The ‘ghost’ fossils of the future past

Fossil Coccolithophores. Image Credit: S.M. Slater, P. Bown et al / Science journal

Calcareous nannoplankton represent a major component of oceanic phytoplankton, ranging in size from 0.25 to 30 μm. The first records are from the Late Triassic. Their calcareous skeletons can be found in fine-grained pelagic sediments in high concentrations and the biomineralization of coccoliths is a globally significant rock-forming process. This heterogeneous group includes coccoliths, discoasters and nannoconids. They are crucial elements for our understanding of past and present oceans. Their skeletons take up chemical signals from the sea water, in particular isotopes of oxygen and carbon.  In the Jurassic and Cretaceous oceans, the calcareous nannoplankton was the most efficient rock-forming group, for that reason the characterization of calcareous nannofloras in OAE (Oceanic Anoxic Events) intervals are used to improve our understanding of the marine ecosystem and biological processes such as photosynthesis (biological pump) and biomineralisation (carbonate pump) that affect the organic and inorganic carbon cycle, as well as adsorption of atmospheric CO2 in the oceans (Erba, 2013). 

Schematic representation of a generic coccolithophore cell. From Flores & Sierro, 2013.

Coccolithophores are unicellular marine golden-brown algae differing from other Chrysophyta in having two flagella and a third flagella-like appendage called a haptonema. They also posses calcified scales, called coccoliths, at some stage in their life as a protective armour that eventually falls to the ocean floor to build deep-sea ooze and fossil chalks. Declines in the abundance of nannofossils through several past global warming events were linked to biocalcification crises caused by climate change and ocean acidification. Now, a new study presents a global record of ‘ghost’ nannofossils that reveals that  nannoplankton were more resilient to past warming events than traditional fossil evidence would suggest.

Scanning electron microscope images of calcareous nannofossil imprints preserved on the surface of organic matter; Toarcian (including T-OAE interval), Yorkshire, UK. From Slater et al., 2022

The new study focused on the Toarcian Oceanic Anoxic Event (T-OAE ∼183 mya) considered as one of the most severe of the Mesozoic era. This event is associated with a major negative carbon isotope excursion, mass extinction, marine transgression and global warming caused by massive volcanism in the Southern Hemisphere. Previous studies had indicated that during the peak of this event calcareous nannofossils collapsed due to ocean acidification. But the new research have found that these fossils had been overlooked due to their tiny size and their mode of preservation.

Ghost nannofossils  in rocks from the T-OAE, OAE1a and OAE2. Credit: S.M. Slater et al., 2022

After their death, nannofossils were buried in soft sediment at the bottom of the sea, while their imprints were preserved in the surfaces of other organic matter, such as pollen or spores. These imprints—or “ghost”—nannofossils were found in sediments through the TOAE in the UK, Germany, Japan and New Zealand, but also from two similar global warming events in the Cretaceous: the early Aptian Oceanic Anoxic Event (OAE1a, 120 Ma) from Sweden, and the Oceanic Anoxic Event 2 (OAE2, 94 Ma) from Italy. These findings provide new tools to understand how the calcareous nannoplankton respond to warming events.



S. M. Slater, P. Bown, R. J. Twitchett, S. Danise, V. Vajda, Global record of ‘ghost’ nannofossils reveals plankton resilience to high-CO2 and warming, Science (2022).

Elisabetta Erba, Calcareous nannofossils and Mesozoic oceanic anoxic events, Marine Micropaleontology 52 (2004) 85 – 106

Doney, D. S. Busch, S. R. Cooley, K. J. Kroeker, The impacts of ocean acidification on
marine ecosystems and reliant Human communities. Annu. Rev. Environ. Resour. 45, 83
112 (2020).

J.-A. Flores, F.J. Sierro, Flores, PALEOCEANOGRAPHY, BIOLOGICAL PROXIES| Coccolithophores. (2013): 783-794,


Maip macrothorax, the shadow of the death

Maip macrothorax. Image credit: Agustín Ozán

Patagonia has yielded the most comprehensive fossil record of Cretaceous theropods from Gondwana, including Megaraptora, a clade of medium-sized and highly pneumatized theropods characterized by their elongate skulls, and the formidable development of their manual claws on digits I and II. The enigmatic nature of this group has been a matter of discussion since the description of Megaraptor namunhaiquii in 1990s . Other representatives of the clade are Aoniraptor libertatem, Aerosteon riocoloradensis, Australovenator wintonensis, Murusraptor barrosaensis, Tratayenia rosalesi and Orkoraptor burkei. The phylogenetic position of Megaraptora is still controversial. But despite the lack of consensus, megaraptorans themselves remain a well-supported clade. Now, a new megaraptoran theropod dinosaur from the Upper Cretaceous of the Santa Cruz Province, Argentina, sheds light on on these enigmatic predators and their evolutionary radiation.

Maip macrothorax is a large-bodied megaraptorid from lower Maastrichtian Chorrillo Formation in Santa Cruz Province, Argentina. The holotype (MPM 21,545) includes the axis (only lacking both prezygapophyses and its right postzygapophysis), several dorsal and caudal vertebrae, three incomplete cervical ribs, numerous incomplete or fragmentary dorsal ribs, numerous gastral elements, left coracoid, distal end of a second metatarsal, and fragments of the scapula. The generic name, Maip, is derived from an evil entity in Aonikenk mythology that represents “the shadow of the death”. The specific name, macro, derives from the Greek word makrós (meaning long), and the Latin word thorax (meaning chest) in reference to its wide thoracic cavity (which has, approximately, more than 1.20 m width).

Axis of Maip in lateral (A, A´), anterior (B, B´), posterior (C, C´), dorsal (D, D´) and ventral (E, E´). Scale bar: 5 cm. From Aranciaga et al., 2022

The new specimen was discovered in 2019, but due the outbreak of the COVID-19 pandemic in early 2020 the dig was temporarily interrupted. The most striking feature of Maip is its large size. Maip macrothorax was between nine and 10 meters (30-33 feet) and weight about 5 tons. Several vertebrae and ribs of Maip show striations or rugosities interpreted as the attachment sites for the costovertebral and costotransversarium ligaments, a condition not commonly observed in other theropods.

The new study, lead by Aranciaga Rolando, recovered two new clades comprising some derived megaraptorids from South America. The first one, Clade A, comprises Megaraptor, Murusraptor, most of the Cenomanian–Turonian Patagonian forms with 6 or 7 m in length. The second one, Clade B, includes Orkoraptor, Tratayenia, Aerosteon and Maip, most of the Santonian through Maastrichtian megaraptorids from South America, with 8 or 10 m in length. This clade is supported by two synapomorphies: dorsal vertebrae with a bifurcated lamina anterior to the transverse process and forming an accessory fossa, and round and large articular facets of pre- and postzygapophyses of proximal caudal vertebrae. Additionally, the work suggests that after the Turonian, megaraptorids showed an increase in the body size and (with other theropod groups) replaced carcharodontosaurids in the role of apex predators within the Southern continents in the course of the Late Cretaceous.



Aranciaga Rolando, A.M., Motta, M.J., Agnolín, F.L. et al. A large Megaraptoridae (Theropoda: Coelurosauria) from Upper Cretaceous (Maastrichtian) of Patagonia, Argentina. Sci Rep 12, 6318 (2022).

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