to Asian faunas. Those lineages include titanosauriforms, crocodyliforms, enanthiornitean birds, and the gobiconodontid mammal Spinolestes xenarthrosus related to Gobiconodon and Repenomamus.


Borja Holgado, Rodrigo V. Pêgas, José Ignacio Canudo, Josep Fortuny, Taissa Rodrigues, Julio Company & Alexander W.A. Kellner, 2019, “On a new crested pterodactyloid from the Early Cretaceous of the Iberian Peninsula and the radiation of the clade Anhangueria”, Scientific Reports 9: 4940

The mounting of the cast of Diplodocus carnegii at the Museo de La Plata.

Diplodocus carnegii at the Museo de La Plata, 1912 (From Otero and Gasparini, 2014)

Diplodocus carnegii at the Museo de La Plata, 1912 (From Otero and Gasparini, 2014).

Diplodocus is one of the most popular dinosaurs of all time. The first remains of a Diplodocus were found by Benjamin Mudge and Samuel Wendell Williston, in the Upper Jurassic outcrops of Cañon City, Colorado, United States, in 1877. One year later, Othniel Charles Marsh named the species Diplodocus longus on the basis of remains of the hind limb and tail. The name Diplodocus means ‘double beam’ in reference to the particular two-pronged morphology of the posterior hemal arches. D. carnegii, was discovered in 1899 during an expedition carried out by the Carnegie Museum to the Upper Jurassic Morrison Formation of Wyoming. John Bell Hatcher dedicated the new species of to Andrew Carnegie.

A sketch from the of Diplodocus carnegii, which Carnegie had framed and mounted on a wall at his castle in Scotland.

William Jacob Holland, director of the Carnegie Museum, sent a sketch of the skeleton of Diplodocus to Andrew Carnegie. At the time, the steel tycoon was at his Castle, Skibo, in Sutherland County, Scotland. The King Edward VII of England, saw the sketch and asked Carnegie to give him a specimen for the British Museum of Natural History in London. Holland proposed to Carnegie to make a life-sized replica of D. carnegii to be given to the British Museum of Natural History. On May 12, 1905, the long skeleton was unveiled to a crowd of 300 people, and became an instant star.

Mounting of the cast of Diplodocus carnegii at the Museo de La Plata, Argentina. Arthur Coggeshall and William Holland are second and third from left (Adapted from ‘Caras y Caretas’ magazine, 1912).

Nine replicas of D. carnegii were made and donated to kings and presidents of Europe and Latin America. On November of 1911, Argentinean president Dr. Roque Saenz Peña communicated to Andrew Carnegie his request to have a replica of D. carnegii. His request was accepted, and on July 1, 1912, 34 boxes containing the cast of the animal were sent to Argentina on the S.S. ‘Sallust’. William Holland and Mr. Arthur Coggeshall were in charge of mounting the replica. The site where the replica would be mounted in the Museo de La Plata, would be the Sala III, which was dedicated to invertebrates and plants. Holland insisted that the plans used for the mounting of D. carnegii at Vienna were followed in mounting the skeleton in La Plata. After the skeleton was mounted, the Director of the Museum, Dr. Samuel Lafone-Quevedo, gave a speech expressing his gratitude to Andrew Carnegie and his representatives, in which William Holland was designated an Honorary Member of La Plata University.


Alejandro Otero and Zulma Gasparini “The History of the Cast Skeleton of Diplodocus carnegii Hatcher, 1901, at the Museo De La Plata, Argentina,” Annals of Carnegie Museum 82(3), (2014). doi:

BARRETT, P., P. PARRY, AND S. CHAPMAN. 2010. Dippy: The Tale of a Museum Icon. Natural History Museum, London. 48 pp.


Introducing Moros intrepidus, the harbinger of doom.

Moros intrepidus. Credit: Jorge Gonzalez

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

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

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

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



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

Introducing Bajadasaurus pronuspinax.

