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TWO

Building Bodies

The Great Barrier Reef swarms with life. The tides surging through the coral heads charge the water with oxygen and the tropical sun warms it and fills it with light. All the main kinds of sea animals seem to flourish here. Phosphorescent purple eyes peer out from beneath shells; black sea urchins swivel their spines as they slowly perambulate on needle tip; starfish of an intense blue spangle the sand; and patterned rosettes unfurl from holes in the smooth surface of coral. Dive down through the pellucid water and turn a boulder. A flat ribbon, striped yellow and scarlet, dances gracefully away and an emerald green brittle star careers over the sand to find a new hiding place.

The variety at first seems bewildering, but leaving aside primitive creatures like jellyfish and corals which we have already described, and the much more advanced backboned fish, nearly all can be allocated to one of three main types: shelled animals, like clams, cowries and sea snails; radially symmetrical creatures, like starfish and sea urchins; and elongated animals with segmented bodies varying from wriggling bristle worms to shrimps and lobsters.

The principles on which these three kinds of bodies are built are so fundamentally different that it is difficult to believe that they can be related to one another except right at the very roots of the evolutionary tree. The fossil record bears this out. All three groups, being sea-dwellers, have left behind abundant remains, and the details of their separate dynastic fortunes can be traced through the rocks for hundreds of millions of years. The walls of the Grand Canyon show that animals without backbones, invertebrates, came into existence long before the vertebrates such as fish. But just below the layer of gently folded limestones that contain the earliest of the invertebrate fossils, the strata change radically. Here the rocks are highly contorted. They had once formed mountains. These were eroded and eventually covered with the sea that deposited the limestone now lying above them. The episode occupied many millions of years and during all that time there were no deposits. As a consequence, this junction in the rocks represents a huge gap in the record. To trace the invertebrate lines back to their origins, we must find another site where rocks were not only deposited continuously throughout this critical period, but have survived in a relatively undistorted condition.

Such places are few, but one lies in the Atlas Mountains of Morocco. The bare hills behind Agadir in the west are built of blue limestones so hard that they ring under the fossil hunter’s hammer. The beds of rock are slightly tilted but otherwise undistorted by earth movements. On the crest of the passes, the rocks yield fossils. They are not very many, but if you look hard enough you can collect quite a range of species. All fossils found anywhere in the world in rocks of this age can be placed in one or other of those three main groups we identified on the reef. There are tiny shells, the size of your little fingernail, called brachiopods; radially symmetrical organisms looking like stalked flowers called crinoids; and trilobites, segmented creatures that resemble woodlice.

The limestones at the top of the Moroccan succession are about 560 million years old. Beneath them lie more layers extending downwards for thousands of metres, seemingly unchanged in character. In them, surely, must be evidence about the origins of those three great invertebrate groups.

But it is not so. As you clamber down the mountainside over the strata, the fossils suddenly disappear. The limestone seems to be exactly the same as that at the head of the pass, so the seas in which it was laid down must surely have been very similar to those that produced fossiliferous rocks. There are no signs of a revolutionary change in physical conditions. It is simply that at one time the ooze covering the seafloor contained shells of animals – and before that it did not.

This abrupt beginning to the fossil record is not just a Moroccan phenomenon, though you can see it here more vividly than in most places. It occurs in almost all the rocks of this age throughout the world. The microfossils from the cherts of Lake Superior and South Africa show that life had started long, long before. In the theoretical year of life, shelled animals do not appear until early November. So the bulk of life’s history is undocumented in the rocks. Only at this late date, about 600 million years ago, did several separate groups of organisms begin to leave records of any abundance by secreting shells. Why this sudden change should have come about, we do not know. Perhaps before this time the seas were not at the right temperature or did not have the chemical composition to favour the deposition of the calcium carbonate from which most marine shells and skeletons are constructed. Whatever the reason, we have to look elsewhere for evidence of the origins of the invertebrates.


A living crinoid: the great west indian sea lily (Cenocrinus asterius), 180–250 metres depth, Caribbean.


Flatworm (Maiazoon orsaki) Raja Ampat, Irian Jaya, Indonesia, Pacific Ocean.

