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 Milankovich Effect during Snowball Earth episodes

The idea that the Earth was like a giant snowball during the Neoproterozoic Era arose from the discovery of rocks of that age that could only have formed as a result of glaciation. However, unlike the Pleistocene ice ages, evidence for these much older glacial conditions occurs on all continents. In some locations remanent magnetism in sedimentary rocks of that age is almost horizontal; i.e. they had been deposited at low magnetic latitudes, equivalent to the tropics of the present day. Frigid as it then was, the Earth still received solar heating and magmatic activity would have been slowly adding CO2 to the atmosphere so that less heat escaped – a greenhouse effect must have been functioning. Moreover, an iced-over world would not have been supporting much photosynthetic life to draw down the greenhouse gas into solid carbohydrates and carbonates to be buried on the ocean floor. As far as we know the Solar System’s geometry during the Neoproterozoic was much as it is today. So changes in the gravitational fields induced by the orbiting Giant Planets would have been affecting the shape (eccentricity) of Earth’s orbit, the tilt (obliquity) of its rotational axis and the precession (wobble) of its rotation as they do at present through the Milankovich effect. These astronomical forcings vary the amount of solar energy reaching the Earth’s surface. It has been suggested that a Snowball Earth’s climate system would have been just as sensitive to astronomical forcing as it has been during the last 2 million years or more. Proof of that hypothesis has recently been achieved, at least for one of the Snowball events (Mitchell, R.N. and 8 others 2021. Orbital forcing of ice sheets during snowball Earth. Nature Communications, v. 12, article 4187; DOI: 10.1038/s41467-021-24439-4).

Another of the enigmas of the Neoproterozoic is that after and absence of more than a billion years banded iron formations (see: Banded iron formations (BIFs) reviewed, December 2017) began to form again. BIFs are part of the suite of sedimentary rocks that characterise Snowball Earth events, often alternating with glaciogenic sediments. Throughout each cold cycle – the Sturtian (717 to 663 Ma) and Marinoan (650 to 632 Ma) glacial periods – conditions of sediment deposition varied a great deal from place to place and over time. Some sort of cyclicity is hinted at, but the pace of alternations has proved impossible to check, partly because the dominant rocks (glacial conglomerates or diamictites) show little stratification and others that are bedded (various non-glacial sandstones) vary from place to place and give no sign of rates of deposition, having been deposited under high-energy conditions. BIFs, on the other hand are made up of enormous numbers of parallel layers on scales from millimetres to centimetres. Bundles of bands can be traced over large areas, and they may represent repeated cycles of deposition.

Typical banded iron formation

How BIFs formed is crucial. They were precipitated from water rich in dissolved iron in its reduced Fe2+ (ferrous) form, which originated from sea-floor hydrothermal vents. Precipitation occurred when the amount of oxygen in the water increased the chance of electrons being removed from iron ions to transform them from ferrous to ferric (Fe3+). Their combination with oxygen yields insoluble iron oxides. Cyclical changes in the availability of oxygen and the balance between reducing and oxidising conditions result in the banding. In fact several rhythms of alternation are witnessed by repeated packages at deci-, centi- and millimetre scales within each BIF deposit. Overall the packages suggest a constant rate of deposition: a ‘must-have’ for precise time-analysis of the cycles. BIFs contain both weakly magnetic hematite (Fe2O3) and strongly magnetic magnetite (Fe3O4), their ratio depending on varying geochemical conditions during deposition. Ross Mitchell of Curtin University, Western Australia and his Chinese, Australian and Dutch colleagues measured magnetic susceptibility at closely spaced intervals (1 and 0.25 m) in two section of BIFs from the Sturtian glaciation in the Flinders Ranges of South Australia. Visually both sections show clear signs of two high-frequency and three lower frequency kinds of cycles, expressed in thickness.