Bajadasaurus reconstruction (Museo Municipal Ernesto Bachmann, Villa El Chocón, Neuquén).

Dicraeosauridae is a family of mid-sized sauropod dinosaurs characterized by a distinctive vertebral column with paired, long, neural spines. The group was first described in 1914 by Werner Janensch with the discovery of the nearly complete skeletons of Dicraeosaurus in the expeditions to the upper Jurassic beds of Tendaguru, Tanzania. Dicraeosauridae includes  Amargasaurus, Pilmatueia, Suuwassea, and Brachytrachelopan. Now, the description of Bajadasaurus pronuspinax gen. et sp. nov., from the Early Lower Cretaceous of Bajada Colorada Formation in Northern Patagonia, Argentina), shed new light on the function of its spines and the defense behavior in sauropod dinosaurs.

Bajadasaurus was discovered in 2013, by a team of paleontologists from CONICET, Fundación Félix de Azara, Universidad Maimónides, and Museo Paleontológico Ernesto Bachmann. The generic name derived from Bajada (Spanish, in reference to the locality Bajada Colorada) and saurus (Greek, lizard). The specific name derived from pronus (Latin, bent over forward) and spinax (Greek, spine), referring to the anteriorly pointed, curved, neural spines of the cervical vertebrae.

Skeletal elements of Bajadasaurus pronuspinax. From Gallina et al., 2019.

The holotype, MMCh-PV 75, includes a nearly complete skull (left maxilla, left lacrimal, both prefrontals, both frontals, both parietals, both postorbitals, both squamosals, left quadratojugal, both pterygoids, both quadrates, supraoccipital, exoccipital-opisthotic complex, basioccipital, basisphenoid, both prootics, both laterosphenoids, both orbitosphenoids, both dentaries, left surangular, both angulars, both splenials, left prearticular, left articular, isolated upper tooth row), both proatlases, atlantal neurapophyses, axis and the fifth cervical vertebra.

The skull of Bajadasaurus is gracile, with dorsally exposed orbits, dorsoventrally compressed occipital condyle, extremely narrow basipterygoid processes, elongate and slender anterior processes of the squamosals, medially extended post-temporal fenestrae, short lateral temporal fenestrae and a reduced dentition in the maxilla and dentary, that largely differs from other known taxa within Dicraeosauridae. But the most striking feature of Bajadasaurus is the presence of extremely long cervical neural spines that curve anteriorly. Amargasaurus exhibit the same development of cervical neural spine elongation as Bajadasaurus, but the spines of the former point backwards rather than forwards. Dicraeosaurus and Brachytrachelopan show anteriorly inclined neural spines in the cervical vertebrae, but the spines are much shorter than in Bajadasaurus.

A group of Bajadasaurus. Illustration: Jorge A. González.

The discovery of Amargasaurus cazaui in 1991, from the Early Cretaceous beds of La Amarga Formation of Northern Patagonia, renewed the discussion on the peculiar vertebral anatomy of these sauropod dinosaurs, including interpretations as a support structure for a thermoregulatory sail, a padded crest as a display and/or clattering structure, a dorsal hump, or as internal cores of dorsal horn. Those explanation, except the last one, require that these long and extremely gracile bone projections, now recognized in Bajadasaurus as well, can support enough physical stress to avoid fracturing. Bone is stronger and stiffer in passive situations, however, horns and other keratin-based materials are tougher and highly resistant to impact-related fractures. Therefore, the keratinous sheath in Amargasaurus and perhaps Bajadasaurus provides a better mechanical solution against a potential fracture.



Gallina, Pablo A., Apesteguía, Sebastián, Canale, Juan I., Haluza, Alejandro (2019), A new long-spined dinosaur from Patagonia sheds light on sauropod defense system, Scientific Reports volume 9, Article number: 1392 DOI:

Janensch, W. Die Wirbelsäule der Gattung Dicraeosaurus. Palaeontographica Supplement 7, 37–133 (1929).

