Category Archives: Geobiology, palaeontology, and evolution

Origin of the arms race

Global paleogeographic reconstruction of the E...

Global paleogeographic reconstruction of the Earth in the early Cambrian period 540 million years ago. (credit:Ron Blakey, Northern Arizona University)

Palaeontologists generally agree on one broad aspect of animal evolution: the central role of predation versus defence in animal diversification to occupy different ecological niches. Indeed that co-relation has to an extent been responsible for the diversification of potentially habitable niches themselves. Armour and arms form a dialectic within the animal world, but one that only rose to dominate when hard materials became an integral part of animal morphology, allowing some to bite, gnaw or rasp and others to develop shelly or horny skeletons. The Kingdom Animalia within the domain of the eukaryotes – organisms based on cells that bear a nucleus – is united by one life style, that of feeding directly or indirectly on other living things. They are heterotrophs unable to generate energy and tissue through the fundamental harnessing of chemistry and physics to use the inorganic world directly, as do autotrophs.  One of the earliest discoveries about the history of animals was that fossils in the widely accepted meaning of the word appeared suddenly in the geological record, earlier rocks containing virtually no tangible signs of life: fossils explode in numbers from the start of the Cambrian Period at 542 Ma. Subsequently, geologists did discover imprints of clearly quite complicated organisms in rocks a few tens of million years older than the start of the Cambrian. But these were flaccid, bag like creatures that recent research has shown to rely on filtering microorganisms from water or directly absorbing organic matter through their skin.

Cropped and digitally remastered version of an...

An animal from the late Precambrian(Photo credit: Wikipedia)

Another feature of sediments of the oldest Cambrian is that in many parts of the world they rest with or=profound unconformity on deformed older rocks of Precambrian age. Throughout Britain the lowest Cambrian rocks are almost pure quartz sandstones that rest upon older more complex rocks ranging from only a few tens of million years older than 542 Ma to some of the oldest rocks in Europe, the Lewisian complex dating back 3 billion years. Many of the hills of North West Scotland have a gleaming white cap of Lower Cambrian quartzite above what has been termed the Great Unconformity where it occurs in Arizona’s Grand Canyon. Sedimentary sequences that continuously record the Precambrian to Cambrian transition and the biological explosion at the juncture are rare. But they show two curious features in sediments that immediately predate those bearing recognisable fossils: a complete lack of evidence for burrowing and millimetre-scale shell-like bodies made of calcium phosphate and carbonate, which are thought to have adorned the skins of otherwise unprotected animals.

Português: Classe Radiodonta

Creatures from the Cambrian Period (credit: Wikipedia)

Calcium, while a very common element is one of the most dangerous to life. Traces are essential for the signalling that goes on in cell metabolism, and too little snuffs out those vital processes.  Yet too much – still a very low concentration in cell cytoplasm – results in the growth of minute mineral crystals within cells, also spelling cell death. This results from the limited solubility of calcium in water, compared with those of other common metals.  At an early stage in evolution cells developed means of restricting the admission of calcium ions and efficient means of expelling excess amounts of calcium. The ubiquitous occurrence of Ca-rich marine limestones throughout the geological record bears witness to two things: the abundance of calcium ions in seawater; a closer look reveals that a great many limestones, going back some 3.5 billion years show traces of biomineralisation that helped form the limey sediments. In the second case, the calcium carbonate in most Precambrian limestones was secreted by photosynthetic blue-green bacteria in minutely thing layers, probably in the form of a slimy film excreted to avoid calcium toxicity. Palaeontologists have long suspected that the earliest skeletal materials formed by animals evolved from the need to excrete biomineralised films by turning a metabolic necessity into functional and integral parts of their body plans: arms and armour. Yet limestones are not rare signs of the presence of a dissolved calcium threat, so why the sudden adoption of waste products in this way?

