Mitochondria, oxygen toxicity and the quahog

One of the many crises through which life passed during its evolution was the widespread appearance of oxygen.  This occurred once the release of soluble iron-2 to the oceans from sea-floor processes fell below a rate that buffered the photosynthetic generation of oxygen through the precipitation of iron-3 oxides in marine sediments.  Oxygen is life-threatening, largely through its encouraging the formation of simple compounds that are more potent oxidizers than oxygen (O2) itself, such as O, H2O2 and HO.  In cells they can lead to genetic degeneration, progressive ageing and eventually cell death.  Free oxygen in the environment was a stealthy threat to all life forms that existed around 2200 Ma.  A possible evolutionary response that may have opened the way for the later rise of the Eucarya, and the huge diversification that permitted, is nicely summarized by Doris Abele in Nature of 7 November 2002 (Abele, D. 2002.  The radical life-giver.  Nature, v. 420, p. 27).  The main strand of her argument is that mitochondria, the energy converters in eukaryote cells, also serve to keep oxygen levels inside cells high enough for metabolism, yet low enough to minimise the formation of threatening oxidants.  Her object of study has been the noble ocean quahog, Arctica islandica (incidentally, a clam often referred to by Herman Melville in Moby Dick) which mysteriously burrows into anoxic muds for a while and drops its metabolism alarmingly.  By this habit, the quahog has achieved what middle-aged Californians yearn for; spectacular life extension to as much as 220 years.  Abele believes that this protective function of mitochondria  is deployed by the quahog, having arisen in the earliest Eucarya, after the oxygenation of the planet.  However, as Lyn Margulis observed in developing her endosymbiotic hypothesis for the emergence of eukaryotes, mitochondrial RNA is very like that of oxygen-respiring purple bacteria.  Anti-oxidant mechanisms may therefore be more ancient.  The other main defence against free radicals takes the form of a range of vitamins and other complex compounds, some of which seem to have their origins in heat-shock proteins; possibly harking back to life’s origins near deep-ocean hydrothermal vents.

In a similar vein, linked to the rise of oxygen concentrations, doubt has been cast on the role of photosynthesising cyanobacteria since the earliest times..  Most geologists hold them responsible for creating stromatolites since 3500 Ma, and also for providing an early source of oxygen that was rapidly scavenged by the precipitation of iron oxides in banded iron formations.  Carrine Blank, a palaeobiologists at Washington University in St Louis, has genetically compared cyanobacteria with a range of other living Bacteria, to asses their relatedness.  Her work suggests that the blue-greens were late additions to early life, perhaps long after the first BIFs appeared (report on the annual meeting of the Geological Society of America, in New Scientist 9 November 2002, p. 25).

Climate change data from satelites

Among the many regions of the electromagnetic spectrum, that spanning microwaves most easily penetrates the atmosphere.  Most people are familiar with radar images, produced by actively illuminating the Earth’s surface with microwaves.  However, the Earth also radiates microwaves, depending on surface temperature.  A great deal of information can be gleaned from this upwelling radiation, about surface temperature and at different levels in the atmosphere, rainfall, wind speed and soil moisture.  NASA, in partnership with several US universities, has launched public access to passive microwave data from the Tropical Rainfall Measuring Mission and NOAA-15 satellites at pm-esip.nsstc.nasa.gov .  Data go back 5 years, and include comparisons of daily air temperatures with the 20-year average.

Snowball Earth hypothesis challenged, again

Palaeomagnetic data from localities famed for their Neoproterozoic glaciogenic rocks point persuasively to several epochs between 750 and 550 Ma when widespread continental glaciation took place at low latitudes.  It is this evidence, along with theoretical consideration of drastic changes in the Earth’s albedo that would result from tropical land ice, that encouraged the idea of pole to pole ice cover.  Only a build-up of volcanogenic CO2 in the atmosphere could prevent such a “Snowball Earth” lasting indefinitely, and even with such relief it would have endured for millions of years.  Much of the geological evidence cited by those who support and promote this neo-catastrophic idea comes from excellent, but geographically quite limited occurrences of tillites or glaciomarine sediments, such as those of Namibia.  Some occurrences have never been seriously analysed, except as examples that superficially support the hypothesis.  One such sequence is that of Arabia, easily accessed in northern Oman and described by a British-Swiss team (Leather, J. et al. 2002.  Neoproterozoic snowball Earth under scrutiny: Evidence from the Fiq glaciation of Oman.  Geology, v. 30, p. 891-894).

