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,


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

A mid-Cretaceous enantiornithine frozen in time

Overview of HPG-15-1 in right lateral view. (From Xing et al., 2017)

Overview of HPG-15-1 in right lateral view. (From Xing et al., 2017)

Amber from the Hukawng Valley in northern Myanmar, called Burmese amber, has been commercially exploited for millennia. Of the seven major deposits of amber from the Cretaceous Period, Burmese amber has probably the most diverse paleobiota, including the tail of a non-avian coelurosaurian theropod, and three juvenile enantiornithine birds. The third specimen, HPG-15-1, is the most complete fossil bird discovered in Burmese amber. It comes from the Angbamo site, and measures approximately 86 mm x 30 mm x 57 mm, and weighs 78 g. It  was encapsulated during the earliest stages of its feather production, and  plumage preserves an unusual combination of precocial and altricial features unlike any living hatchling bird.

 Details of the head in HPG-15-1. A, x-ray µCT reconstruction in left lateral view

Details of the head in HPG-15-1. A, x-ray µCT reconstruction in left lateral view (From Xing et al., 2017)

The skull was split when the amber was cut. The rostrum is preserved in one section and the neck and most of the braincase in the other. The skull is mesorostrine. A  single tooth is visible in the left premaxilla. As in Early Cretaceous enantiornithines, the premaxillary corpus is short, forming approximately one-third of the rostrum. The exoccipitals contributed to the dorsal portion of the condyle and were unfused at the time of death. The frontals articulate for most of their length with a small gap between their rostral ends as in Archaeopteryx.  The inner ear and its semicircular canals are preserved. There are at least six articulated cervical vertebrae, including the atlas and axis, preserved in articulation with the skull. The post-axial vertebrae are rectangular with large neural canals, low and caudally displaced neural spines, and a ventral keel as in many enantiornithines. The articulated skull and series of cervical vertebrae bear plumage in dense fields. The individual feathers  are dark brown in color, and appear to consist of tufts of four or more barbs. Skin is preserved as a translucent film in unfeathered regions of both the head and neck.

Microstructure and pigmentation of feathers on wing and body of HPG-15-1. Scale bars equal 1 mm in (A, C); 0.5 mm in (B, D). From Xing et al., 2017

The new specimen also preserves a partial distal wing, the distal right tibiotarsus and complete right foot as well as part of the left pes. Both skeletal material and integumentary structures from the wing’s apex are well-preserved. The plumage consists of fragments of some of the primaries, and alula feathers, some of the secondaries and coverts, and traces of contours from the wing base. The hind limbs preserve feathers and traces of skin. The absence of fusion between the tarsals indicates that the specimen is ontogenetically immature. The proportions of the pedal digits suggest an arboreal lifestyle. Plumage within the femoral and crural tracts consists of neoptile feathers with a short or absent rachis. These feathers are nearly transparent, suggesting that they were pale or white. The skin beneath the crural tract is thin and smooth. The tip of the tail clearly preserves the remains of a single large sheathed rectrix.

The slow post-natal growth results in a protracted period of vulnerability, which is reflected in the Enantiornithes by the large number of juveniles found in the fossil record, whereas young juveniles of other Cretaceous bird lineages are unknown.



Lida Xing, Jingmai K. O’Connor, Ryan C. McKellar, Luis M. Chiappe, Kuowei Tseng, Gang Li, Ming Bai , A mid-Cretaceous enantiornithine (Aves) hatchling preserved in Burmese amber with unusual plumage, (2017), doi: 10.1016/

Murusraptor barrosaensis, a new species in the megaraptorid clade.

Body reconstruction of Murusraptor barrosaensis (From Coria et al., 2016)

Body reconstruction of Murusraptor barrosaensis (From Coria et al., 2016)

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 Megaraptor, Orkoraptor and Aerosteon, 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 (Novas et al, 2013). The enigmatic nature of this group has been a matter of discussion since the description of the first megaraptoran, Megaraptor namunhaiquii. For years, Megaraptor has been alternatively interpreted as belonging to different theropod lineages: as basal coelurosaurians (Novas,1998), basal tetanurans (Calvo et al.,2004; Smith et al., 2008), and allosauroids closely related with carcharodontosaurids (Smith et al., 2007; Benson et al., 2010; Carrano et al., 2012). The main reason for so many different interpretations is the incomplete nature of most available megaraptorid skeletons and the little information about their cranial anatomy.

Murusraptor barrosaensis, from the Upper Cretaceous of Neuquén Province, Argentina, belongs to a Patagonian radiation of megaraptorids together with Aerosteon, Megaraptor and Orkoraptor. Murusraptor, meaning “Wall Raptor”, was discovered in a canyon wall in 2001 during an expedition to Sierra Barrosa in northwestern Patagonia. The holotype specimen includes much of the skull, axial skeleton, pelvis and tibia. The braincase is intact and most of the sutures are still visible, indicating that this was not a fully mature animal.