A fairly old hypothesis is that calcium in seawater must have risen above a threshold that posed toxic threat to all living things and excretion had to increase to maintain the balance, perhaps matched with increasing sizes of animals in the late Precambrian. Only recently has support been found for this suggested evolutionary trigger, initially from analysis of brines trapped in crystalline materials within sediments, such as salt (NaCl). But the very presence of such halite in a sediment is a universally accepted sign of evaporation increasing ionic concentrations in isolated seawater lagoons, whereas a general increase in marine calcium would be needed to present sufficient chemical stress that the whole of animal evolution would require a step-change for survival.  It turns out that support for the hypothesis stems from two isotopic systems most usually associated with dating the formation and weathering of continental  crust: those of strontium and neodymium. The global record of ratios of 87Sr/86Sr and 143Nd/144Nd show unusually large changes in the run-up to the Cambrian Period, the first rising to the highest level recorded in geological history and the second reaching a historic nadir during the Cambrian. This anti-correlation signifies the greatest chemical weathering of older continental crust in the history of the Earth (Peters, S. & Gaines, R.R. 2012. Formation of the ‘Great Unconformity’ as a trigger for the Cambrian explosion. Nature, v. 484, p. 363-366). Not only would this have poured dissolved ions, including those of calcium, into the oceans and raised their concentrations in seawater, but vast areas of the continents would have been eroded to form huge coastal plains, ripe for marine inundation. The last is exactly what the near-universal unconformity at the base of the Cambrian signifies. Presaging this long drawn-out grinding of continents to their gums had been a protracted bout of continental assembly to form the Rodinia supercontinent around 1000 Ma though collision and mountain building. Then Rodinia broke apart, its fragments being driven by plate tectonics to reassemble, along with vast chains of new crust formed in volcanic island arcs, by yet more orogenesis: tectonically high-energy times matched by the processes of denudation on land.

A nice example of planetary interconnectedness on the largest scale with the greatest conceivable consequences, for we vertebrates anyhow. This comes as a great comfort to me in the twilight of my career, since in 1999 I stuck out my neck with a similar concept in Stepping Stones only to meet a suitably stony silence.

A cuddly tyrannosaur

Feathered Dinosaurs 1

Feathered dinosaur Deinonychus (Photo credit: Aaron Gustafson)

Feathered and fluffy dinosaurs in the families that may have led to birds have become almost commonplace, thanks to wonderful preservation in some Chinese Mesozoic sedimentary rocks (see http://earth-pages.co.uk/2003/03/01/flying-feathers/)  and what has become a cottage industry for local people, under professional direction. Most have been small theropods in the Coelurosauria taxonomic branch that span the Jurassic and Cretaceous Periods. The famous Lower Cretaceous Liaoning lagerstätte in NE China recently yielded something truly awesome: three well-preserved specimens of a feathered dinosaur almost as large as the giant tyrannosaurs of the Late Cretaceous (i.e. > 1 tonne in life) (Xu, X. et al.2012. A gigantic feathered dinosaur from the Lower Cretaceous of China. Nature, v. 484. P. 92-95). In fact Yutyrannus huali (‘beautiful feathered tyrant)is a member of the same subgroup as the Upper Cretaceous T. rex and was clearly a top predator in its day. Equally fortuitous is that the three specimens  comprise one with a living body weight of about 1.4 t, the other two being between 500 and 600 kg. Various differences between the largest and the two smaller individuals suggest that thee find represents two generations, the largest perhaps 8 years older than the two smaller ones. All three preserve densely packed filaments suggesting that they were fluffy rather than truly feathered. So why the difference from its probably scaly relative tyrannosaurs from about 50 Ma later?

Around 125 Ma global climate was considerably cooler than the Late Cretaceous greenhouse world, Liaoning probably having mean annual air temperatures around 10°C compared with 18°C late in the Period. Yutyrannus huali and some of its contemporary theropods probably evolved high TOG insulation to ensure all-season sprightliness. It is also possible that a display function was also involved, as seems to have been the case for other dinosaurs.

