An incomplete history of the fossil record People have been finding fossils for a very long time, and by the 1950’s, by using some new techniques and instruments, scientists began finding what are probably the oldest fossils possible. They are about 3,850,000,000 years old. To discover why these are the oldest fossils possible, please read the text which accompanies the images below. Source: (left) http://www.fossilmuseum.net/Tree_of_Life/Stromatolites.htm (right) http://www.blackwellpublishing.com/ridley/image_gallery/Precambrian_jellyfish.asp Stromatolites are the oldest fossils we have discovered. It must be remembered that because of plate tectonics (i.e. continental drift – the subject of another post), very old fossils are often destroyed as old bits of the earth’s crust (from the Archaean era) are more likely to have been subducted by the movement of the earth’s plates, and to have melted in the heat, in addition, there are doubtless countless fossils which have been eroded into nothingness by wind and water, or otherwise lost to us, so beyond a certain point, the older they are, the rarer the fossils which survive become. Also, very old life-forms were all simple single-celled or multicelled organisms, and these, because they have no hard parts, only fossilize under exceptionally ideal conditions. So, what does the fossil evidence tell us? All of the oldest fossils we have found (between about 3,850,000,000 years old, and about 2,500,000,000 years old) are fossils of simple single-celled organisms, rather like bacteria, called prokaryotes (these are mostly single-celled organisms without a nucleus). From this, we can conclude that for a bit over 1,000,000,000 years, the only life-forms on the planet were these simple organisms. (Now, I would just like to put that in perspective: from the Cambrian revolution, when life diversified like mad to now is only half that time; from the start of the dinosaurs to now is only a quarter of that time, and the from the very first proto-towns to now is one hundred thousandth of that time.) In other words, the simple single-celled forms had our earth all to themselves for an enormously long time indeed. This has some interesting implications for areas like exobiology, and SETI (the search for extraterrestrial intelligence). Presuming, as I do, that life is probably fairly commonplace in the universe, and that there are loads of large lumps of rock (planets, and perhaps moons etc) ‘out there’ with life on them, the chances are quite high that these are going to be simple single-celled forms, with whom we will not even be having involved conversations – let alone alien sex (tabloid revelations to the contrary notwithstanding)! Source: http://library.thinkquest.org/C004535/media/prokaryote.gif A prokaryote. Below are some photographs of simple bacteria: Source: http://www.biology4kids.com/extras/show_kingdoms/03.jpg Around 3,500,000,000 years ago, some of these life-forms began producing a chemical called chlorophyll (which as you know, is green in colour, and is found in most plants today). This chemical allowed these organisms to make their own food, and this was a very important ability, as we shall shortly see. Source: http://kentsimmons.uwinnipeg.ca/16cm05/16lab05/lb1pg2_files/image007.jpg Cyanobacteria structure. Source: http://universe-review.ca/I11-30-cyanobacteria.jpg Some examples of chlorophyll-containing bacteria, the cyanobacteria. At some point, these simple single-celled organisms also clumped together, and eventually became multi-cellular organisms. This however is a complicated story. First of all, we have to define what exactly we mean by a multi-celled organism: At the most basic level, we have the eukaryotes. Eukaryotes differ from prokaryotes (see above) in that their cells contain membranous sacs called organelles, including mitochondria, chloroplasts, and the nucleus. They look rather like they were assembled from a variety of different cells, and as a result, many scientists think these organelles are descended from formerly free-living prokaryotic organisms which banded together. Thus, many important functions of eukaryotic cells, such as photosynthesis, and respiration (the process by which organisms use oxygen to metabolize organic compounds to produce energy, giving off carbon dioxide) may well have been acquired through a symbiosis of previously independent forms of life. Source: http://www.fas.org/irp/imint/docs/rst/Sect20/celltypes.jpg If this scenario is true, then Eukaryotes were the first multi-cellular organisms because they were originally two (or more) organisms which somehow joined together to become one symbiotic living entity (though we are not yet sure how exactly this may have happened). Anyway, though we do not know enough about this, we do know that eukaryotes flourished as the environment became richer in oxygen (which was generated by photosynthesizing organisms). Perhaps this was in part due their more complex intracellular function. Source: http://cosmology.net/images/AcritarchFossil9.jpg