Origin of animals at a time of chaotic oxygen levels

Every organism that you can easily see is a eukaryote, the vast majority of which depend on the availability of oxygen molecules. The range of genetic variation in a wide variety of eukaryotes suggests, using a molecular ‘clock’, that the first of them arose between 2000 to 1000 Ma ago. It possibly originated as a symbiotic assemblage of earlier prokaryote cells ‘bagged-up’ within a single cell wall: Lynn Margulis’s hypothesis of endosymbiosis. It had to have happened after the Great Oxygenation Event (GOE 2.4 to 2.2 Ga), before which free oxygen was present in the seas and atmosphere only at vanishingly small concentrations. Various single-celled fossil possibilities have been suggested to be the oldest members of the Eukarya but are not especially prepossessing, except for one bizarre assemblage in Gabon. The first inescapable sign that eukaryotes were around is the appearance of distinctive organic biomarkers in sediments about 720 Ma old. The Neoproterozoic is famous for its Snowball Earth episodes and the associated multiplicity of large though primitive animals during the Ediacaran Period (see: The rise of the eukaryotes; December 2017).

The records of carbon- and sulfur isotopes in Neo- and Mesoproterozoic sedimentary rocks are more or less flat lines after a mighty hiccup in the carbon and sulfur cycles that followed the GOE and the earliest recorded major glaciation of the Earth. The time between 2.0 and 1.0 Ga has been dubbed ‘the Boring Billion’. At about 900 Ma, both records run riot. Sulfur isotopes in sediments reveal the variations of sulfides and sulfates on the seafloor, which signify reducing and oxidising conditions respectively.  The δ13C record charts the burial of organic carbon and its release from marine sediments related to reducing and oxidising conditions in deep water. There were four major ‘excursions’ of δ13C during the Neoproterozoic, which became increasingly extreme. From constant anoxic, reducing conditions throughout the Boring Billion the Late Neoproterozoic ocean-floor experienced repeated cycles of low and high oxygenation reflected in sulfide and sulfate precipitation and by fluctuations in trace elements whose precipitation depends on redox conditions. By the end of the Cambrian, when marine animals were burgeoning, deep-water oxic-anoxic cycles had been smoothed out, though throughout the Phanerozoic eon anoxic events crop up from time to time.

Atmospheric levels of free oxygen relative to that today (scale is logarithmic) computed using combined carbon- and sulfur isotope records from marine sediments since 1500 Ma ago. The black line is the mean of 5,000 model runs, the grey area represents ±1 standard deviations. The pale blue area represents previous ‘guesstimates’. Vertical yellow bars are the three Snowball Earth events of the Late Neoproterozoic (Sturtian, Marinoan and Gaskiers). (Credit: Krause et al., Fig 1a)

The Late Neoproterozoic redox cycles suggest that oxygen levels in the oceans may have fluctuated too. But there are few reliable proxies for free oxygen. Until recently, individual proxies could only suggest broad, stepwise changes in the availability of oxygen: around 0.1% of modern abundance after the GOE until about 800 Ma; a steady rise to about 10% during the Late Neoproterozoic; a sharp rise to an average of roughly 80% at during the Silurian attributed to increased photosynthesis by land plants. But over the last few decades geochemists have devised a new approach based on variations on carbon and sulfur isotope data from which powerful software modelling can make plausible inferences about varying oxygen levels. Results from the latest version have just been published (Krause, A.J. et al. 2022. Extreme variability in atmospheric oxygen levels in the late Precambrian. Science Advances, v. 8, article 8191; DOI: 10.1126/sciadv.abm8191).

Alexander Krause of Leeds University, UK, and colleagues from University College London, the University of Exeter, UK and the Univerisité Claude Bernard, Lyon, France show that atmospheric oxygen oscillated between ~1 and 50 % of modern levels during the critical 740 to 540 Ma period for the origin and initial diversification of animals. Each major glaciation was associated with a rapid decline, whereas oxygen levels rebounded during the largely ice-free episodes. By the end of the Cambrian Period (485 Ma), by which time the majority of animal phyla had emerged, there appear to have been six such extreme cycles.

Entirely dependent on oxygen for their metabolism, the early animals faced periodic life-threatening stresses. In terms of oxygen availability the fluctuations are almost two orders of magnitude greater than those that animal life faced through most of the Phanerozoic. Able to thrive and diversify during the peaks, most animals of those times faced annihilation as O2 levels plummeted. These would have been periods when natural selection was at its most ruthless in the history of metazoan life on Earth. Its survival repeatedly faced termination, later mass extinctions being only partial threats. Each of those Phanerozoic events was followed by massive diversification and re-occupation of abandoned and new ecological niches. So too those Neoproterozoic organism that survived each massive environmental threat may have undergone adaptive radiation involving extreme changes in their form and function. The Ediacaran fauna was one that teemed on the sea floor, but with oxygen able to seep into the subsurface other faunas may have been evolving there exploiting dead organic matter. The only signs of that wholly new ecosystem are the burrows that first appear in the earliest Cambrian rocks. Evolution there would have ben rife but only expressed by those phyla that left it during the Cambrian Explosion.