Mesozoic fleas

Giant Mesozoic fleas from China, 1.4 and 0.8 cm long. From Huang et al. (2012)

Strange as it might seem, rather than bringing to mind the opening pages of Michael Crichton’s Jurassic Park ancient fleas suggest to me Frederick Engels’s Dialectics of Nature (1883). In his lampoon of determinism, which might today be directed at a famous evolutionary biologist, Engels wrote:

‘…last night I was bitten by a flea at four o’clock in the morning, and not at three or five o’clock, and on the right shoulder and not on the left calf – these are all facts which have been produced by an irrevocable concatenation of cause and effect, by an unshatterable necessity of such a nature indeed that the gaseous sphere, from which the solar system was derived, was already so constituted that these events had to happen thus and not otherwise.’

But a paper about fossil fleas from the time of the dinosaurs was always going to catch the eye (Huang, D. et al. 2012. Diverse transitional giant fleas from the Mesozoic era of China. Nature, v. 483, p. 201-204), and that they come from China does have an element of inevitability that arises from that country’s rich endowment with sites of exceptional preservation. The fleas are not at all like the shiny creatures that are so difficult to trap in the fur of a cat’s ear, and they are big: up to 2 cm long. Two species come from Middle Jurassic and one from the Lower Cretaceous. The fascinating thing about fleas, however, is that they evolved to live and thrive in fur and feathers.  This niche is signified by their claws, whose form and articulation avoid entanglement with fibres: which is why cat fleas are so nimble. While cat fleas are flattened laterally to help them slip though fur and have powerful legs that allows them to leap from host to host, the Mesozoic fleas are flat from back to front and are not so leggy.

English: This photo was taken by Andy Brookes ...

Cat flea ~1.5 mm long. Image via Wikipedia

Being so large, it seems unlikely that these Mesozoic fleas would have parasitized mammals that were probably far smaller on average than now. But by the Jurassic fossil evidence, largely from China, shows that dinosaurs had developed feather-like cover. Their evolution itself created a niche occupied thereafter by fleas and other bloodsuckers. They are wingless relatives of flies (Order: Diptera) that first appear in the Triassic fossil record, both thought to have stemmed from more primitive scorpionflies (Order: Mecoptera)

Late Devonian: mass extinction or mass invasion?

A hand made lookalike for User:Dragons flight'...

Image via Wikipedia

The later part of the Devonian (the Frasnian and Famennian Stages) once marked the second largest marine mass extinction (~375 Ma) of the Phanerozoic Eon: it was one of the ‘Big Five’. For the last decade the drop in marine biodiversity in that interval has come under scrutiny: partly because it may have involved several  events;  no well-supported extinction mechanism has emerged; and extinctions seem have been concentrated on three animal groups – trilobites, brachiopods and reef corals. Something large did happen, as reef-building corals almost disappeared and coral reefs only returned with the rise of modern (scleractinian) corals in the Mesozoic. While the end of the Devonian still figures widely as having experienced a mass extinction event, more detailed palaeontological work at the genus and species level suggests another possibility.

‘Officially’ a mass extinction event must exceed the background extinction rate throughout the Phanerozoic and be above that in immediately preceding and following stages: statistically significant, that is. They always give rise to a marked reduction in biodiversity, but another mechanism can do that without extinctions suddenly increasing. The rate at which new species emerge can fall below that of species extinctions, when the overall number of living species falls. As far as ecosystems are concerned both processes are equally severe, but the causes may be very different.

Hederelloids encrusting a Spiriferida brachiop...

Brachiopod from the Devonian of Ohio, USA. Image via Wikipedia

Reviewing detailed records of Devonian species of two genera of brachiopods and one bivalve genus (50 species in all) in five North American stratigraphic sequences, Alycia Stigall of Ohio University, USA noted apparently significant variations in the local assemblages (Stigall, A. L. 2012. Speciation collapse and invasive species dynamics during the Late Devonian ‘Mass Extinction’. GSA Today, v. 22(1), p. 4-9). Speciation overall fell in the Frasnian and the preceding Givetian, while rate of extinction barely changed. For the three studied genera ,speciation reached low values in the Frasnian and Famennian, but that was accompanied by an equally large fall in extinctions. In this narrow sample we seem to be seeing not an extinction crisis but one of biodiversity. Why?