The tricky step was converting the magneto-stratigraphic data to a time series. High-precision zircon U-Pb dating of volcanic rocks in the sequence suggested a mean BIF deposition rate of 3.7 to 4.4 cm per thousand years. This allowed the thickness of individual bands and packages to be expressed in years, the prerequisite for time-series analysis of the BIF magneto-stratigraphic sequence. This involves a mathematical process known as the Fast-Fourier Transform, which expresses the actual data as a spectral curve. Peaks in the curve represent specific frequencies expressed as cycles per metre. The rate of deposition of the BIF allows each peak to be assigned a frequency in years, which can then be compared with the hypothetical spectrum associated with the Milankovich effect. One of the BIF sequences yielded peaks corresponding to 23, 97 and 106 ka. These match the effects of variation in precession (23 ka) and ‘short’ orbital eccentricity (97 and 106 ka) found in Cenozoic sea-floor sediments and ice cores. The other showed peaks at 405, 754 and 1.2 Ma corresponding to ‘long’ orbital eccentricity and long-term features of both obliquity and precession. Quite a result! But how does this bear on Snowball Earth events? Cyclical changes in solar heating would have affected the extent of ice sheets and sea ice at all latitudes, forcing episodes of expansion and contraction and thus changes in sediment supply to the sea floor. That helps explain the many observed variations in sedimentation other than that of BIFs. Rather than supporting a ‘hard’ Snowball model of total marine ice cover for millions of years, it suggests that such an extreme was relieved by period of extensive open water, much as affected the modern Arctic Ocean for the last 2 million years or so. There could have been global equivalents of ice ages and interglacials during the Sturtian and Marinoan. ‘Hard’ conditions would have shut down much of the oceans’ biological productivity, periodically to have been reprieved by more open conditions: a mechanism that would have promoted both extinctions and evolutionary radiations. Snowball events may have driven the takeover of prokaryote (bacteria) dominance by that of the multicelled eukaryotes that is signalled by the Ediacaran faunas that swiftly followed glacial epochs and the explosion of multicelled life during the Cambrian. As eukaryotes, we may well owe our existence to Snowball.

The East African Orogen: Neoproterozoic tectonics on display

Over a period of about 300 Ma the fragmentation of a supercontinent, Rodinia, drove a round of sea-floor spreading and continental drift that culminated in reassembly of the older continental pieces and entirely new crust in a new supercontinent, Gondwana. The largest source of evidence for this remarkable tectonic turnaround is a belt stretching N-S for over 3000 km from southern Israel through East Africa to Mozambique. At its widest the belt exposes Neoproterozoic  rocks and structures for some 1700 km E-W from west of the Nile in northern Sudan almost to Riyadh in Saudi Arabia.  This Arabian-Nubian Shield tapers southwards to thin out completely in northern Tanzania between far older cratons and in a state of high-grade metamorphism.

This East African Orogen has long been considered the best exposed bowels of former mountain building that there are: results of continent-continent collision and the bulldozing together of many oceanic arcs and remnants of oceanic lithosphere that once separated the cratons. This was much more complex than a case of head-on tectonics, the northward-swelling Arabian-Nubian Shield showing all the signs of being like a gigantic ‘pip’ squeezed out northwards from two cratonic jaws during the last stages of what is often called the Pan African Orogeny. Interestingly, the line of the orogen is roughly followed by East Africa’s other giant feature, the Rift Valley; actually two of them following Pan African terranes. A continental scale anisotropy has been reactivated and subject to extensional tectonics, and maybe in future a new round of sea-floor spreading as has begun in the Red Sea, some half a billion years after it formed.

Simplified geological map of the East African Orogen courtesy of the authors of Fritz et al 2013
Simplified geological map of the East African Orogen courtesy of the authors of Fritz et al 2013

Now there is an opportunity for anyone to download and read a digest of East African orogenic processes compiled by researchers from several countries along the belt and their colleagues from North America, Europe and Australia who have been privileged to work in this vast area (Fritz, H and 13 others 2013. Orogen styles in the East African orogen: A review of the Neoproterozoic to Cambrian tectonic evolution. Journal of African Earth Sciences, v. 86, p. 65-106 Click on the link, scroll to the Open Access article to download the 12 Mb PDF version). The authors present superb simplified geological maps of each major part of the orogen, a vast array of references and well-written accounts of its sector-by-sector tectonic and metamorphic evolution, variations in style and broad tectonic setting.

Early animals and Snowball Earth

"SNOWBALL EARTH" - 640 million years ago
The Earth 640 million years ago during the Marinoan ‘Snowball’ event (credit: Cornell University via Flickr)