Salgado, L. & Bonaparte, J. F. Un nuevo saurópodo Dicraeosauridae, Amargasaurus cazaui gen et sp. nov., de la Formación La Amarga, Neocomiano de la provincia del Neuquén, Argentina. Ameghiniana 28, 333–346 (1991).

The hyperthermals of the geological record

During the last 540 million years five mass extinction events shaped the history of the Earth. Those events were related to extreme climatic changes. The geological records show that large and rapid global warming events occurred repeatedly during the course of Earth history.
Our planet’s climate has oscillated between two basic states: the “Icehouse”, and the “Greenhouse”, and superimposed on this icehouse–greenhouse climate cycling, there are a number of geologically abrupt events known as hyperthermals, when atmospheric CO2 concentrations may rise above 16 times (4,800 ppmv). Although each hyperthermal is unique, they are consequence from the release of anomalously large inputs of CO2 into the atmosphere and are relatively short-lived (with the exception of the Permian–Triassic boundary).

A summary of the most significant hyperthermals in the last 300 Myr. From Foster et. al., 2018.

The emplacement of large igneous provinces (LIPs) is commonly associated with hyperthermals, for example, the Siberian Traps at the P–T boundary. The CO2 emissions caused global warming. The SO2 emissions on mixing with water vapour in the atmosphere, caused acid rain, which in turn killed land plants and caused soil erosion. Warmer oceans melted frozen methane located in marine sediments which pushed the global temperatures to higher levels. Additionally, the increased continental weathering induced by acid rain and global warming led to increased marine productivity and eutrophication, and so oceanic anoxia, and marine mass extinctions.

The hyperthermal at the P–T boundary was associated with the most severe terrestrial and oceanic mass extinction of the last 541 Myr, where 96% of species became extinct. It comprises two killing events, one at the end of the Permian (EPME) and a second at the beginning of the Triassic, separated by 60000 years. In terms of carbon isotope excursion, the P–T boundary hyperthermal and the PETM share many similarities, but the warming after the P-T boundary was more extreme and extended for longer than PETM.

Flow chart summarizing proposed cause-and-effect relationships during the end-Permian extinction (From Bond and Wignall, 2014)

The End-Triassic Extinction is probably the least understood of the big five. It has been linked to the eruption of the Central Atlantic Magmatic Province (CAMP), a large igneous province emplaced during the initial rifting of Pangea. Most mammal-like reptiles and large amphibians disappeared, as well as early dinosaur groups. In the oceans, this event eliminated conodonts and nearly annihilated corals, ammonites, brachiopods and bivalves. In the Southern Hemisphere, the vegetation turnover consisted in the replacement to Alisporites (corystosperm)-dominated assemblage to a Classopollis (cheirolepidiacean)-dominated one.

The early Toarcian Oceanic Anoxic Event (T-OAE; ∼183 mya) in the Jurassic Period is considered as one of the most severe of the Mesozoic era. The T-OAE is thought to have been caused by increased atmospheric CO2 triggered by Karoo–Ferrar volcanism. Results from the Paris Bassin indicates that the increasing greenhouse conditions may have caused acidification in the oceans, hampering carbonate bio-mineralisation, and provoking a dramatical loss in the CO2 storage capacity of the oceans.

Tentative changes in mid-latitude vegetation patterns during OAE2. (a) Araucariaceae, (b) other conifers incl. Cheirolepidiaceae, (c) Cupressaceae, (d) angiosperms incl. Normapolles-producing forms, (e) ferns. From Heimhofer et al., 2018.

The early Aptian Oceanic Anoxic Event (OAE1a, 120 Ma) represents a geologically brief time interval characterized by rapid global warming, dramatic changes in ocean circulation including widespread oxygen deficiency, and profound changes in marine biotas. During the event, black shales were deposited in all the main ocean basins. It was also associated with the calcification crisis of the nannoconids, the most ubiquitous planktic calcifiers during the Early Cretaceous. Their near disappearance is one of the most significant events in the nannoplankton fossil record.