Isotopic studies of carbonates from glaciogenic sediments (see Meltdown for Snowball Earth? in Earth Pages News for February 2002) seriously undermined several arguments by “Snowball Earth” supporters, but are open to various interpretations.  Hard geological evidence is less easy to rationalize.  A growing number of  Neoproterozoic glaciogenic sequences, such as the Port Askaig Tillite of the Scottish Dalradian Supergroup and others from the Congo and Kalahari cratons, and Laurentia, show dropstone-rich diamictites interbedded with sediments that show little if any sign of a glacial influence (Condon, D.J. et al. 2002.  Neoproterozoic glacial-rainout intervals: Observations and implications.  Geology, v. 30, p. 35-38).  Such evidence can be explained by climatic change and a fully functioning hydrological cycle.  The report on the Omani example by Leather and colleagues highlights splendid examples of sediments that mark cycles of glacial advance and retreat, reminiscent of those of the Pleistocene glacial epoch and more or less the same as in many Neoproterozoic occurrences.  It can only be a matter of time before Australian geologists enter the fray decisively, for glaciogenic sediments comprise up to 30% of the many-kilometres thick Umberatana Group in the Neoproterozoic of the Flinders Range in South Australia, and there are several other stratigraphically distinct diamictite sequences.

It seems likely that the “Snowball Earth” hypothesis is waning; an embarrassment for those geologists who have promoted it so assiduously over the last several years.  However, the enigma of low-latitude glaciation on a vast scale is likely to remain, unless, that is, all the diamictites can be shown to have non-glacial origins, which is not as unlikely as it might seem.  The Fiq sequence of Oman, like the Dalradian example in Scotland, formed in an actively extending basin.  Repeated seismicity on rift-bounding faults could have launched debris flows to deposit diamictites (a purely descriptive term for sediments containing a wide variety of clast sizes).  The most spectacular diamictite in the Dalradian Supergroup, and perhaps anywhere, is the Great Breccia of the Garvellachs.  Recent work suggests strongly that it is not glaciogenic, but the product of such a debris flow (Arnaud, E. & Eyles, C.H. 2002.  Catastrophic mass failure of a Neoproterozoic glacially influenced continental margin, the Great Breccia, Port Askaig Formation, Scotland.  Sedimentary Geology, v. 151, p. 313-333).  The supposedly clinching evidence for diamictites’ origin from iceberg armadas is the way in which some clasts (“dropstones”) puncture underlying stratification.  All that is required is a means of puncturing, and sediment compaction around large, resistant clasts in a water saturated matrix is quite capable of doing that.  Even the long-held belief that glaciation is uniquely signified by polished and striated surfaces beneath diamictites containing similarly scratched clasts is coming into question.  Sites of large impacts, such as the Ries crater in Germany, include exactly similar features caused by ejecta blasted from the crater, cited by Vern Oberbeck, formerly of NASA, in a little-cited paper that proposed an impact origin for diamictites (Oberbeck, V.R. et al. 1993.  Impacts, tillites and the breakup of Gondwanaland.  Journal of Geology, v. 101, p. 1-19).

Post-apocalypse weathering in the Early Triassic

Environmental crises do not come bigger than that at the end of the Permian, when marine ecosystems virtually collapsed, and similar extinctions of terrestrial flora and fauna are becoming clear.  Whereas the Siberian Traps may indeed have been a triggering mechanism, there are carbon-isotope indicators that vast amounts of methane entered the atmosphere shortly afterwards, rapidly being oxidised to CO2.  The density of respiratory openings (stomata) in fossil leaves from the lowest Triassic is unusually low, indicating an abundance of CO2 in the atmosphere and probably enhanced “greenhouse” conditions.  Hot and humid conditions encourage weathering of the continental surface, and there are many Early Triassic palaeosols, some which mimic those in the tropics being found at unusually high palaeolatitudes.  Such soils harbour crucial evidence for surface conditions, and the high-latitude ones present a surprise (Sheldon, N.D. & Retallack, G.J. 2002.  Low oxygen levels in earliest Triassic soils. Geology, v. 30, p. 919-922).  Unlike tropical laterites, which are rich in kaolinite, high-latitude soils are dominated by illitic clays that signify incomplete breakdown of silicates.  The surprise comes in the form of an unusual mineral, berthierine; a green, serpentine-like mineral that is easily confused with chlorites in hand specimen.  It can form by reaction between clays and ferric oxy-hydroxides, but only under highly reducing conditions.  Because most soils since about 2000 Ma ago have formed in contact with an increasingly oxygen-rich atmosphere, achieving suitably reducing conditions demands input of a reductant to the soil “atmosphere”.  The most likely candidate is methane, whose oxidation would consume oxygen.  However, methane’s residence time in the air is around 10 years, because it is quickly oxidised to CO2, so methane release following the P-Tr boundary event seems as if it was sufficiently prolonged to influence considerably longer term soil formation.