The Late Devonian saw repeated ups and downs in sea level, which repeatedly connected and disconnected shallow- to moderate-depth marine basins. The fossil record shows repeated cases of species from one basin colonising another, each invasion accompanying rapid marine transgression.. One means whereby species arise is through prolonged isolation of separate populations of the ancestral species through independent genetic drift and mutation. The episodic connection of basins may have prevented such allopatric speciation. Interestingly, the invading species  were dominantly animals with a broad tolerance for environmental conditions.

Whether this mechanism applied to all three main animal groups whose diversity plummeted in Late Devonian times remains to be seen, and it begs the question ‘why didn’t it happen among other animal groups that were less affected by whatever the events were?’ One of the problems associated with decreasing biodiversity in modern marine (and terrestrial) settings is growth in the numbers of invasive species, so the work on 375 Ma fossils might help understand and mitigate current ecological issues. The only difference is that for many of the hyper-successful invader species the means of invasion has been provided by human activities. brachiopod brachioopod

Excitement over early animals dampened

Alga (Volvox sp.)

Volvox cyst. Image via Wikipedia

The Neoproterozoic lagerstätte in the Doushantuo Formation in the south of China was until recently thought to be a source of astonishing information about Earth’s earliest animals (See Ancestral animal? in EPN August 2004) that preceded the appearance of those with hard parts at the start of the Phanerozoic.  It contains well-preserved fossils that resemble embryos, algae, acritarchs, and small bilaterians. Dated at between 580 to 600 Ma(See Age range of early fossil treasure trove  in EPN February 2005), the Doushantuo directly overlies cap carbonates representing the emergence of Earth’s climate from a Snowball epoch represented by a tillite beneath the carbonate sequence. A detailed examination using synchrotron X-ray tomography of the putative animal embryos does show clear signs of cell doubling or palintomy (Huldtgren, T. et al. 2011. Fossilized nucluei and germination structures identify Ediacaran ‘animal embryos’ as encysting protists. Science. V. 334, p. 1696-1699) but also internal cell features most likely to be nuclei, but which have no counterparts in animal embryos. The organisms which the fossils most resemble are indeed eukaryotes, but of a kind separate from animals known as Holozoa. Yet there are striking resemblances with eukaryotes more distant from animals, such as the modern Volvox, a type of alga (Butterfield, N.J. 2011. Terminal developments in Ediacaran embryology. Science. V. 334, p. 1655-1656), that developed from an ancestor further back in time than the separation of metazoan animals from holozoans.

Life at the battery terminal

Mussels of species Bathymodiolus childressi (B...

Hydrothermal-vent mussel Bathymodiolus. Image via Wikipedia

Having an interior that is dominated by reducing conditions and oxidising surface environments since free oxygen gradually permeated from its initial build up in the atmosphere to the ocean depths, the Earth has been likened to a massive self-charging battery. Electrons flow continually as a consequence of the nature of the linked oxidation-reduction: in terms of electrons, oxidation involves loss while reduction involves gain (the OILRIG mnemonic). Although there are natural electrical currents, most of the electron flow is in the form of reduced compounds rich in electrons that make their way through the flow of fluids from the deep Earth – effectively an anode – towards the surface  where the reduced compounds lose electrons to create the equivalent of a cathode. Reduction-oxidation (redox) is therefore a power source. Inorganic reactions, such as the precipitation on the sea floor of sulfides from hydrothermal fluids at ‘black smokers’ dissipate energy. Yet the power has considerable potential for organic life. Some bacteria oxidise hydrogen sulfide carried by hydrothermal fluids and others do the same to upwelling methane. In 1977 a teeming biome of worms, molluscs and higher animals was discovered in a totally dark environment around ocean-floor vents. It soon became clear that it could only subsist on chemical energy of this kind, rather than any form of photosynthesis. The key to some metazoans’ success had to be symbiosis with bacteria that could perform the chemical tricks possible in the cathode region of the Earth’s electron flow. There are several candidate compounds: H2S, CH4, NH4, metal ions and even hydrogen gas.