The mid-Cretaceous Oceanic Anoxic Event 2 (OAE2, 93 Ma) marks the onset of an extreme phase in ocean temperatures known as the “Cretaceous thermal maximum”. It has been postulated that the OAE2 was triggered by a massive magmatic episode.

Comparison of the effects of anthropogenic emissions (total of 5000 Pg C over 500 years) and PETM carbon release (3000 Pg C over 6 kyr) on the surface ocean saturation state of calcite. From Zeebe, 2013

The Paleocene-Eocene Thermal Maximum (PETM; 55.8 million years ago), was a short-lived (~ 200,000 years) global warming event attributed to a rapid rise in the concentration of greenhouse gases in the atmosphere. It was suggested that this warming was initiated by the melting of methane hydrates on the seafloor and permafrost at high latitudes. During the PETM, around 5 billion tons of CO2 was released into the atmosphere per year, and temperatures increased by 5 – 9°C. This event was accompanied by other large-scale changes in the climate system, for example, the patterns of atmospheric circulation, vapor transport, precipitation, intermediate and deep-sea circulation and a rise in global sea level. But unlike other hyperthermals, the PETM is not associated with significant extinctions.

Anthropogenic climate change and ocean acidification resulting from the emission of vast quantities of CO2 and other greenhouse gases pose a considerable threat to ecosystems and modern society. The combination of global warming and the release of large amounts of carbon to the ocean-atmosphere system during the PETM has encouraged analogies to be drawn with modern anthropogenic climate change. The current rate of the anthropogenic carbon input is probably greater than during the PETM, causing a more severe decline in ocean pH and saturation state. Also the biotic consequences of the PETM were fairly minor, while the current rate of species extinction is already 100–1000 times higher than would be considered natural. This underlines the urgency for immediate action on global carbon emission reductions.


Foster GL, Hull P, Lunt DJ, Zachos JC. (2018) Placing our current‘hyperthermal’ in the context of rapid climate change in our geological past. Phil. Trans. R. Soc. A 376: 20170086

Benton MJ. (2018) Hyperthermal-driven mass extinctions: killing models during the Permian–Triassic mass extinction. Phil. Trans. R. Soc. A 376: 20170076.

Penn, J. L., Deutsch, C., Payne, J. L., & Sperling, E. A. (2018). Temperature-dependent hypoxia explains biogeography and severity of end-Permian marine mass extinction. Science, 362(6419), eaat1327. doi:10.1126/science.aat1327 

Ernst, R. E., & Youbi, N. (2017). How Large Igneous Provinces affect global climate, sometimes cause mass extinctions, and represent natural markers in the geological record. Palaeogeography, Palaeoclimatology, Palaeoecology, 478, 30–52. doi:10.1016/j.palaeo.2017.03.014

Turgeon, S. C., & Creaser, R. A. (2008). Cretaceous oceanic anoxic event 2 triggered by a massive magmatic episode. Nature, 454(7202), 323–326. doi:10.1038/nature07076

Ulrich Heimhofer, Nina Wucherpfennig, Thierry Adatte, Stefan Schouten, Elke Schneebeli-Hermann, Silvia Gardin, Gerta Keller, Sarah Kentsch & Ariane Kujau (2018) Vegetation response to exceptional global warmth during Oceanic Anoxic Event 2, Nature Communications volume 9, Article number: 3832

Zeebe RE and Zachos JC. 2013 Long-term legacy ofmassive carbon input to the Earth system: Anthropocene versus Eocene. Phil Trans R Soc A 371: 20120006.


Top fossils discoveries of 2018.

Ingentia prima outcropping from the soil.