Different appendicular elements of Murusraptor in their original burial positions (From Coria et al., 2016)

Different appendicular elements of Murusraptor in their original burial positions (From Coria et al., 2016)

Murusraptor barrosaensis is unique in having anterodorsal process of lacrimal longer than height of preorbital process; sacral ribs hollow and tubelike; short ischia distally flattened and slightly expanded dorsoventrally.

Murusraptor shares with all Megaraptoridae two unambiguous synapomorphies: teeth with no enamel wrinkles (interpreted as a reversion to primitive condition in Theropoda); and anterior caudal vertebrae with neural arch bearing prominent centrodiapophysial laminae that define a deep infradiapophysial fossa. Murusraptor also exhibits some characters that are interpreted as convergencies of this taxon with non-tyrannosauroid theropods, including lacrimal with a small pneumatic recess; and a highly pneumatic braincase (Coria et al., 2016)


Rodolfo A. Coria, Philip J. Currie. A New Megaraptoran Dinosaur (Dinosauria, Theropoda, Megaraptoridae) from the Late Cretaceous of Patagonia. PLOS ONE, 2016; 11 (7): e0157973 DOI: 10.1371/journal.pone.0157973

Porfiri, J. D., Novas, F. E., Calvo, J. O., Agnolín, F. L., Ezcurra, M. D. & Cerda, I. A. 2014. Juvenile specimen of Megaraptor (Dinosauria, Theropoda) sheds light about tyrannosauroid radiation. Cretaceous Research 51: 35-55.


Palynological reconstruction of the Antarctic Cretaceous-Paleocene climate.

Artist’s impression of the eastern flank of the Antarctic Peninsula during theMaastrichtian (Artist: James McKay, University of Leeds.)

Artist’s impression of the eastern flank of the Antarctic Peninsula during the Maastrichtian (From Bowman et al, 2014, Artist: James McKay, University of Leeds.)

Past fluctuations in global temperatures are crucial to understand Earth’s climatic evolution. During the Late Cretaceous the global climate change has been associated with episodes of outgassing from major volcanic events, orbital cyclicity and tectonism before ending with the cataclysm caused by a large bolide impact at Chicxulub, on the Yucatán Peninsula, Mexico.
The Antarctic Peninsula is an area of specific interest to modern and past climatic studies, as it seems particularly sensitive to change (Kemp et al., 2014). Most of the studies are focused on Seymour Island which has one of the most expanded Cretaceous–Paleogene successions known. The K-Pg boundary occurs in the uppermost part of the López de Bertodano Formation, where it is marked by a minor iridium anomaly.

The terrestrial palynomorph record at the López de Bertodano Formation was divided into six phases. The first one contains an assemblage dominated by Nothofagidites spp. and Podocarpidites spp., with aquatic fern spores (Azolla spp., Grapnelispora sp.) and rare freshwater algal spores, suggesting a cool and relatively humid period.


Two examples of grains pollen from the Lopez de Bertodano Formation: Podocarpidites sp. (left) and Nothofagidites asperus (right)

Two examples of grains pollen from the Lopez de Bertodano Formation: Podocarpidites sp. (left) and Nothofagidites asperus (right). From Bowman et al, 2014.

In the phase two the increased abundance of Phyllocladidites mawsonii implies a gradual increase in humidity. During phase three, bryophytes began to increase. The phase four is characterised by relatively high abundances of Podocarpidites spp. and relatively low levels of Nothofagidites spp.
The phase 5 is characterised by a rapidly changing sequence of abundance peaks of different taxa, which may indicate a successional turnover in forest composition. The phase six suggests a return to a cool climatic conditions with high abundances of Araucariacites australis and Nothofagidites at the top of the section. It seems that Araucariaceae were capable of surviving long periods of adverse climatic conditions during the Early Pleistocene, but most modern araucarians have subtropical to mesothermal climatic preferences.

The nature of vegetational change in the south polar region suggests that terrestrial ecosystems were already responding to relatively rapid climate change prior to the K–Pg catastrophe. The composition of the terrestrial palynoflora indicates that the Maastrichtian climate fluctuated from cool, humid conditions, through a rapid warming about 2 million years prior to the K–Pg event – which is consistent with the evidence from the marine palynomorph record –  followed by cooling conditions in the earliest Danian.


Two examples of spores from the  Lopez de Bertodano Formation: Grapnelispora sp. (left) and Azolla sp.(right).

Two examples of spores from the Lopez de Bertodano Formation: Grapnelispora sp. (left) and Azolla sp.(right). From Bowman et al, 2014.



Vanessa C. Bowman, Jane E. Francis, Rosemary A. Askinb, James B. Riding, Graeme T. Swindles, Latest Cretaceous–earliest Paleogene vegetation and climate change at the high southern latitudes: palynological evidence fromSeymour Island, Antarctic Peninsula, Palaeogeography, Palaeoclimatology, Palaeoecology, 408. 26-47. 10.1016/j.palaeo.2014.04.018
David B. Kemp, Stuart A. Robinson, J. Alistair Crame, Jane E. Francis, Jon Ineson, Rowan J. Whittle, Vanessa Bowman, and Charlotte O’Brien, A cool temperate climate on the Antarctic Peninsula through the latest Cretaceous to early Paleogene, Geology (2014) doi: 10.1130/G35512.1


The Winter of Our Discontent: short-term cooling following the Chicxulub impact.