Palaeobiologists generally believe that without a significant boost to oxygen levels in the oceans macroscopic eukaryotes, animals in particular, could not have evolved. Although the first signs of a rise in atmospheric oxygen enter the stratigraphic record some 2.4 billion years ago and eukaryote microfossils appeared at around 2 Ga, traces of bulky creatures suddenly show up much later at ~610 Ma with possible fossil bilaterian embryos preserved in 630 Ma old sediments. An intriguing feature of this Ediacaran fauna is that it appeared shortly after one of the Neoproterozoic global glaciations, the Marinoan ‘Snowball’ event: a coincidence or was there some connection? It has looked very like happenstance because few if any signs of a tangible post-Marinoan rise in environmental oxygen have been detected. Perhaps the sluggish two billion-year accumulation of free oxygen simply passed the threshold needed for metazoan metabolism. But there are other, proxy means of assessing the oxidation-reduction balance, one of which depends on trace metals whose chemistry hinges on their variable valency. The balance between soluble iron-2 and iron-3 that readily forms insoluble compounds is a model, although iron itself is so common in sediments that its concentration is not much of a guide. Molybdenum, vanadium and uranium, being quite rare, are more likely to chart subtle changes in the redox conditions under which marine sediments were deposited.

English: Cropped and digitally remastered vers...
Dickinsonia; a typical Ediacaran animal. Scale in cm (credit: Wikipedia)

Swapan Sahoo of the University of Nevada and colleagues from the USA, China and Canada detected a marked increase in the variability of Mo, V and U content of the basal black shales of the Doushantuo Formation of southern China, which contain the possible eukaryote embryos (Sahoo, S.K and 8 others 2012. Ocean oxygenation in the wake of the Marinoan glaciation. Nature, v. 489, p. 546-549). These rocks occur just above the last member of the Marinoan glacial to post-glacial sedimentary package and are around 632 Ma old. Since the black shales accumulated at depths well below those affected by surface waves that might have permitted local changes in the oxygen content of sea water the geochemistry of their formative environment ought not to have changed if global chemical conditions had been stable: the observed fluctuations may represent secular changes in global redox conditions. The earlier variability settles down to low levels towards the top of the analysed sequence, suggesting stabilised global chemistry.

What this might indicate is quite simple to work out. When the overall chemistry of the oceans is reducing Mo, V and U are more likely to enter sulfides in sediments, thereby forcing down their dissolved concentration in sea water. With a steady supply of those elements, probably by solution from basalt lavas at ocean ridges, sedimentary concentrations should stabilise at high levels in balance with low concentrations in solution. If seawater becomes more oxidising it holds more Mo, V and U in solution and sediment levels decline. So the high concentrations in sediments mark periods of global reducing conditions, whereas low values signal a more oxidising marine environment. Sahoo et al.’s observations suggest that marine geochemistry became unstable immediately after the Marinoan glaciation but settled to a fundamentally more oxidising state than it had been in earlier times, perhaps by tenfold increase in atmospheric oxygen content. So what might have caused this and the attendant potential for animals to get larger in the aftermath of the Snowball Earth event? One possibility is that the long period of glaciers’ grinding down continental crust added nutrients to the oceans. Once warmed and lit by the sun they hosted huge blooms of single-celled phytoplankton whose photosynthesis became an oxygen factory and whose burial in pervasive reducing conditions on the sea bed formed a permanent repository of organic carbon. The outcome an at-first hesitant oxygenation of the planet and then a permanent fixture opening a window of opportunity for the Ediacarans and ultimately life as we know it.

Pan African Review

A terrane boundary close to the Nile in the Sudan, detected by radar from the Space Shuttle: the Keraf Suture. From NASA

Undoubtedly the best exposed and one of the biggest examples, the accretionary orogen of the Arabian-Nubian Shield (ANS) is a witness to the creation of a supercontinent from the remnants of an earlier one. At about 1 Ga, most of the Earth’s continental material was clumped together in the Rodinia supercontinent that existed for a quarter of a billion years. At a time of massive mantle upheaval that left most crust of that age affected by basaltic magmatism, in the form of lava flows and dyke swarms, Rodinia began to break up at 800 Ma to scatter continental fragments. Subduction zone accommodated this continental drift to form many ocean and continental-margin volcanic arcs. The ANS is a repository for many of these arcs which episodically accreted between earlier cratons to the west in Africa and those comprising Somalia and the present Indian subcontinent. Primarily the terranes are oceanic in origin and formed in the aftermath of the dismemberment of Rodinia, although a few slivers of older, reworked crust occur in Saudi Arabia and Yemen. Among the various components are ophiolites marking sutures and other major tectonic features of the orogen. The shape of the Shield is unlike that of any other major orogen of later times, for it shrinks from a width estimated at ~2000 km in Arabia to the north to vanish just south of the Equator in southern Kenya. This ‘pinched’ structure has suggested to some that the bulk of the new crust was forced laterally northwards when the African and Indian cratons collided, in the manner of toothpaste from a trodden-on tube.