More confusion over Bangladesh arsenic crisis

Millions of Bangladeshi people risk arsenic poisoning if they drink water drawn from tube wells (see British Geological Survey sued over arsenic, October 2002 Earth Pages News).  Since the disaster first came to light, UNICEF have tested 1.3 million of the estimated 10 million tube wells that are potentially hazardous.  Those deemed safe are painted green, while those which are risky are now red.  Unfortunately, doubts are being cast on the reliability of the commercial test kits that UNICEF use to estimate dissolved arsenic concentrations.  It is claimed that the analytical method have never been validated by controlled field experiments, and also that the minimum level of arsenic that they can detect is ten times higher than the safe level set by the WHO.  A positive contribution to solving the problem is to drill deeper, since it seems as if the condition for release of arsenic from bonding in sedimentary iron minerals is related to bacterial action that creates reducing conditions.  Although deep by comparison with traditional hand-dug wells, the tube wells go down only 50 to 80 metres and do not penetrate the zone in which reducing bacteria survive.

Source:  Pearce, F. & Hecht, J. 2002.  Flawed water tests put millions at risk.  New Scientist, 16 November 2002, p, 4-5.

Seismic bathymetry and Mediterranean debris flows

Tsunamis are an ever present threat in coastal areas, and can be set in motion by submarine debris flows as well as by earthquakes.  As more evidence for ancient tsunamis emerges on coastlines, such as characteristic features in Alaska (seismically induced), the Hawaiian islands and Bahamas (induced by landslips on unstable volcanic islands), and even the east coast of Britain (submarine debris flow off western Norway) their perceived threat has grown.  A team of oceanographers from Spain, Canada, Belgium, Britain and France has re-examined seismic reflection data from the western Mediterranean, to extract detailed topography of the sea floor (Lastras, G. and 6 others 2002.  Seafloor imagery from the BIG’95 debris flow, western Mediterranean.  Geology, v. 30, p. 871-874).  Although the western Mediterranean is seismically quiet, compared with around Italy and Greece, it is floored by products of turbidity flows.  A particularly large example (BIG’95) off the Spanish coast has an estimated volume greater than 26 km3.  Lastras et al. provide exceptional detail of the internal structure and surface shape of this debris flow, which enables them to suggest how it formed.  It coincides with an interface between deep volcanic rocks and a thick cover of soft sediments, along which gradual detachment eventually resulted in a normal fault propagating to the sea floor.  The mechanical instability seems to have been due to rapid deposition from the Ebro river system at a time of low sea level in the Mediterranean around the beginning of the Holocene.  The flow is marked by fluid escape structures, which the authors suggest may have been connected with a rise in bottom-water temperatures.  Is this another example of gas hydrate being involved, as seems likely for the Storegger slide that caused tsunamis along Britain’s east coast (see Collapsing islands, March 2002 Earth Pages News) about 7200 years ago?  The authors do not speculate on that.  However, the detail that they provide about the conditions that culminated in BIG’95 should provide a benchmark for seeking areas prone to such massive and potentially catastrophic events.