We can find some living clues back on the reef. Fluttering over the coral heads, hiding in the crevices or clinging to the underside of rocks, are flat leaf-shaped worms. Like jellyfish, they have only one opening to their gut through which they both take in food and eject waste. They have no gills and breathe directly through their skin. Their underside is covered with cilia which by beating enable them to glide slowly over surfaces. Their front end has a mouth below and a few light-sensitive spots above so that the animal can be said to have the beginnings of a head. These flatworms are the simplest creatures to show signs of such a thing.

Eye-spots, to be of any use, must be linked to muscles so that the animal can react to what it senses. In flatworms all that exists is a simple network of nerve fibres. There are a few thickenings in some of them, but these can hardly be described as brains. Yet the flatworms can learn the kind of things that would help even this simplest of animals to survive, such as avoiding a particularly dangerous place or remembering where food can be found.

Today we know of some 3,000 species of flatworm in the world. Most are tiny and water-living. You can find freshwater ones in most streams simply by dropping a piece of raw meat or liver into the water. If the underwater vegetation is thick, flatworms are likely to glide out in some numbers and settle on the bait. In humid tropical forests, the ground is usually moist enough for some species to live on land, and many are likely to appear, undulating on the mucus that they secrete from their undersides. One of these species grows to a length of about 60 centimetres. Other flatworms have taken to the parasitic life and live unseen within the bodies of other animals – including us.

Liver flukes still retain the typical flatworm form. Tapeworms are also members of the group, though they look very different, for after burying their heads in the walls of their host’s gut, they bud off egg-bearing sections from their tail end. These segments remain attached while they mature, eventually forming a chain that may be as much as 10 metres long. The whole creature, as a result, looks as though it is divided into segments, but in fact these separate living packets of eggs are quite different from the permanent internal compartments of a truly segmented creature like an earthworm.

Flatworms are very simple creatures. Members of one free-swimming group lack a gut altogether and look very like the tiny free-swimming coral organisms before they settle down to a sedentary life. So there is little difficulty in believing those researchers who conclude from a study of the detailed structure of both adult and larva that the flatworms are descended from simpler organisms like corals and jellyfish.

During the period when these first marine invertebrates were evolving, between 600 and 1,000 million years ago, erosion of the continents was producing great expanses of mud and sand on the seabed around the continental margins. This environment must have contained abundant food in the form of organic detritus falling from the waters above as the single-celled organisms that floated in the surface waters died and drifted downwards. It also offered concealment and protection for any creature that lived within it. The flatworm shape, however, is not suited to burrowing. A tubular form is much more effective, and eventually worms with such a shape appeared. Some became active burrowers, tunnelling through the mud in search of food particles. Others lived half buried with their front end above the sediment. Cilia around their mouths created a current of water and from it they filtered their food.

Some of these creatures lived in a protective tube. In time, the shape of the top of this was modified into a collar with slits in it. This improved the flow of water over the tentacles. Further modification and mineralisation eventually produced a two-part protective shell around the front end. These were the first brachiopods, including Lingulella, an example of a species that has existed virtually unchanged for hundreds of millions of years.

The front end of a brachiopod is really quite complicated. Within the shell, it has a mouth surrounded by a group of tentacles. They are covered with beating cilia which create a current in the water. Any food particles in it are caught by the tentacles and then passed by them down to the mouth. While doing this, the tentacles perform another and important function, for the water brings with it dissolved oxygen which the animal needs in order to respire. The tentacles absorb it and so, in effect, they become gills. The shell enclosing the tentacles not only gives protection to these soft delicate structures, but concentrates the water into a steady stream so that it flows more effectively over them.

The brachiopods elaborated this design considerably over the next million years or so. One group developed a hole at the hinge end of one of the valves through which the worm-like stalk emerged to fasten the animal into the mud. This gave the shell the look of an upside-down Aladdin oil lamp, with the stalk as the wick, and so the group as a whole gained the name of lamp shell. The tentacles within the shell eventually became so enlarged that they had to be supported by delicate spirals of limestone.