Today the ANS is a harsh place, some off-limits to geologists either for political reasons or the sheer hostility and remoteness of the environment. Yet a picture has emerged, bit by bit, over the last 30 years. So a detailed review of the most extensive and varied part from 7° to 32°N and 26° to 50°E – in Egypt, Saudia Arabia, eastern Sudan, Eritrea, Yemen and northern Ethiopia is especially welcome (Johnson, P.R. et al. 2011. Late Cryogenian–Ediacaran history of the Arabian–Nubian Shield: A review of depositional, plutonic, structural, and tectonic events in the closing stages of the northern East African Orogen. Journal of African Earth Sciences, v. 61, p. 167-232). Peter Johnson himself compiled a vast amount of information during his career with the US Geological Survey Mission in Saudi Arabia and has blended the inevitably diverse ideas of his 7 co-workers – but by no means all the ideas that are in the literature. The result is a readable and well illustrated account of how the ANS assembled tectonically during times when a near-global glaciation took place, and the first macroscopic animals appear in the fossil record. Tillites and other glaciogenic rocks from the Marinoan ‘SnowBall’ occur from place to place in the ANS, as do banded iron formations that made a surprise return after a billion-year or longer absence in the Cryogenian Period . Coincidentally, glacial conditions returned to the region twice in Ordovician and Carboniferous to Permian times, forming distinctive, tectonically undisturbed sediments in the Phanerozoic cover that unconformably overlies the Neoproterozoic orogen.

Except in a few areas only recently explored, geologists have assiduously dated events in the ANS, showing nicely that all the basement rock formed after 800 Ma, and that orogenic events culminated before the start of the Cambrian period, although one or two unusual granites intruded as late as the Ordovician. The deformation is immense in places, with huge nappes, often strike-slip shear zones and exposure ranging from the lowest metamorphic grade to that in which water and granitic magma was driven from the lower Pan African crust. The range of exposed crustal levels stems partly from the tectonics, but owes a lot to the 2-3 km of modern topographic relief, unique to NE Africa and Arabia. Yet it is not uncommon to come upon delicate features such as pillowed lavas, conglomerates and finely laminated volcanoclastic tuffs. Following tectonic welding, more brittle deformation opened subsiding basins that contain exclusively sedimentary rocks derived from the newly uplifted crust, both marine and terrestrial in formation (basins of this type, in Eritrea and Ethiopia, unfortunately do not figure in the regional maps). Much of the ANS is currently the object of a gold rush, encouraged by a rising world price for the ‘inflation-proof’ comfort blanket provided by the yellow metal. Consequently, newcomers to the stampede will be well advised to mug-up on the regional picture of occurrences and gold-favourable geology provided in the review, and may be interested by other exploration possibilities for rare-earth metals and other rising stars on the London Metal Exchange, such as tin, which are often hosted in evolved granites, that stud the whole region.

Snowball Earth melting hypothesis weakened

"SNOWBALL EARTH" - 640 million years ago
Artist's impression of the Neoproterozoic Earth during a Snowball episode. Image by guano via Flickr

The combination of glaciogenic sediments with palaeomagnetic evidence for their formation at low-latitudes, together with dates that show glacial events were coeval in just two or three Neoproterozoic episodes are the linchpins for the Snowball Earth hypothesis.  There is little doubt that the latest Precambrian Era did witness such extraordinary climatic events. Evidence is also accumulating that, in some way, they were instrumental in that stage of biological evolution from which metazoan eukaryotes emerged: the spectacular Ediacaran fossil assemblages follow on the heels of the last such event (see Bigging-up the Ediacaran in Earth Pages for March 2011). One of the difficulties with the ‘hard’ Snowball Earth hypothesis is how the middle-aged planet was able to emerge from a condition of pole-to-pole ice cover; hugely increased reflectivity of that surface should have driven mean global temperature down and down. Clearly the Earth did warm up on each occasion, and the leading model for how that was possible is massive release of greenhouse gases from sea-floor sediments or deep-ocean waters to increase the heat-retaining powers of the atmosphere; sufficiently voluminous release from volcanic action seems less likely as there is little evidence of upsurges in magmatism coinciding with the events. Almost all glaciogenic units from the Neoproterozoic have an overlying cap of carbonate rocks, indicating that hydrogen carbonate (formerly bicarbonate) ions together with those of calcium and magnesium suddenly exceeded their solubilities in the oceans.