An Acritarch fossil, one of the earliest multi-cellular organisms. This however is only one version of multi-cellularity. Organisms also began grouping together, first into simple agglomerations, and then into interdependent ones, and finally in coordinated forms, and these were all different ways of being multi-cellular. The oldest known possible multi-cellular eukaryote is Grypania spiralis (see immediately below), a coiled, ribbon-like fossil two millimetres wide and over ten centimetres long. It looks very much like a coiled multi-cellular alga and has been identified in banded iron formations found in what is today Michigan which date to 2,1000,000,000 years ago. Source: http://www.fas.org/irp/imint/docs/rst/Sect20/pmg31.jpg Grypania may, or may not, be a eukaryote, but another, unrelated colonial eukaryote, Horodyskia (see below), is known from sedimentary rocks which are 1,500,000,000 years old, and which were found western North America as well as in rocks more than 1,000,000,000 years old, which were found in Western Australia. Source: http://macroevolution.narod.ru/fedonk1.jpg Some of earliest known eukaryote fossils are those of acritarch, shown directly above, which date from around 2,100,000,000 years ago. In fact, acritarchs are the most common fossils of the late Proterozoic. The earliest known extended multi-cellular animals are to be found amongst the Ediacaran fauna (see directly below), which are named for the Ediacaran hills of South Australia: Some of these Ediacaran animals resemble modern jellyfish and segmented worms, found in great numbers in the seas today. Others are unlike any known organisms and cannot be classified with any certainty. One feature they have in common is that without exception, they all lack the rigid, supporting skeletons and protective shells that characterize the first fossils of the Cambrian Period (see further below). Source: http://arjournals.annualreviews.org/na101/home/literatum/publisher/ar/journals/production/ earth/2005/33/1/annurev.earth.33.092203.122519/images/medium/ea330421.f2.gif Anyway, getting back to the story of the development of photosynthesis, perhaps around 2,000,000,000 years ago, the ability to synthesize their own food from sunlight and carbon dioxide (along with some minerals) allowed certain organisms to eventually leave the water and to live (albeit initially only for short periods, but eventually permanently) whilst exposed to the air (in other words, to live on the dry land), which up to this time had been totally uninhabited; there were neither animals nor plants until this colonisation of the land by early organisms (which initially were algae and possibly bacteria) began. Source: http://www.uic.edu/classes/bios/bios104/mike/mycorr.gif Fossil of an early terrestrial algae. These plants probably resembled the rather slippery algae which one sees today growing on rocks which are periodically exposed by the tides. Although I write that the incorporation of chlorophyll allowed some plants to leave the water, the story, in fact, is somewhat more complex than I am letting on: A likely mechanism by which plants moved onto dry land is that some of these early plants which grew underwater, but near the water’s edge would have been exposed by tides or by the drying out of the pond / pool in which they were growing, or possibly by changes in water-levels due to some other factor. This would have forced them to eventually evolve structures which made it possible for them to adapt to full terrestrial living – and this is exactly how natural selection works. Growing in water is a very different proposition from growing in air, and is much easier, at least in physiological terms: In water, you do not have to worry about being dehydrated (although growing in salt water can bring on its own special problems, especially if your salinity balances are not right), and you do not need to worry about physically supporting your own weight. Once out of the water, you need thicker, less permeable and stronger cell-walls to address both of these problems; Also, once you are out of the water, there are not likely to be many nutrients freely floating around, and available for you to consume, thus the ability to synthesize your own food out of carbon dioxide, water, minerals and sunshine, via the intermediation of chlorophyll would have been of incalculable use on land (for more on this, see boxed section below). The Silurian land was populated by early land plants as well as a variety of insects. Both plants and animals had a number of challenges when they moved from the water to land. 1. Drying out. Once removed from water and exposed to air, organisms must deal with the need to conserve water. A number of approaches have developed, such as the development of waterproof skin (in animals), living in very moist environments (amphibians, bryophytes), and production of a waterproof surface (the cuticle in plants, cork layers and bark in woody trees). 