Paraphrasing Dickens, 2018 was the best of years, and it was the worst of years. Marked by extreme weather, earthquakes, and an intense volcanic activity, 2018 is also noted by amazing fossil discoveries. My top list include:

  • The oldest Archaeopteryx

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

The Archaeopteryx story began in  the summer of 1861, two years after the publication of the first edition of Darwin’s Origin of Species, when workers in a limestone quarry in Germany discovered the impression of a single 145-million-year-old feather. Over the years, eleven Archaeopteryx specimens has being recovered. The new specimen from the village of Schamhaupten, east-central Bavaria is the oldest representative of the genus (earliest Tithonian). The shoulder girdles and arms, as well as the skull have been slightly dislocated from their original positions, but the forelimbs remain in articulation. The skull is triangular in lateral outline and has approximately 56 mm long. The orbit is the largest cranial opening (approximately 16 mm long), and the lateral temporal fenestra is collapsed. There are probably four tooth positions in the premaxilla, nine in the maxilla and 13 in the dentary. The postcranial skeleton was affected by breakage and loss of elements prior to or at the time of discovery.

  • Tratayenia rosalesi

Fossilized vertebrae and right hip bone of Tratayenia rosalesi. From Porfiri et al., 2018

Patagonia has yielded the most comprehensive fossil record of Cretaceous theropods from Gondwana, including Megaraptora, a clade of medium-sized and highly pneumatized theropods represented by Fukuiraptor, Aerosteon, Australovenator, Megaraptor, Murusraptor, and Orkoraptor, and characterized by the formidable development of their manual claws on digits I and II and the transversely compressed and ventrally sharp ungual of the first manual digit. Tratayenia rosalesi is the first megaraptoran theropod described from the Santonian Bajo de la Carpa Formation of the Neuquén Group. The genus name is for Tratayén, the locality where the holotype was collected. The specific name honors Diego Rosales, who discovered the specimen in 2006. Tratayenia is also the largest carnivorous taxon known from Bajo de la Carpa Formation, reinforcing the hypothesis that megaraptorids were apex predators in South America from the Turonian through the Santonian or early Campanian, following the extinction of carcharodontosaurids.

  • Lingwulong shenqi

Skeletal reconstruction and exemplar skeletal remains of Lingwulong shenqi. Scale bars = 100 cm for a and 5 cm for b–o. From Xi et al., 2018

Sauropods were the largest terrestrial vertebrates. Their morphology is easy recognizable: a long, slender neck and a tail at the end of a large body supported by four columnar limbs. Sauropods dominated many Jurassic and Cretaceous terrestrial faunas. Although they were globally distributed, the absence of Diplodocoidea from East Asia has been interpreted as a biogeographic pattern caused by the Mesozoic fragmentation of Pangea. Lingwulong shenqi — literally the “amazing dragon from Lingwu” — is the first well-preserved confirmed diplodocoid from East Asia (23 synapomorphies support the placement of Lingwulong within Diplodocoidea with 10 of these being unequivocal). The holotype, (LM) V001a, is a partial skull comprising the braincase, skull roof, and occiput, and an associated set of dentary teeth. The paratype, (LGP) V001b, comprises a semi-articulated partial skeleton including a series of posterior dorsal vertebrae, complete sacrum, the first caudal vertebra, partial pelvis, and incomplete right hind limb. The Lingwulong specimens were found in the Yanan Formation at Ciyaopu, in northwest China. The presence of a conchostracans assemblage (including Palaeoleptoestheria, Triglypta, and Euestheria) is indicative of a Middle Jurassic age. The discovery of Lingwulong undermines the EAIH (East Asian Isolation Hypothesis), forcing a significant revision of hypotheses concerning the origins and early radiation of Neosauropoda.