As hydrothermal fluids cycle ocean water into the basaltic crust and underlying peridotite mantle, they not only hydrate the olivines and pyroxenes that dominate the oceanic lithosphere but trigger other reactions one of whose products is hydrogen. As well as a reaction being eyed by those keen on a cheap source of clean fuel, it generates more energy potential for biological metabolism in the guise of hydrogen than those which form other common compound in the returning fluids. Although the nature of hydrogen’s organic use has been elusive, it has now come to light in a surprising guise (Petersen, J.M. and 14 others 2011. Hydrogen is an energy source for hydrothermal vent symbioses. Nature, v. 476, p. 176-180).

One highly successful animal in ocean-floor hot spring systems is a mussel called Bathymodiolus. Genetic experiments by the German-French-US team revealed that a gene known as hupL is present in the mussels’ gill tissue; a gene found in bacteria that use either carbon monoxide or hydrogen as an electron donor. The hupL gene encodes for enzymes known as hydrogenases that are needed to set off the reaction H2 = 2H+ + 2e- that provides electrons needed in bacterial metabolism; a sort of living fuel cell. Hydrogen-using bacteria interact symbiotically with the mussels, which would otherwise be unable to live in the pitch black environment. Genomic sequencing of tube worms and shrimps that occur in the vent communities also contain the bacterial hupL gene. Hydrogenase enzymes are proteins with an iron-nickel core, and probably evolved far back in bacterial evolution around metal-rich hot springs. Interesting as the specific detail of hydrogen-based symbiosis is, the general concept of Earth’s redox systems’ having battery-like behaviour is very useful. On land groundwater sometimes comes into contact with sulfide ore bodies that are oxidised to yield hydrogen and sulfate ions ,while the groundwater is reduced: a battery comes into being with a cathode in the aerated groundwater and electrons flow from the unaltered orebody towards it. Such currents are useful in revealing hidden orebodies using the ‘self-potential’ or SP method. Indeed the downward change from oxidising to reducing groundwater, caused by the redox reactions involved in weathering and soil formation also result in weak negative and positive ‘electrodes’ with a sluggish flow of compounds that bacteria can exploit and thereby encourage metazoan life through symbiosis. In doing so, changes in redox conditions affect the inorganic load of the slowly moving groundwater so that reduced metal ions can be precipitated once they rise into the oxidising horizon. The general enrichment of the upper horizons of soils in iron oxides and hydroxides, and metal depletion in lower horizons probably stem from the ‘Earth battery’ produced by an interplay between inorganic and organic redox reactions. Be on the look-out for more on this topic as the quest for hydrogen fuels becomes more urgent. A former colleague, Gordon Stanger, investigating groundwater in the Semail ophiolite of the Oman for his PhD in the 1970s discovered to his surprise that in outcrops of the mantle sequence there were springs from which hydrogen bubbled freely: fortunately he was not a smoker…

Feathers will fly: Archaeopteryx relegated

Archaeopteryx

A not unimaginative reconstruction of Archaeopteryx. Image via Wikipedia

This year, 2011, is the 150th anniversary of the first Archaeopteryx specimen being unearthed from the famous Solnhofen  limestone lagerstätte. With its feathered, lizard-like tail; two-clawed, stubby wings; a bill-shaped muzzle with teeth but no keratin coating; feet capable of perching and unlike those of small dinosaurs; a ‘wishbone’ and lightweight bones, Archaeopteryx was just the half-and-half missing link in the fossil record so desperately needed to support Darwin’s Origin of Species, published two years beforehand.  It has remained controversial ever since, even having been claimed to be a forgery by such luminaries as cosmologist Fred Hoyle in 1985, despite its superbly preserved intricacies and the existence at the time of 6 slightly different specimens from the same source some discovered long after Hoyle’s supposed master craftsman must have died. Creationists soon after the first discovery claimed it was simply a bird created on a Friday together with fish (Genesis 1:20) and must have predated dinosaurs by a day, as they were created on the 6th Day along with all the ‘cattle and creeping thing and beast of the earth’ (Genesis 1:24-31). That scurrilous sect will certainly leap gleefully on the new discovery of a feathered dinosaur from the ever productive Late Jurassic Tiaojishan Formation in NE China (Xu, X. et al.2011. An Archaeopteryx-like theropod from China and the origin of the Avialae. Nature, v. 475, p. 465-470) because ironically, by itself, it could be said to be a missing link too.