Changing composition of seawater

Using carbonate sediments and fossil shells to assess how the composition of seawater has changed is a long-standing technique in sedimentary geochemistry.  Isotopes of strontium and oxygen have provided revolutionising windows on the pace of continental weathering and fluctuations in sea-surface temperature and continental ice cover for over 30 years.  The magnesium to calcium ratio in fossil shells has given insights into deep-water temperatures for the Cenozoic, more recently.  However, tracking changes in the bulk composition of seawater through time, through analyses of carbonates, is plagued by the continual chemical interaction between rocks and the waters with which they are in contact.  The Mg/Ca ratio of sea water is a potential proxy for the amount of hydrothermal activity on the sea floor, and thus the rate of sea-floor spreading.  This is not because oceanic basalts are magnesium rich compared with continental crust that provides much of the dissolved matter that enters the oceans, but because hydrothermal reactions tend to mop up dissolved magnesium and release calcium..  Unfortunately, magnesium also easily replaces calcium in carbonates during diagenetic processes, particularly dolomitisation.  There are two means of overcoming this hindrance, by analysing seawater trapped as fluid inclusions in evaporite minerals and the shells of echinoderms that still contain minute structures formed in life and are unlikely to have been altered (Dickson, J.A.D. 2002.  Fossil echinoderms as monitor of the Mg/Ca ratio of Phanerozoic oceans. Science, v. 298, p. 1222-1224).  Early results seem to match a prediction that while supercontinents existed, the length of mid-ocean ridges and therefore ocean floor hydrothermal activity were at a minimum.  Around the Precambrian boundary and during the Carboniferous to Jurassic periods, Mg/Ca was high at the time of the Vendian and Pangaea supercontinents.  During major bouts of continental break up – the Lower Palaeozoic and Mesozoic – the ratio is low.  Oddly, the ratio has risen to unprecedented high levels during the Cenozoic Era, when clearly there is high hydrothermal activity.

Despite the fact that the Mg-Ca record of the oceans is limited to just a few short time spans in the 545 Ma record of the Phanerozoic, plenty of geochemists and palaeobiologists are speculating about the possible consequences for evolution of changes in the bulk composition of seawater.  There have been major swings in the proportion of calcite to dolomite in carbonate sediments throughout geological time (see Bacteria and dolomites, January 2001 Earth Pages News).  Discussion now centres on the possible effect of changing Mg/Ca ratios on the waxing and waning of important carbonate secreting organisms, ranging from corals and molluscs that build reefs to the minute coccoliths that formed the Cretaceous Chalk.  Perhaps different groups responded differently to changing water composition, and maybe the Cambrian Explosion of shelly faunas was triggered somehow by a critical shift in the ratio.

See also: Kerr, R.A. 2002.  Inconstant ancient seas and life’s path.  Science, v. 298, p. 1165-1166

Deep carbon cycling, and gold mineralization

One of the more speculative aspects of the carbon cycle concerns the fate of carbonate sediments that descend subduction zones.  One popular hypothesis, with an acronym that is likely to amuse colloquially inclined, British readers (the BLAG model named after its three originators Berner, Lasaga and Garrels) avows that such carbonates contribute to CO2 emissions from volcanoes above subduction zones by reacting with silica.  The presence in blueschists of abundant aragonite associated with silica suggests that if that does happen, not all carbonate is consumed and a great deal enters very long-term storage in the mantle.  Indeed, aragonite-magnesite associations are stable to pressures that are equivalent to depths of 240 km.  Rocks formed under exceptionally high-pressure conditions, which might shed further light on the deep part of the carbon cycle, are exceptionally rare.  One such occurrence is the Kokchetav massif of Kazakhstan, in which dolomitic marbles accompany eclogites.  Notable for the occurrence of metamorphic diamonds, Kokchetav rocks probably equilibrated deeper than 250 km, so the carbonates are particularly interesting.  Yongfeng Zhu and Yoshihide Ogasawara of Beijing University in China and Waseda University in Japan have found evidence for dissociation of dolomite in them (Zhu, Y. & Ogasawara, Y. 2002.  Carbon recycled into deep Earth: Evidence from dolomite dissociation in subduction-zone rocks.  Geology, v. 30, p. 947-950) during reactions that generate garnet and clinochlore.  The mineral textures reveal equilibria that involve the production of carbon and oxygen, rather than CO2, so it is quite possible that reflux of CO2 from subduction zones to the atmosphere may not be as significant as the “BLAGgers” suppose.

Interestingly, the same issue of Geology includes a paper on the geochemical conditions under which gold and copper enter subduction-zone magmas to source major ore deposits (Mungall, J.E. 2002.  Roasting the mantle: Slab melting and the genesis of major Au and Au-rich Cu deposits. Geology, v. 30, p.915-918).  Mungall focuses on the inability of chalcophile metals to enter magmas when sulphides are stable in the mantle.  Under those condition Au and Cu tend to enter sulphide melts whose density and immiscibility separate them from silicate melts.  Oxidation of sulphur is needed to overcome this tendency, and that requires high oxygen fugacity at the depths involved, suggested by him to accompany abundant iron-3 in the subducted materials.  That may be so, but release of molecular oxygen by high-pressure carbonate dissociation, as described by Zhu and Ogasawara, seems an even more likely means of freeing chalcophile metals to magmas.