White flocculent mats in and around the extrem...
Modern sea-floor hydrothermal vent. Image via Wikipedia

To seek out a possible source for sufficient carbon release in gaseous form geochemists have turned to C-isotopes in the cap carbonates. Early studies revealed large deficits in the heavier stable isotope of carbon (13C) that seemed to suggest that the releases were from large reservoirs of carbon formed by burial of dead organisms: photosynthesis and other kinds of autotrophy at the base of the trophic pyramid selectively take up lighter 12C in forming organic tissues compared with inorganic chemical processes). As in the case of the sharp warming event at the Palaeocene-Eocene boundary around 55.8 Ma ago (See The gas-hydrate ‘gun’ in June 2003 Earth Pages), these negative d13C spikes have been interpreted as due to destabilisation of gas hydrates in sea-floor sediments to release organically formed methane gas. This powerful greenhouse gas would have quickly oxidised to CO2 thus acidifying the oceans by jacking up hydrogen carbonate ion concentrations.  Detailed carbon-, oxygen- and strontium-isotope work in conjunction with petrographic textures in a Chinese cap carbonate (Bristow, T.F. et al. 2011. A hydrothermal origin for isotopically anomalous cap dolostone cements from south China. Nature, v. 274, p. 68-71) suggests an alternative mechanism to produce the isotopically light carbon signature at the end of Snowball events. The greatest 13C depletion occurs in carbonate veins that cut through the cap rock and formed at temperatures up to 378°C and even the early-formed fine grained carbonate sediment records anomalously high temperatures. So, it seems as if the cap-rock was thoroughly permeated by hydrothermal fluids, more than 1.6 Ma after it formed on the sea floor. This triggered oxidation of methane within the sediments themselves, with little if any need for an atmospheric origin through massive methane release from destabilised gas hydrates elsewhere.

Eukaryote conquest of the continents

NW end of a classic example of a mesa form of ...
Suilven, a spectacular outlier of Torridonian terrestrial sandstones resting on a buried landscape of Archaean gneisses near Lochinver, Sutherland. Image via Wikipedia

Geologists often assume that the continents were first colonised by plants, insects then vertebrates beginning in the Ordovician Period with preservation of spores very like those of the liverworts, which incidentally can only be removed from gravel driveways by the use of acetic acid, glyphosate, pycloram and flamethrowers having no lasting effect. The most intractable of all organisms found on the land surface today are prokaryotic (nucleus-free cells) cyanobacteria whose biofilms cement desert varnish (see Desert varnish, May 2008 in Subjects: GIS and Remote Sensing). Cyanobacteria have long been suspected to have been the first life forms to adopt a terrestrial habit, and their cells have been discovered in the now-famous Neoproterozoic lagerstätten in the Doushantuo Formation of China (see The earliest lichens, May 2005 in Subjects: Geobiology, palaeontology, and evolution) The oldest un-metamorphosed sediments in Britain, the Torridonian redbeds that form the magnificent scenery of north-western Scotland, now push back the date of the earliest eukaryotic (cells with nuclei) terrestrial life, of which we are one form, half a billion years before the Doushanto cyanobacteria (Strother, P.K. et al. 2011. Earth’s earliest non-marine eukaryotes. Nature, v. 473, p. 505-509). The Torridonian is one of the thickest (~12 km) terrestrial sequences on the planet, and spans a time range of around 200 Ma (1.2 to 1 Ga). It is a repository of almost the entire range of humid continental sedimentary environments: colluvial fan; bajada; alluvial; deltaic and lacustrine build-ups. Grey lake-bed mudstones and phosphate nodules in the Torridonian yield small organic fossils lumped in the sack-term acritarchs. Similar bodies, whose affinities are diverse and generally obscure, have been reported from marine sediments as old as 3.2 Ga. The fascination of those from the Torridonian, other than their terrestrial association, is that some include aggregates of spherical cells with tantalising suggestions of central nuclei and, as a whole assemblage, exhibit a range of morphologies far beyond that of nucleus-free prokaryotes and the signature of cytoskeletal filaments that form a ‘scaffold’ for eukaryote cells. Worth noting is that one of the authors is Martin Brasier of Oxford University, whose meticulous bio-morphological skills in microscopy has made him one of the foremost critics of speculation on Precambrian  microfossils (see Doubt cast on earliest bacterial fossils April 2003 in Subjects: Geobiology, palaeontology, and evolution). The authors opine that the ecological diversity of freshwater and land systems, and the physico-chemical stress associated with repeated wetting and desiccation compared with the marine domain may have been instrumental in origination of the Eucarya, which should give the Torridonian a scientific reputation that extends beyond these shores.