2. Gas exchange. Organisms that live in water are often able to exchange carbon dioxide and oxygen gases through their surfaces. These exchange surfaces are moist, thin layers across which diffusion can occur. Organismal response to the challenge of drying out tends to make these surfaces thicker, waterproof, and to retard gas exchange. Consequently, another method of gas exchange must be modified or developed. Many fish already had gills and swim bladders, so when some of them began moving between ponds, the swim bladder (a gas retention structure helping buoyancy in the fish) began to act as a gas exchange surface, ultimately evolving into the terrestrial lung. Many arthropods had gills or other internal respiratory surfaces that were modified to facilitate gas exchange on land. Plants are thought to share common ancestry with algae. The plant solution to gas exchange is a new structure, the guard cells that flank openings (stomata) in the above ground parts of the plant. By opening these guard cells the plant is able to allow gas exchange by diffusion through the open stomata. 3. Support. Organisms living in water are supported by the dense liquid they live in. Once on land, the organisms had to deal with the less dense air, which could not support their weight. Adaptations to this include animal skeletons and specialized plant cells/tissues that support the plant. 4. Conduction. Single celled organisms only have to move materials in, out, and within their cells. A multi-cellular creature must do this at each cell in the body, plus move material in, out, and within the organism. Adaptations to this include the circulatory systems of animals, and the specialized conducting tissues xylem and phloem in plants. Some multi-cellular algae and bryophytes also have specialized conducting cells. 5. Reproduction. Organisms in water can release their gametes into the water, where the gametes will swim by flagella until they encounter each other and fertilization happens. On land, such a scenario is not possible. Land animals have had to develop specialized reproductive systems involving fertilization when they return to water (amphibians), or internal fertilization and an amniotic egg (reptiles, birds, and mammals). Insects developed similar mechanisms. Plants have also had to deal with this, either by living in moist environments like the ferns and bryophytes do, or by developing specialized delivery systems like pollen tubes to get the sperm cells to the egg. Plants divide into two large groups: vascular plants that contain lignified conducting cells, and the nonvascular plants, which do not. Some Silurian plant fossils might be algae or nonvascular plants. Vascular plants developed during the Silurian period, 400 million years ago. The earliest vascular plants had no roots, leaves, fruits, or flowers. From: http://www.emc.maricopa.edu/faculty/farabee/biobk/BioBookPaleo3.html Before chlorophyll came along, organisms presumably had to depend for their food on the amino acids which were dissolved in the water they were living in. This was a very marginal, iffy kind of existence, and so it is unlikely that there was a very high density of living things present on the planet until chlorophyll made its appearance by introducing the food-synthesizing blue-green organisms onto the scene. These new blue-green organisms, because they could make their own food wherever there was sunlight, were much more independent than their forerunners, and so suddenly life- forms were free to go forth and multiply far more quickly than they had been in the past (for some images of blue-green organisms, please refer to the pictures towards the beginning of this post). Remember, without food, it is very difficult to reproduce, and to live, so life had to solve the problem of provisioning itself efficiently and reliably, before it could move on to conquer the planet to the extent to which it has done so today. Source: http://www.emc.maricopa.edu/faculty/farabee/biobk/BioBookPaleo3.html Would it still be called a family tree even though it’s plants we’re talking about?! A very general sketch of the ancestry of plants. If you’re not sure what the difference between vascular and non-vascular plants is, please look at the boxed text above. This ability to make your own food however was a double-edged sword, and was not wholly and unequivocally a good thing. It certainly allowed one to live in new zones, but it also stimulated the evolution of predators: Since there were now a bunch of these concentrated food-packages (i.e. organisms which made their own food) floating around, other organisms evolved the ability to get their food by simply eating these food producing life-forms. These new organism-eating organisms were the first predators –which eventually became the first animals (all animals sustain themselves by eating other organisms – be they plants or other animals). You may not know this, but ladybirds are actually aggressive predators. Source: http://www.eoearth.org/article/Predation Source: http://australianmuseum.net.au/Uploads/Images/9148/herbivory_021_big.jpg