There are other shelled worms to be found alongside the brachiopods in these ancient rocks. In one kind the elaborated worm did not attach itself to the seafloor but continued to crawl about and secreted a small conical tent of shell under which it could huddle when in danger. This was the ancestor of the most successful group of all these shelled worms, the molluscs, and it too has a living representative, a tiny organism called Neopilina, which was dredged up in 1952 from the depths of the Pacific. Today there are about 80,000 different species of molluscs with about as many again known from their fossils. You can find some of them in your garden; they are the snails and the slugs.


Brachiopods (Glottidia albida).

The lower part of the molluscan body is called the foot. Its owner moves itself about by protruding the foot from the shell and rippling its undersurface. Many species carry a small disc of shell on the side of it which, when the foot is retracted into the shell, forms a close-fitting lid to the entrance. The upper surface of the body is formed by a thin sheet that cloaks the internal organs and is appropriately called the mantle. In a cavity between the mantle and the central part of the body, most species have gills which are continually bathed by a current of oxygen-bearing water, sucked in at one end of the cavity and expelled at the other.

The shell is secreted by the upper surface of the mantle. One whole group of molluscs has single shells. The limpet, like Neopilina, produces shell at an equal rate right round the circumference of the mantle and so builds a simple pyramid. In other species, the front of the mantle secretes faster than the rear and creates a shell in a flat spiral, like a watch spring. In yet others, maximum production comes from one side so that the shell develops a twist and becomes a turret. The cowrie concentrates its secretion along the sides of the mantle, forming a shell like a loosely clenched fist. From the slit along the bottom, it protrudes not only its foot but two sections of its mantle which in life may extend over each flank of the shell and meet at the top. These lay down the marvellously patterned and polished surface characteristic of cowries.


Blue limpet (Patella coerulea), showing underside.

The single-shelled molluscs feed not with tentacles within the shell like the brachiopods but with a radula, a ribbon-shaped tongue, covered with rasping teeth. Some use it to scrape algae from the rocks. Whelks have developed a radula on a stalk which they can extend beyond the shell and use to bore into the shells of other molluscs. Through the holes they have drilled, they poke the tip of the radula and suck out the flesh of their victim. Cone shells also have a stalked radula but have modified it into a kind of gun. They slyly extend it towards their prey – a worm or even a fish – and then discharge a tiny glassy harpoon from the end. While the tethered victim struggles, they inject a venom so virulent that it kills a fish instantly and can even be lethal to human beings. They then haul the prey back to the shell and slowly engulf it.

A heavy shell must be something of a handicap when actively hunting, and some carnivorous molluscs have taken to a faster if riskier life by doing without it altogether and reverting to the lifestyle of their flatworm-like ancestors. These are the sea slugs (nudibranchs) and they are among the most beautiful and highly coloured of all invertebrates in the sea. Their long soft bodies are covered on the upper side with waving tentacles of the most delicate colours, banded, striped and patterned in many shades. Though they lack a shell, they are not entirely defenceless, for some have acquired secondhand weapons. These species float near the surface of the water on their feathery extended tentacles and hunt jellyfish. As the sea slug slowly eats its way into its drifting helpless prey, the stinging cells of the victim are taken into its gut, complete and unsprung. Eventually these migrate within the sea slug’s tissues and are concentrated in the tentacles on its back. There they give just the same protection to their new owners as they did to the jellyfish that developed them.

Other molluscs, such as mussels and clams, have shells divided into two valves lke those of a brachiopod and thus are known as bivalves. These creatures are much less mobile. The foot is reduced to a protrusion that they use to pull themselves down into the sand. For the most part, they are filter feeders, lying with valves agape, sucking water in through one end of the mantle cavity and squirting it out through a tubular siphon at the other. Since they do not need to move, great size is no disadvantage. Giant clams on the reef may grow to be a metre long. They lie embedded in the coral, their mantles fully exposed, a zigzag of brilliant green flesh spotted with black, which pulsates gently as water is pumped through it. They can certainly be quite big enough for a diver to put his foot into, but he would have to be very incautious indeed to get trapped. Powerful though the clam’s muscles are, it cannot slam its valves shut. It can only heave them slowly together, and that gives plenty of notice of its intentions. What is more, even when the valves of a really large specimen are fully closed, they only meet at the spikes on the edge. The gaps between them are so big that if you plunge your arm through into the mantle, the clam is quite unable to grip it – though the experiment is a little less unnerving if it is tried first with a thick stick.