  • Ingentia prima

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

Ingentia prima — literally the “first giant” in Latin — from the Late Triassic of Argentina shed new lights on the origin of gigantism in this group. The holotype, PVSJ 1086, composed of six articulated posterior cervical vertebrae, glenoid region of right scapula and right forelimb lacking all phalanges, has been recovered from the southern outcrops of the Quebrada del Barro Formation, northwestern Argentina. Discovered in 2015 by Diego Abelín and a team led by Cecilia Apaldetti of CONICET-Universidad Nacional de San Juan, Argentina, this new fossil weighed up to 11 tons and measured up to 32 feet (10 meters) long. Ingentia was unearthed with three new specimens of Lessemsaurus sauropoides. The four dinosaurs belongs to the clade Lessemsauridae, that differs from all other Sauropodomorpha dinosaurs in possessing robust scapulae with dorsal and ventral ends equally expanded; slit-shaped neural canal of posterior dorsal vertebrae; anterior dorsal neural spines transversely expanded towards the dorsal end; a minimum transverse shaft width of the first metacarpal greater than twice the minimum transverse shaft of the second metacarpal; and bone growth characterized by the presence of thick zones of highly vascularized fibrolamellar bone, within a cyclical growth pattern.

  • Caelestiventus hanseni

A 3D printed model of the C. hanseni skull discovered in Utah

Caelestiventus hanseni, from the Upper Triassic of North America, is the oldest pterosaur ever discovered, and it predates all known desert pterosaurs by more than65 million years. The holotype, BYU 20707, includes the left maxilla fused with the jugal, the right maxilla, the right nasal, the fused frontoparietals, the right and left mandibular rami, the right terminal wing phalanx and three fragments of indeterminate bones. The maxilla, jugal, frontoparietal, and mandibular rami of the specimen are pneumatic. The unfused skull and mandibular elements suggest that BYU 20707 was skeletally immature or had indeterminate growth. Based on the relationship between the length of the terminal wing phalanges and wing span in other non-pterodactyloid pterosaurs the new taxon would have a wing span greater than 1.5 m. The significance of C. hanseni lies in its exceptional state of preservation, and its close phylogenetic relationship with Dimorphodon macronyx, indicating that dimorphodontids originated by the Late Triassic and survived the end-Triassic extinction event.

  • Macrocollum itaquii

Skull of Macrocollum itaquii (From Müller et al 2018)

Macrocollum itaquii is the oldest long-necked dinosaur known. Discovered in 2012, from rocks belonging to the upper part of the Candelaria Sequence constrained as about 225 Ma, the three individuals described as M. itaquii are relatively well preserved. The holotype specimen (CAPPA/UFSM 0001a) consists of an almost complete and articulated skeleton. The two paratype specimens (CAPPA/UFSM 0001b and CAPPA/UFSM 0001c) are both articulated skeletons with one missing a skull and its cervical series. The clustered preservation of the three skeletons also represents the oldest evidence of gregarious behaviour in sauropodomorphs, a pattern seen in other Triassic associations, such as the ‘Plateosaurus bonebed’ from Central Europe, and the Mussaurus remains from the Laguna Colorada Formation, Argentina. M. itaquii was only 3.5 meters long and weighed about 101.6 kilograms. In contrast to most Carnian members of the group, the teeth of M. itaquii and other Norian taxa are fully adapted to an omnivore/herbivore diet. The neck elongation may also have provided a competitive advantage for gathering food resources, allowing members of the group to reach higher vegetation. The modifications of the hindlimb of M. itaquii could be related to the progressive loss of cursorial habits.

  • Soft-tissue evidence in a Jurassic ichthyosaur.

Stenopterygius specimen from the Holzmaden quarry. Credit: Johan Lindgren

During the Norian, the evolution of ichthyosaurs took a major turn, with the appearance of the clade Parvipelvia (ichthyosaurs with a small pelvic girdle). They were notably similar in appearance to extant pelagic cruisers such as odontocete whales. An exquisitely fossilized parvipelvian Stenopterygius from the Early Jurassic (Toarcian) of the Holzmaden quarry in southern Germany, indicates that their resemblance with dolphin and whales is more than skin deep. The specimen (MH 432; Urweltmuseum Hauff, Holzmaden, Germany) reveals endogenous cellular, sub-cellular and biomolecular constituents within relict skin and subcutaneous tissue. The external surface of the body is smooth, and was presumably comparable in life to the skin of extant cetaceans. The histological and microscopic examination of the fossil, evinced a multi-layered subsurface architecture. The approximately 100-μm-thick epidermis retains cell-like structures that are likely to represent preserved melanophores. The subcutaneous layer is over 500 μm thick, and comprises a glossy black material superimposed over a fibrous mat. The anatomical localization, chemical composition and fabric of the subcutaneous material is interpreted as fossilized blubber, a hallmark of warm-blooded marine amniotes.