The K-T impact by Don Davis.

The Cretaceous–Paleogene extinction that followed the  Chicxulub impact was one of the five great Phanerozoic  mass extinctions. The impact released an estimated energy equivalent of 100 teratonnes of TNT and produced high concentrations of dust, soot, and sulfate aerosols in the atmosphere. Model simulations suggest that the amount of sunlight that reached Earth’s surface was reduced by approximately 20%.This decrease of sunlight caused a drastic short-term global reduction in temperature. This phenomenon is called “impact winter”.

Cold and darkness lasted for a period of months to years.  Photosynthesis stopped and the food chain collapsed. This period of reduced solar radiation may only have lasted several months to decades. Three-quarters of the plant and animal species on Earth disappeared. Marine ecosystems lost about half of their species while freshwater environments shows low extinction rates, about 10% to 22% of genera.

Three factors can be associated with the impact winter in marine and fresh water enviroments. First, starvation caused by the stop of photosynthesis. Second, the loss of dissolved oxygen. Third, the low temperatures. The flux of organic detritus to the sea floor also

A paleogeographic map of the Gulf of Mexico at the end of the Cretaceous (From Vellekoop, 2014)

Three factors can be associated with the impact winter in marine and fresh water environments. First, starvation caused by the stop of photosynthesis. Second, the loss of dissolved oxygen. Third, the low temperatures. Because the late Cretaceous climate was warm, a major challenge for aquatic organisms, especially in inland waters, may have been the persistence of low temperatures. Additionally, the vapour produced by the impact  could have led to global acid rain and a dramatic acidification of marine surface waters.

Fossil evidence for this impact winter was recovered in the Brazos River region of Texas.  The biostratigraphy of the section presents the Ir anomaly, and impact-related tsunami beds. The age of the outcrops was updated using  planktonic foraminifera and  dinocysts.

The “impact winter”  model is supported by a migration of cool, boreal dinoflagellate species into the subtropic Tethyan realm directly across the K–Pg boundary interval and the ingression of boreal benthic foraminifera into the deeper parts of the Tethys Ocean, interpreted to reflect millennial timescale changes in ocean circulation following the impact (Vellekoop, 2014).


Johan Vellekoop, Appy Sluijs, Jan Smit, Stefan Schouten, Johan W. H. Weijers, Jaap S. Sinningh Damsté, and Henk Brinkhuis, Rapid short-term cooling following the Chicxulub impact at the Cretaceous–Paleogene boundary, PNAS (2014) doi: 10.1073/pnas.1319253111

Douglas S. Robertson, William M. Lewis, Peter M. Sheehan and Owen B. Toon, K-Pg extinction patterns in marine and freshwater environments: The impact winter model, Journal of Geophysical Research: Biogeosciences, JUL 2013, DOI: 10.1002/jgrg.20086.

Brief paleontological history of planktonic foraminifera.


Neogloboquadrina dutertrei. (Credit: Dr Kate Darling).

Planktonic foraminifera 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.

planktonic foraminifera evolution

The evolution of early planktonic foraminifera (From Boudagher-Fadel, 2013)

All species of Late Triassic and Jurassic planktonic foraminifera are members of the superfamily Favuselloidea. They present a test composed by aragonite, with microperforations, and sub-globular adult chambers. After the major End Triassic event, the Jurassic period saw warm tropical greenhouse conditions worldwide. The surviving planktonic foraminifera were usually dominated by small globular forms.

It was suggested  that a second transition from a benthic to a planktonic mode of life took place at the Jurassic, which occurred under conditions similar to those that triggered planktonic speciation in the Late Triassic (hot and dry global climate, and low sea levels).

During the Cretaceous,  the favusellids must have made the transition from being aragonitic to calcitic.  Also, in the Late Aptian there was a significant number of planktonic foraminiferal extinctions, but these were compensated by the establishment of a large number of new genera at the Aptian–Albian boundary.

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

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

The Paleogene assemblage of planktonic foraminifera was derived from the few species that survive the mass extinction event at the end of the Cretaceous.

In the Early Miocene, the planktonic foraminifera were most abundant and diverse in the tropics and subtropics, and after the Mid-Miocene Climatic Optimum, many species were adapted to populate temperate and sub-polar oceans.

During the Middle and Late Pliocene, the final closure of the Central American seaway, changed oceanic circulation and drove a significant number of species extinctions. Most modern, living species originated in the Pliocene and Pleistocene.


Armstrong, H. A., Brasier, M. D., 2005. Microfossils (2nd Ed). Blackwell, Oxford.

Boudagher-Fadel, MK; (2013) Biostratigraphic and Geological Significance of Planktonic Foraminifera. (2nd ed.)