Archaeopteryx lithographica, specimen displaye...

Cast of the first-described Archaeopteryx fossil. Image via Wikipedia

In fact, Xiaotingia zhengi possesses features very like those displayed by Archaeopteryx but convincingly close affinities to deinonychosaurian dinosaurs. The shared features show that neither is a bird (Avialae) and nor are they part of the clade that evolved to birds: they are part of the growing group of feathered dinosaurs that may well have glided or even flown. As Lawrence Witmer of Ohio University has observed (Witmer, L.M. 201. An icon knocked off its perch. Nature, v. 475, p. 458-459), ‘This finding is likely to be met with considerable controversy (if not outright horror)…’. However, Witmer still considers Archaeopteryx to have iconic status, indeed yet more, for its taxonomy and that of its related feathery dinosaurs provides compelling evidence that the origin and evolution of life was a ‘rather messy affair’. Undoubtedly, more feathered creatures hundreds of million years old will be unearthed; it is even possible that further finds will push the beast of Solnhofen back onto its avian perch. Let the celebrations begin!

Added 12 August 2011: Ironically, yesterday the German mint issued a €10 silver coin commemorating the 150th anniversary of the first discovery of Archaeopteryx, artwork of the skeleton with fully fledged arms on the reverse side of the coin compared with the stylised German eagle on the front. This event coincides with the greatest crisis facing the eurozone in its short history, though Germany still retains its ‘triple A’ financial status unlike France and the US. See: http://witmerlab.wordpress.com/2011/01/31/evolution-icon-archaeopteryx-turns-150-this-year-how-are-we-celebrating/

From small beginnings

Camarasaurus, Brachiosaurus, Giraffatitan, Euh...

Some really cool sauropods. Image via Wikipedia

The great vegetarian sauropod dinosaurs, such as Brachiosaurus, were the biggest animals to walk the Earth, weighing up to 100 tonnes, as long as 60 m from snout to the end of their tails and more than 10 m tall. So big, indeed, that even the largest contemporary predators would have been unable to get sufficient purchase with their jaws to do them much damage. This vast bulk, unlike even bigger modern whales, was unsupported by water and would have posed major problems had the sauropods not evolved very porous, low-density neck and tail bones and kept their heads small relative to the rest of their bodies. Such small heads needed to take in up to a tonne of vegetation each day to keep the monsters alive and  ambling. Their teeth are not those of a chewer, being peg- or spoon-like and pointed forwards; specialised for raking in leaves and twigs, swallowed unchewed in great gulps. Once that style of eating developed in their precursors, with no need for massive chewing muscles it became possible to evolve necks up to 15 m long with increasingly diminutive heads. Studies of large numbers of some species of sauropod precursors indicate that juveniles grew astonishingly quickly, essential if their initial vulnerability was to be outpaced; newly hatched they would have weighed little more than 10 kg. At the growth rates of modern reptiles, the largest sauropods would only have reached full size in about a century. The estimated growth rates suggest warm bloodedness, research suggesting that they maintained body temperatures up to 12°C higher than do alligators. Clearly, sauropod dinosaurs were highly specialised, and their evolution is now known to have been lengthy.