A stick insect eating a leaf. Source: http://www.zoology.ubc.ca/courses/bio332/Labs/CiliateProject/DidEatpara.jpg An organism in the process of consuming its fellow organism – it’s a dog-eat-dog (or would that be blob-eat-blob?) world under the microscope too! Anyway, from the beginnings of life on earth, until about 570,000,000 years ago, things were pretty slow: There were a bunch of tiny blobs floating around synthesizing their own food out of air, water, minerals and sunlight; there were some other blobs which developed the predatory habit of swallowing their fellow blobs; and a bit later, there were also some larger blobs composed of agglomerations of little blobs in various stages of mutual integration, and that was about it. Then, for some as yet unknown reason, in a matter of a few million years, a whole bunch of different, and rather strange looking life-forms evolved. This explosion of life is known as the Cambrian revolution. It was at this time that vaguely familiar ancient animals like trilobites, and the ancestors of prawns, crabs and lobsters (as well as those of humans and most other things which are around today) first emerged; it was also the time when many forms which were utterly bizarre, utterly unfamiliar, and which have left no descendants today – organisms with five eyes, and so on – also first made their appearance on the scene. As an aside, I want to note that some religious fundamentalists have tried to argue that the Cambrian explosion is evidence of the hand of god at work, but I find this a bit of an odd argument for a fundamentalist to make: did a supposedly omnipotent, omniscient, omnipresent and benevolent god make a mistake, not think things through properly, or perhaps change its mind, in deciding to suddenly cause life to radiate out in new and really rather radical directions? This does not seem to fit well with the 4 classical properties normally ascribed to god. Doubtless though, the response of the fundamentalists (might I say irrationalists) would be that the ways of god are inscrutable to the intelligence of man: a convenient verbal-conceptual trick which allows one to dodge difficult questions. Source: http://www.emc.maricopa.edu/faculty/farabee/biobk/BioBookPaleo3.html The thickness of the lines indicates relative diversities of the various forms. Notice that the chordates (which includes us, or rather our distant direct ancestors) did not clamber onto the stage until relatively late in the story, whilst molluscs and arthropods (this includes clams, squid, snails and slugs) are far more blue-blooded than any peer of the realm, since they really can trace their ancestry back to the dim past. Source: http://webecoist.com/2009/09/22/gone-wild-7-extinct-wonders-of-the-animalkingdom/ Opabinia – a five-eyed predator with an unusual feeding appendage from 510,000,000 years ago. There are no descendants of this guy around today: how do we know? Do you see any 5-eyed animals with one feeding-appendage around? Source: http://www.emc.maricopa.edu/faculty/farabee/biobk/BioBookPaleo3.html Meet anomalocaris (whose name means anomalous shrimp), the biggest (ca. 1 metre long), baddest predator of the Cambrian period – it was in fact the first great predator of the Cambrian period, and its prey included trilobites. It had an unusual, disk-like mouth which was composed of 32 overlapping plates, four large and 28 small, resembling pineapple rings, with their centres replaced by a series of serrated prongs. The mouth could constrict to crush prey, but never completely close, and the tooth-like prongs continued down the walls of the gullet. Truly an animal fit for a starring role in an alien horror-movie. Source: http://paleobiology.si.edu/burgess/imgBurgess/anamalocaris1.gif A photograph of the fossilised mouth of anomalocaris. The Cambrian was also the time – with the first appearance of hard, more easily fossilised parts – that the fossil record picks up the pace. Source: http://www.didmancreategod.com/Chapter_excerpts/images/C3%20Marrella.jpg Above is marrella, a delicate looking arthropod often referred to as a ‘lace-crab’. Source: http://paleobiology.si.edu/burgess/imgBurgess/hallucigenia.gif The perhaps aptly named hallucigenia. When it was first discovered, some scientists thought it walked on the long spines, rather like a stilt-walker does today. It has since been discovered that the spines were on its dorsal (back) side, not on its ventral (stomach) side. Source: http://digsfossils.com/fossils/pics/trilobites/morocco-trilobite001a.jpeg There are about 5,000 genera, encompassing over 17,000 species of trilobites (the name means 3 lobes). They first appeared 526,000,000 years ago, and finally died out about 250,000,000 years ago (that’s a longer span of time than that which separates us from the earliest proto-dinosaurs by the way). They had a variety of life-styles: some swam, some crawled; some were predators, some were scavengers; some were spiny, some were smooth; some were tiny (ca. 1 mm), some were quite large (ca. 720 