Some filter feeders like the scallops do manage to travel – by convulsively clapping their valves together and so making curving leaps through the water. By and large, however, adult bivalves live rather static lives and the spreading of the species into distant parts of the seabed is carried out by the young. The molluscan egg develops into a larva, a minuscule animated globule striped with a band of cilia, which is swept far and wide by ocean currents. Then, after several weeks, it changes its shape, grows a shell and settles down. The drifting phase of its life puts it at the mercy of all kinds of hungry animals, from other stationary filter feeders to fish, so in order that its species can survive, a mollusc must produce great numbers of eggs. And indeed it does. One individual may discharge as many as 400 million.

One branch of the molluscs, very early in the group’s history, found a way of becoming highly mobile and yet retaining the protection of a large and heavy shell – they developed gas-filled flotation tanks. The first such creature appeared about 500 million years ago. Its flat-coiled shell was not completely filled with flesh as is that of a snail, but had its hind end walled off to form a gas chamber. As the animal grew, new chambers were added to provide sufficient buoyancy for the increasing weight. This creature was the nautilus, and we can get an accurate idea of how it and its family lived because a few nautilus species, just like Lingulella and Neopilina, have survived to the present day.


A nudibranch (Eubranchus tricolor) on the seabed of a Scottish loch.

One of these species, the pearly nautilus, grows to about 20 centimetres across. A tube runs from the back of the body chamber into the flotation tanks at the rear so that the animal can flood them and adjust its buoyancy to float at whatever level it wishes. The nautilus feeds not only on carrion but on living creatures such as crabs. It moves by jet propulsion, squirting water through a siphon in a variation of the current-creating technique developed by its filter-feeding relatives. It searches for its prey with the help of small stalked eyes and tentacles that are sensitive to taste. Its molluscan foot has become divided into some ninety long grasping tentacles which it uses to grapple with its prey. In the centre of them it has a horny beak, shaped like that of a parrot and capable of delivering a lethal, shell-cracking bite.

About 400 million years ago, after some 100 million years of evolution, the nautiluses gave rise to a variant group with many more flotation chambers to each shell, the ammonites. These became much more successful than their nautilus relatives, and today their fossilised shells can be found lying so thickly that they form solid bands in the rocks. Those of some species grew as big as lorry wheels. When you find one of these giants embedded in the honey-coloured limestones of central England or the hard blue rocks of Dorset, you might think that such immense creatures could do little but lumber massively across the seabed. But where erosion has removed the outer shell, the elegant curving walls of the flotation chambers that are revealed remind you that these creatures may well have been virtually weightless in water and able, like the nautilus, to jet-propel themselves at some speed through the water.

About 100 million years ago, the ammonite dynasty began to dwindle. Perhaps there were ecological changes that affected their egg-laying habits. Maybe new predators had appeared. At any rate, many species died out. Other lines gave rise to forms in which the shells were loosely coiled or almost straight. One group took the same path as the sea slugs did in more recent times and lost their shells altogether. Eventually all the shelled forms except the pearly nautilus disappeared. But some shell-less ones survived and became the most sophisticated and intelligent of all the molluscs, the squids and cuttlefish and the octopus. These are the cephalopods.

The relics of the cuttlefish’s ancestral shell can be found deep within it. This is the flat leaf of powdery chalk, the cuttlebone, that is often washed up on the seashore. The octopus has no trace of a shell even within the flesh of its body, but one species, the argonaut, secretes from one of its arms a marvellous paper-thin version shaped very like a nautilus shell but without chambers, which it uses not as a home for itself but as a delicate floating chalice in which to lay its eggs.

The squid and cuttlefish have many fewer tentacles than the nautilus – only ten – and the octopus, as its name makes obvious, has only eight. Of the three creatures, the squids are much the more mobile and have lateral fins running along their flanks which undulate and so propel the animal through the water. All cephalopods can, like the nautilus, use jet propulsion on occasion.