Slab pull versus subduction suction

The dominant forces that drive plate tectonics are those created by subduction.  Slab pull is transmitted throughout a plate system when subducted oceanic lithosphere remains mechanically attached to its parent plate.  However, detached slabs that descend into the mantle, excite viscous flow that might exert traction on the base of the lithosphere, thereby sucking plates along.

This item and others about Tectonics can be read at Earth-logs in the Tectonics archive for 2002

Florida Department of Environmental Protection – www.dep.state.fl.us/geology/

The department’s site combines high-quality educational media and scientific data about those environmental aspects of Florida that are unique.  You can access downloadable AutoCAD *.dxf files showing geological maps from the county level to smaller scales plus other GIS files, lithological logs from boreholes and detailed information on the State’s oil and gas industry.  Currently featured on its home page is a related web site about Florida’s unique hydrogeology and its famous springs.  A well-designed, easy to use site.

A considered view

Find after find of hominid remains (Bonanza time for Bonzo – August 2002) undoubtedly forces physical anthropologists to reflect on what their still tiny collections of fossils might signify about the descent of humans.  There are two ways of looking at that; as a “tidy” tree and one that is essentially “untidy”.  The first seeks a means of connecting the earliest remains to later ones by the simplest possible connections – a touch of Occam’s Razor.  However, more diversity and ever increasing ranges of ages and localities for the remains inevitably challenges this kind of palaeontological “good housekeeping”.   Bernard Wood of George Washington University has long regarded evolution as untidy, and the finds of Sahelanthropus tchadensis and Orrorin tugenensis, around 6 to 7 Ma old, reinforce his trenchant views (Wood, B. 2002.  Who are we?  New Scientist, 26 October 2002, p. 44-47).

Because the genetic similarity between humans and their nearest relatives, chimpanzees, seems to suggest that the two clades diverged between 5 and 10 Ma ago, Sahelanthropus and Orrorin may be pretty close in age to that division.  But what were they?  Wood’s view is interesting, and a worry to the advocates of a parsimonious set of connections.  Connectivity in proposed clades rests, for obvious reasons, on purely physical characteristics.  There are many examples from the fossil record of animals whose outwardly similar characters, for example those shared by sharks and dolphins, do not signify inheritance from common ancestry.  This is homoplasy, and raises the awkward possibility that special characters, regarded as essentially human, need not have arisen only the once and been carried by linear descendants.  The often quoted “golden characters” of big brains and upright gait, that confer an opportunity to develop consciousness through freeing of the hands, may well have arisen more than once.  The truly odd thing about Sahelanthropus is just how “modern” its face looks.  Beetling brows, thick jaw and un-apelike canine teeth would put it on a sort of par with fossils of species of Homo that arose 4 to 5 million years later.  Yet none of the fossils in between have this combination.; in the “tidy” scheme of things they are more “primitive”, and “therefore” cannot be our ancestors.  Quite a muddle! Faces, the most sought after bits of bone, isolated in time and place could well have led many up the proverbial garden path.  Why, suggests Wood, shouldn’t early hominids have been dead ends morphologically, with “primitive” characters making repeated comebacks?  Why, too, shouldn’t they have been ancestral chimps, or even neither chimp nor human?  The dearth of late-Miocene and Pliocene non-hominid fossils of primates leaves all this as possible.  He reckons the search for “missing links” has always been a non-starter.  Whatever, by expanding enormously the area of potentially fruitful ground from the narrow confines of the East African Rift, the Sahelanthropus find in Chad may yet lead to a big increase in the number of hominid and other primate fossils over which physical anthropologists can ponder.

Africa’s first ice core record

Melting of low-latitude glaciers in Africa is so rapid that, unless they are cored soon, their content of long-term climate data may soon be gone forever.  So the first detailed isotopic record from Africa’s highest glacier on Kilimanjaro is cause for some relief.  Intrepid glaciologist Lonnie Thompson welded a large team together for this important task (Thompson, L. 2002. Kilimanjaro ice core records: evidence of Holocene climate change in tropical Africa.  Science, v. 298, p. 589-593).  The annually layered ice goes back only about 12 ka, but nonetheless gives a precious account of climate change at the heart of the continent, far more detailed than sparse lake-bed cores from various places.