mm); but none of them played jazz, and none of them sang the blue; and though there is some debate about it, the general consensus is that there are no direct trilobites descendants around today. Why they disappeared is a bit of a mystery, but their numbers seem to have diminished drastically once predatory fish made their big entrance, so there may b a connection here. Source: http://en.wikipedia.org/wiki/Trilobite Some more examples showing the enormous diversity of trilobites. Source: http://www.emc.maricopa.edu/faculty/farabee/biobk/BioBookPaleo3.html Above is pikaia, it is the oldest known chordate – in so being, it turns out that it is likely also your direct ancestor, as well as being the ancestor of every other vertebrate (lions and tigers and bears – oh my!) and a bunch of other animals too. Very soon after this came the first fishes, many of which, although very strange looking, were still recognizable as fish, albeit that some had a rather terrifying aspect. Source: http://www.fish.state.pa.us/pafish/prehstrm.jpg This is one of a whole slew of armoured fish which for a while swam in the oceans of the planet, before going extinct. The largest of these armoured fish was Dunkleosteus, at 10 m long. It all brings to mind Omar Khayyam: Think, in this batter'd Caravanserai Whose Portals are alternate Night and Day, How Sultan after Sultan with his Pomp and Glory, Abode his destined Hour, then was sent his way. – The Rubaiyat of Omar Khayyam Source: http://www.topnews.in/files/ancient-fish-fossils.jpg It was also the beginning of the first arms race on the planet of which we have a record: Some fish, in order to protect themselves from other, predatory, fish began developing defensive heavy armour – bony plates which were presumably hard to bite through. Arms races however can have no positive final outcome, and so this gambit was finally abandoned by the forces of evolution, – though apparently not by human heads of state / government, who really could learn some lessons from palaeontology! Source: http://www.emc.maricopa.edu/faculty/farabee/biobk/1ostraco.jpg A reconstruction of an ostracoderm. Around this time (450,000,000 years ago), the first ‘proper’ plants (as opposed to algae and such) also began growing on the land. Again, these first land plants were, by our standards, rather odd looking. Source: http://www.emc.maricopa.edu/faculty/farabee/biobk/cook.jp This is a fossil of Cooksonia, member of an extinct group of early land plants. Source: http://website.nbm-mnb.ca/Doliodus/Devonian%20mural.jpg A Devonian scene as interpreted by an artist. These plant, in turn, were followed by some early arthropods (centipede-like animals), which could now live on land, as there was something there for them to eat. This model is of a rather large arthropod called Arthropleura shows you just how big they were. Source: http://www.icteach.nl/Pagina's/Uitgestorven/Inhoud/Phanerozoïcum/Paleozoïcum/Carboon/B oven%20Carboon/Fauna/Geleedpotigen/arthropleura07.jpg Source: http://ncse.com/files/images/Tetrapod_transition.preview.jpg Then, about 400,000,000 years ago, the first amphibians evolved, probably out of some sort of fish (which is why you, your dog / cat and your chicken dinner are all really just somewhat evolved bony fish [osteichthyes] at heart). These animals, of which Ichthyostega is perhaps the most well known by the public at large, were still highly dependent on water, and like modern-day amphibians (e.g. frogs), they had to lay their soft, shell-less eggs in the water, (or in a watery medium like foam), but they could breathe air, and they could walk around on land. They were still on a fairly short, water-linked tether though, because their skins could not tolerate extremely dry environments, and had to be kept moist. Source: http://webspace.webring.com/people/cu/um_1925/eustenovsichthy.jpg Sorry it’s not in English, but you can work out what all the bits are: yet another way in which science is sometimes a universal language (just kidding!). None-the-less, this ability to live, or at least spend lots of time on land was a great boon, as it now permitted vertebrates to colonise a part of the earth where they were not present before. These early amphibians eventually evolved into all sorts of terrestrial (and some marine) vertebrates. They also inherited bilateral symmetry, and a basic body-plan from their fish ancestors (which is why on top of being bony fish at heart, you, your dog / cat, and your chicken-dinner all share so many skeletal features in common – 4 limbs, each of which has a1-bone-then-2-bone-then-a-bunch-of-bones-layout for example). As far as the story of fossils goes, the newer they are, the more common, or rather the less rare they become, and this is well reflected in the fossil record, in the frequency with which fossils from different periods are found. Thus, non-rational (irrational?) religious objections to evolution notwithstanding, the facts are that the record of evolution from fish to terrestrial animal is already quite well reflected