  • Pterosaurs and feathers


Type 3 filaments (arrows) and similar structures (triangles). Scale bars: 10 mm in a, c and d; 1 mm in b. From Yang et al., 2018

Feathers were once considered to be unique avialan structures. Recent studies indicated that non avian dinosaurs, as part of Archosauria, possessed the entirety of the known non keratin protein-coding toolkit for making feathers. Primitive theropods, such as Sinosauropteryx and the tyrannosaurs Dilong and Yutyrannus, and some plant-eating ornithischian dinosaurs, such as Tianyulong and Kulindadromeus, are known from their spectacularly preserved fossils covered in simple, hair-like filaments called ‘protofeathers’. Other integumentary filaments, termed pycnofibres, has been reported in several pterosaur specimens, but there is still a substantial disagreement regarding their interpretation. J. Yang and colleagues described two specimens of short-tailed pterosaurs (NJU–57003 and CAGS–Z070) from the Middle-Late Jurassic Yanliao Biota, in northeast China (around 165-160 million years ago) with preserved structural fibres (actinofibrils) and four different types of pycnofibres. The specimens resemble Jeholopterus and Dendrorhynchoides, but they are relatively small. Pterosaurs were winged cousins of the dinosaurs and lived from around 200 million years ago to 66 million years ago. In the early 1800’s, a fuzzy integument was first reported from the holotype of Scaphognathus crassirostris. A recent study on this specimen shows a subset of pycnofibers and actinofibrils. The discovery of integumentary structures in other pterosaurs, such as Pterorhynchus wellnhoferi(another rhamphorhynchoid pterosaur), and these exquisitely preserved pterosaurs from China, suggest that all Avemetatarsalia (the wide clade that includes dinosaurs, pterosaurs and close relatives) were ancestrally feathered.


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

Porfiri, J.D., Juárez Valieri, Rubé.D., Santos, D.D.D., Lamanna, M.C., A new megaraptoran theropod dinosaur from the Upper Cretaceous Bajo de la Carpa Formation of northwestern Patagonia, Cretaceous Research (2018), doi: 10.1016/j.cretres.2018.03.014.

Xing Xu, Paul Upchurch, Philip D. Mannion, Paul M. Barrett, Omar R. Regalado-Fernandez, Jinyou Mo, Jinfu Ma and Hongan Liu. 2018. A New Middle Jurassic Diplodocoid Suggests An Earlier Dispersal and Diversification of Sauropod Dinosaurs. Nature Communications.9, 2700.  DOI:  10.1038/s41467-018-05128-1 

Cecilia Apaldetti, Ricardo N. Martínez, Ignacio A. Cerda, Diego Pol and Oscar Alcober (2018). An early trend towards gigantism in Triassic sauropodomorph dinosaurs. Nature Ecology & Evolution.

Brooks B. Britt et al. Caelestiventus hanseni gen. et sp. nov. extends the desert-dwelling pterosaur record back 65 million years, Nature Ecology & Evolution (2018). DOI: 10.1038/s41559-018-0627-y

Müller RT, Langer MC, Dias-da-Silva S. 2018, An exceptionally preserved association of complete dinosaur skeletons reveals the oldest long-necked sauropodomorphs. Biol. Lett. 14: 20180633.

Lindgren, J., Sjövall, P., Thiel, V., Zheng, W., Ito, S., Wakamatsu, K., … Schweitzer, M. H. (2018). Soft-tissue evidence for homeothermy and crypsis in a Jurassic ichthyosaur. Nature. doi:10.1038/s41586-018-0775-x

Yang Z. et al., 2018. Pterosaur integumentary structure with complex feather-like branching. Nature Ecology and Evolution

On Pterosaurs and feathers.