There is a clear, empirical link between redox shifts and very large-scale glacial and deglaciation events. Seeking a cause for the dramatic cycles of climate, oxygen and life is not easy. The main drivers of the greenhouse effect COand methane had to have been involved, i.e. the global carbon cycle. But what triggered the instability after the ‘Boring Billion’? The modelled oxygen record first shows a sudden rise to above 10% of modern levels at about 900 Ma, with a short-lived tenfold decline at 800 Ma. Could the onset have had something to do with a hidden major development in the biosphere: extinction of prokaryote methane generators; explosion of reef-building and oxygen-generating stromatolites? How about a tectonic driver, such as the break-up of the Rodinia supercontinent? Then there are large extraterrestrial events … Maybe the details provided by Krause et al. will spur others to imaginative solutions. See also: How fluctuating oxygen levels may have accelerated animal evolution. Science Daily, 14 October 2022

Signs of massive hydrocarbon burning at the end of the Triassic

One of the ‘Big Five’ mass extinctions occurred at the end of the Triassic Period (~201 Ma), whose magnitude matches that of the more famous end-Cretaceous (K-Pg) event. It roughly coincided with the beginning of break-up of the Pangaea supercontinent that was accompanied by a major episode of volcanism preserved in the Central Atlantic Magmatic Province (CAMP). Eastern North America, West Africa and northern South America reveal scattered patches of CAMP flood basalts, swarms of dykes and large intrusive sills. Like all mass extinctions, that at the Triassic-Jurassic boundary left a huge selection of vacant or depleted ecological niches ready for evolution to fill by later adaptive radiation of surviving organisms. Because it coincided with continental break-up and drift, unlike other such events, evolution proceeded in different ways on the various wandering land masses and in newly formed seas (see  an excellent animation of the formation and break-up of Pangaea – move the slider to 3 minutes for the start of break-up). The Jurassic was a period of explosive evolution among all groups of organisms. The most notable changes were among marine cephalopods, to give rise to a bewildering variety of ammonite species, and on land with the appearance and subsequent diversification of dinosaurs.

Pangaea at the end of the Triassic (top) and in Middle Cretaceous times (Credit: screen shots from animation by Christopher Scotese)

Many scientists have ascribed the origin of these events to the CAMP magmatic activity and the release of huge amounts of methane to trigger rapid global warming. In October 2021 one group focused on a special role for the high percentages of magma that never reached the surface and formed huge intrusions that spread laterally in thick sedimentary sequences to ‘crack’ hydrocarbons to their simplest form, CH4 or methane. A sedimentary origin of the methane, rather than its escape from the mantle, is indicated by the carbon-isotope ‘signature’ of sediments deposited shortly after the Tr-J event. The lighter isotope 12C rose significantly relative to 13C, suggesting an organic source – photosynthesis selectively takes up the lighter isotope.

By examining the element mercury (Hg) in deep ocean sediments from a Tr-J sedimentary section now exposed in Japan, scientists from China, the US and Norway have added detail to the methane-release hypothesis (Shen, J et al. 2022. Mercury evidence for combustion of organic-rich sediments during the end-Triassic crisis. Nature Communications, v. 13, article 1307; DOI:10.1038/s41467-022-28891-8). The relative proportions of Hg isotopes strongly suggest that the mercury had been released, as was the methane, from organic-rich sediments rather than from the CAMP magmas (i.e. ultimately from the mantle) through gasification and then burning at the surface.

The hypothesis is enlivened by a separate study (Fox C.P. et al. 2022. Flame out! End-Triassic mass extinction polycyclic aromatic hydrocarbons reflect more than just fire. Earth and Planetary Science Letters, v. 584, article 117418; DOI: 10.1016/j.epsl.2022.117418) that sees magmatic heating as being not so important. Calum Fox and colleagues at Curtin University, Western Australia analysed sediments from a Triassic-Jurassic sedimentary sequence near the Severn Bridge in SW England, focusing on polycyclic hydrocarbons in them. Their results show little sign of the kinds of organic chemical remnants of modern wildfires. Instead they suggest a greater contribution from soil erosion by acid rain that increased input of plant debris to a late Triassic marine basin

See also: How a major volcanic eruption paved the way for the rise of the dinosaurs Eureka Alert 23 March 2022;  Soil erosion and wildfire: another nail in coffin for Triassic era. Science Daily, 21 March 2022