in the fossil record (as you can clearly see from the graphic above entitled: ‘Evidence of Macroevolution’), and with time, and the future disinterring of more fossil-discoveries, this particular story is only going the get more and more detailed. Source: http://download.naturkundemuseumberlin.de/presse/Darwin/Stuttgart/Fossil_Sclerocephalus.jpg Now, while this was going on, the plants were also evolving. They were growing bigger, and more complex. By about 350, 000, 000 years ago, the land was covered with all sorts of plants, mostly ferns and mosses, although there were also a number of different kinds of trees to be found. Much of the earth’s surface at this time was swampy, so many of the plant-forms were adapted to living in swampy conditions. These plants died, were buried, and eventually became coal. So coal too is in fact a sort of trace of the history of life on the planet, and can give us more indicators of how life on earth evolved. Source: http://www.emc.maricopa.edu/faculty/farabee/biobk/pro_plfr.gif The width of the blue or green sections is proportional to the number of species. Source: http://www.sparththenandnow.org.uk/prehistoric/sparthfossils/528/ Fossilised tree-root Also around this time, the first reptiles evolved, probably from one of the amphibians. These new animals had a great advantage over their amphibian cousins: Because they had developed hard-shelled eggs, the reptiles didn’t need to lay their eggs in water. Therefore, the reptiles were able to live very far away from water (they needed it to drink of course, but they didn’t need large pools of it in which to lay their eggs). Gradually, these reptiles grew in size and in complexity, and some of them evolved into other things (which is why you, your dog / cat, your chicken-dinner, and any lizard you come across all share the same basic skeletal plan which I mentioned above, and which you can clearly see in the fossil below). Source: http://www.tucsonshow.com/reports/tucson2002/p1.shtml Discosauriscus pulcherrimus After 100,000,000 years or so, evolution again produced some new types of animals: mammals, and dinosaurs. These early mammals evolved out of some rather odd looking mammal-like reptiles like good old moschops and thrinaxodon below (again note the skeletal similarities between you, your dog / cat,…); the dinosaurs meanwhile evolved out of a different group of reptiles. Source: http://www.volkoff.net/Images/links/Moschops.gif Source: http://upload.wikimedia.org/wikipedia/commons/d/d1/Thrinaxodon_Lionhinus.jpg Thrinaxodon The oldest dinosaur fossils we have found are those of Eoraptor and Herrerasaurus, both of which are about 230,000,000 years old. Source: http://maniraptora.com/dinosaurs_world/macn/eoraptor/eoraptor_final.gif Eoraptor – dawn thief – one of the earliest dinosaurs of which we have fossil evidence. Presumable, it was descended from some earlier form, but we have yet to find fossil evidence of this ancestor. One thing to note, dinosaurs were initially bipedal, so somewhere along the way they must have evolved from the standard reptilian quadrupedal form, but that story is as yet incomplete. Source: http://www.zum.de/Faecher/Materialien/beck/bilder/herrera.gif Herrerasaurus. Out of these animals (and perhaps a few others early dinosaurs, which may not have left fossil remains) evolved all the other dinosaurs, many of which are familiar to you already. One important thing to note here is that about 180,000,000 years separates these first dinosaurs, from the last dinosaurs to have lived. Thus, a lot of evolving was going on in between. The at-one-time-usual pictures, showing diplodocus fighting with t. rex whilst a pterosaur flies overhead is quite simply false. These animals lived at vastly different points in time, and each was already extinct before the next one had evolved. There were also flying reptiles and a bit later birds, but fossils of these animals are rather rare, as their bones were rather fine and delicate. Source: http://media-2.web.britannica.com/eb-media/40/3640-004-D1583DEA.jpg Pterosaur Source: http://www.fossilmuseum.net/fossilpictures-wpd/Archaeopteryx/Archaeopteryx.jpg Archaeopteryx So as you can see, from this little survey post, the fossil record, though not complete (hence the title of my post), is very good, and the more recent the date of fossilisation, the better the fossils, and hence the better and more complete the fossil record becomes. By the time one comes to the end of the Tertiary period (when the dinosaurs were around), and to the Quaternary period, which was after the dinosaurs, the remains more often become not just fossils, but also sub-fossils and such. These are remains which either because the soil chemistry was not right, or simply because they are not old enough, have not fossilised completely, as I noted in another post within this ‘History of life on the planet series’. Furthermore, the remains change from fossils and sub-fossils to actual remains, organic matter – bones, skin, teeth, hair etc, but that is another story.