Reconstruction of one of the studied anurognathid pterosaurs. Credit: Yuan Zhang/Nature Ecology & Evolution.

Feathers were once considered to be unique avialan structures. Recent studies indicated that non avian dinosaurs, as part of Archosauria, possessed the entirety of the known non keratin protein-coding toolkit for making feathers. Primitive theropods, such as Sinosauropteryx and the tyrannosaurs Dilong and Yutyrannus, and some plant-eating ornithischian dinosaurs, such as Tianyulong and Kulindadromeus, are known from their spectacularly preserved fossils covered in simple, hair-like filaments called ‘protofeathers’.

Other integumentary filaments, termed pycnofibres, has been reported in several pterosaur specimens, but there is still a substantial disagreement regarding their interpretation. J. Yang and colleagues described two specimens of short-tailed pterosaurs (NJU–57003 and CAGS–Z070) from the Middle-Late Jurassic Yanliao Biota, in northeast China (around 165-160 million years ago) with preserved structural fibres (actinofibrils) and four different types of pycnofibres. The specimens resemble Jeholopterus and Dendrorhynchoides, but they are relatively small.


Drawing of of (a) NJU–57003 and (b) CAGS–Z070 with skeletal element identification, outline of
preserved integument, and distribution of the four types of pycnofibres. From Yang et al., 2018.

Types 1 and 4 of pycnofibres occur in both specimens, but types 2 and 3 occur only in CAGS–Z070. This may reflect original biological differences or differences in the taphonomy of the two specimens. The pterosaur type 1 filaments resemble monofilaments in the ornithischian dinosaurs Tianyulong and Psittacosaurus and the coelurosaur Beipiaosaurus. The pterosaur type 2 filaments resemble the brush-like bundles of filaments in the coelurosaurs Epidexipteryx and Yi. Type 3 filaments resemble bristles in modern birds, but surprisingly do not correspond to any reported morphotype in non-avian dinosaurs. The pterosaur type 4 filaments are identical to the radially branched, downy feather-like morphotype found widely in coelurosaurs such as Caudipteryx and Dilong. Functions of these structures could include insulation, tactile sensing, streamlining and colouration (primarily for camouflage and signalling), as for bristles, down feathers and mammalian hairs.

Type 3 filaments (arrows) and similar structures (triangles). Scale bars: 10 mm in a, c and d; 1 mm in b. From Yang et al., 2018

Pterosaurs were winged cousins of the dinosaurs and lived from around 200 million years ago to 66 million years ago. In the early 1800’s, a fuzzy integument was first reported from the holotype of Scaphognathus crassirostris. A recent study on this specimen shows a subset of pycnofibers and actinofibrils. The discovery of integumentary structures in other pterosaurs, such as Pterorhynchus wellnhoferi (another rhamphorhynchoid pterosaur), and these exquisitely preserved pterosaurs from China, suggest that all Avemetatarsalia (the wide clade that includes dinosaurs, pterosaurs and close relatives) were ancestrally feathered.


Yang Z. et al., 2018. Pterosaur integumentary structure with complex feather-like branching. Nature Ecology and Evolution

Barrett PM, Evans DC, Campione NE. 2015 Evolution of dinosaur epidermal structures. Biol. Lett. 11: 20150229.

Kai R.K. Jäger, Helmut Tischlinger, Georg Oleschinski, and P. Martin Sander, Goldfuß was right: Soft part preservation in the Late Jurassic pterosaur Scaphognathus crassirostris revealed by reflectance transformation imaging (RTI) and UV light and the auspicious beginnings of paleo-art,

Craig B. Lowe, Julia A. Clarke, Allan J. Baker, David Haussler and Scott V. Edwards, Feather Development Genes and Associated Regulatory Innovation Predate the Origin of Dinosauria, Mol Biol Evol (2015) 32 (1): 23-28. doi: 10.1093/molbev/msu309