A major news feature in Nature (Heeren, F. 201. Rise of the titans. Nature, v. 475, p. 159-161) traces that evolution through several surprising stages. The earliest likely ancestors, which appear in the Late Triassic (~230 Ma), were about the size of a turkey and had teeth adapted for shredding fibrous plant material; other early dinosaurs show clear signs of a predatory lifestyle. There is a limit to the size of predators bound up with the energy balance between flesh consumption and the energy expended in casing down prey and killing them. The limits on the size of plant eaters are mechanical: how much they can stuff in and the strength of their bodies, especially legs. In a world dominated in numbers by predatory dinosaurs, the selection pressure for herbivores to outgrow them and become too big to bite would have been substantial.

Little Triassic Panphagia (‘eater of everything’) was also bipedal, but the fossil record of sauropod precursors clearly shows their growth to the order of 10 m by the Early Jurassic, but not yet a four-legged gait though they had evolved relatively short but sturdy legs, signs of mass-saving porous neck and tail bones, and jaws with a large gape suited to gulping rather than chewing. By the mid-Jurassic Period sauropods were big, strong and four-legged, and by the Cretaceous they reached unmatched dimensions with the titanosaurs. This evolutionary path was not the only one adopted for dinosaurian herbivory. The famous Iguanodon discovered in 1822 by Gideon Mantell in the Early Cretaceous of Sussex was a member of a bipedal group of herbivores, including the duck-billed dinosaurs, that spanned more or less the same time range as sauropods. Fredric Heeren’s article is accompanied by an on-line ‘tour’ of sauropod evolution (go.nature.com/c7zlct), while the American Museum of Natural History has a website for a major exhibition of sauropods (www.amnh.org/exhibitions/wld/ and www.youtube.com/AMNHorg ) that includes footage of  a full-scale animatronic Mamenchisaurus from China which breathes and moves, (Switek, B. 2011. Living it large: review of The World’s Largest Dinosaurs exhibition. Nature, v. 475, p. 172).

Earliest animals from continental environments

Skolithos trace fossil. Scale bar is 10 mm.

Skolithus burrows. Image via Wikipedia

Following closely on discovery in 1 Ga old sediments of the earliest evidence for eukaryote life in continental environments (see Eukaryote conquest of the continents posted June 11, 2011) it seems that metazoan animals colonised non-marine environments earlier than had previously been thought. Up to now most palaeontologists believed that there was a lag of at least 80 Ma between the emergence of marine bilaterian metazoans and their expansion into freshwater, due to a number of physiological hurdles that had to be overcome, such as regulation of trace element chemistry within their cells and bodily fluids. It has been know for more than a century that the first signs of sturdy animals in the marine realm are burrows in tidal sediments that formed more or less at the Cambrian-Precambrian boundary; the earlier sac-like Ediacaran fauna seemed ill-suited to a burrowing or infaunal habitat. A considerable thickness of clastic sediments occur in the Cambrian of eastern California, USA. The earliest are clearly shallow-marine and contain abundant evidence of burrowing. Succeeding them are intensively studied fluviatile sands and silts that have been used a model for sedimentation in the absence of the stabilising influence of land plants. What has been overlooked until recently is evidence for colonisation of the river-laid deposits by burrowing animals (Kennedy, M.J. & Droser, M.L. 2011. Early Cambrian metazoans in fluvial environments, evidence of the non-marine Cambrian radiation. Geology, v. 39, p. 583-586).

The burrows include the vertical U-shaped forms given the name Arenicolites, which is the most common trace fossil, simple vertical tubes (Skolithus) and horizontal, meandering tubes with furrowed sides (Psammichnites). Anyone who has seen the Early Cambrian Pipe Rock of NW Scotland will also have seen these trace fossils, yet the Pipe Rock shows evidence of tidal deposition and is shallow marine. Their non-marine equivalents in California are coeval with the earliest known trilobites in the Cambrian marine sequence. It seems that whatever the burrowing animals were, they easily overcame any physiological or environmental barriers to adopting a life in freshwater, encouraged by the ready sustenance that terrestrially adapted acritarchs and cyanobacteria had provided for half a billion years previously.

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.