The core confirms a broad pattern of warm, wet conditions from 11 to 4 ka, before the long-term cooling and drying of historical times.  These reflect likely weakening of monsoonal conditions in the late Holocene.  However, assigning precise ages to depth in the cores is not as easy as in those from high-latitude ice sheets, because of a lack of good layering (presumably) and dateable carbon.  At about 5200 years ago, the record shows an abrupt fall in d18O, a sign of drying and cooling that took place over perhaps a matter of decades.  This correlates with disruption of early civilisations in India, Egypt and the Middle East, and probably stemmed from cooling in the North Atlantic.  However, an equally rapid deterioration occurred around 6300 years bp, although not so extreme, to presage a millennium of arid conditions at the heart of Africa.  Important as these data are, the team’s estimates of current retreat rates of the Kilimanjaro glaciers are alarming.  Quite probably, the white cap of Africa’s highest mountain will have disappeared within the next 20 years.

Lonnie Thompson is obviously both keyed- and clued up about extracting climatic data from ice at high elevations.  So much so, that Science has printed a lengthy account of his exploits, mainly on low-latitude glaciers (Krajick, K. 2002.  Ice man: Lonnie Thompson scales the peaks for science.  Science, v. 298, p. 518-522

Land plants at the P-Tr boundary

The Permian to Triassic transition involved a transformation from globally cool conditions to a hothouse, as well as the largest mass extinction in the fossil record.  It also spanned a time when most continental lithosphere was clumped in the Pangaea supercontinent.  In the case of plants, it is not easy to sort the effects of climatic shifts from those due to catastrophic events, either the effects of the huge Siberian flood-basalt event (see Earth Pages August 2002, Flood basalts of Siberian Traps doubled at a stroke) or a yet to be proven impact.  Allister Rees of the University of Chicago has painstakingly organised global Permian and Triassic floral data to see if the changes were slow (climatically influenced) or sudden ( possible evidence for a catastrophic collapse),a nd if they coincide from region to region.  He found that in some regions big changes happened quickly around the P-Tr boundary, but in others the shifts were protracted and unrelated to faunal extinctions (Rees, P. McA. 2002.  Land-plant diversity and the end-Permian mass extinction.  Geology, v. 30, p. 827-830).  This clearly implies caution in the interpretation of detailed local records as signs of massive events, and also points out the need to place such records in the contexts of global climate belts and biases that result from varied degrees of biotic preservation.

Dinosaurs did urinate

News is coming in (New Scientist, 19 October 2002, p. 26) of a startling find along a dinosaur trackway in Colorado.  At the October meeting of the Society of Vertebrate Palaeontology, Katherine McCarville of the South Dakota School of Mines and Technology described a bath-sized pit preserved among sauropod footprints.  Seemingly, all the evidence points to it having been excavated by a gargantuan stream of liquid pouring from above.  Ranking as a candidate for the IgNobel Awards of 2003, this evidence for dinosaurian bladder relief may shake the theory that birds are descended from dinosaur ancestors; birds do not urinate.

Continents colonised a billion years ago

The Torridonian of NW Scotland is a thick sequence of mainly terrestrial sediments that accumulated on the Laurentian craton, between 1200 and 1000 Ma ago.  Much of the sequence evidences braided-stream deposition, with brief lacustrine episodes.  Any geologist who examines these mainly siliciclastic rocks will find abundant evidence for subaerial conditions in the form of desiccation cracks, often affecting directional current ripples.  However, it takes a keen eye and some knowledge of biofilms to spot any signs of microbial activity.  In sandstones they manifest themselves by having increased the normally very low cohesiveness of wet sand by their binding action (Prave, A.R. 2002.  Life on land in the Proterozoic: evidence from the Torridonian rocks of northwest Scotland.  Geology, v.  30, p. 811-814).  Prave analysed the shapes of desiccation polygons to show that the Torridonian sands were unusually cohesive, and recognised other features likely to have been formed by microbial crusts.  These finds add to the growing evidence for substantial terrestrial biomass, long before the “official” colonisation by land plants in the Silurian and Devonian.  Whether or not such an expansion of the biosphere added significantly to carbon burial and drawdown of atmospheric CO2, as it did in post-Silurian times, remains to be determined from average carbon contents of quite rare Precambrian terrestrial sediments,