Several nautilus (Nautilus pompilius) on a coral reef at night, Pacific.

Cephalopod eyes are very elaborate. In some ways they are even better than our own, for a squid can distinguish polarised light, which we cannot do, and their retinas have a finer structure, which means, almost certainly, that they can distinguish finer detail than we can. To deal with the signals produced by these sense organs they have considerable brains and very quick reactions.

Some squids grow to an immense size. The aptly named colossal squid lives in the seas around Antarctica. It can reach nearly 100 kilos in weight and measure six metres from the end of its body to the tip of its outstretched tentacles. Its rival for the claim to be the largest species of all is the giant squid. The biggest so far discovered have in fact been slightly smaller and substantially lighter. Although there are records of even larger specimens of this species, it seems that these were not accurate. Nevertheless, we are unlikely to have discovered the biggest individuals of either species, so the record may yet be broken. The eyes of these huge cephalopods are even larger than might be expected. The biggest recorded were 27 centimetres across and are the largest known eyes of any kind of animal, five times bigger, for example, than those of the blue whale. Why the squid should have such gigantic eyes is a mystery.

It could be, however, that they need extremely sensitive eyes to detect the presence of their great enemy – the sperm whale. Squid beaks are often found in the stomachs of sperm whales, and their heads often carry circular scars with diameters that match a squid’s suckers. So there seems little doubt that squids and whales regularly fight in the dark depths of the ocean. Maybe the squids’ huge eyes help them to detect the presence of the only animal big enough to hunt them.

The intelligence of all the cephalopods – octopus, squid and cuttlefish – is well known. Octopus have been observed disguising themselves from an approaching enemy by covering themselves with shells or picking up two halves of a coconut and hiding within. Many species in all three groups have an extraordinary ability to change their colour and shape. They can camouflage themselves by matching almost any environment and also signal to one another with patterns and shapes that sweep across their bodies. A female squid has even been filmed signalling to a male lying alongside her that she is not ready to mate, while at the same time displaying a pattern on the other side of her body to summon another male. Octopus and squid, two of the most advanced animals in the ocean which least resemble human beings, are among the few, it seems, that can rival mammals in their intellectual abilities.

But what of the second great category of animals without backbones, the one represented in ancient rocks by the flower-like crinoids? As these are traced upwards through the rocks, they become more elaborate and their fundamental structure becomes clearer. Each has a central body, the calyx, rising from a stem like the seedhead of a poppy. From this sprout five arms which, in some species, branch repeatedly. The surface of the calyx is made up of closely fitting plates of calcium carbonate, as are the stems and branches. Lying in the rocks, the stems look like broken necklaces, their individual beads sometimes scattered, sometimes still in loose snaking columns, as though their thread had only just snapped. Occasionally gigantic specimens are found with stems 20 metres long. These creatures, like the ammonites, have had their day, but a few species, sometimes called sea lilies, still survive in the ocean depths.


Bigfin squid (Sepioteuthis lessoniana) hovering in open water above a coral reef at night. Dampier Strait, Raja Ampat, West Papua, Indonesia. Tropical West Pacific Ocean.


Crinoid (feather star, centre) on a gorgonian (sea fan, red) with a Dendronephthya soft coral in the background, Andaman Sea, Thailand.

Sea lilies show that the calcium carbonate plates, in life, are embedded just under the skin. This gives their surface a curious prickly feel. In other families, related to the crinoids, the skin has spines and needles attached to it so the creatures are known as echinoderms, ‘spiny-skins’. The basic module on which the echinoderm body is built has a fivefold symmetry. The plates on the calyx are pentagons. Five arms extend from it, and all the internal organs occur in groups of five. Their bodies work by a unique exploitation of hydrostatic principles. Tube feet, each a thin tube ending in a sucker and kept firm by the pressure of water within, wave and curl in rows along the arms. The water for this system circulates quite separately from that in the body cavity. It is drawn through a pore into a channel surrounding the mouth and circulated throughout the body and into the myriads of tube feet. When a drifting particle of food touches an arm, tube feet fasten on to it and pass it on from one to another until it reaches the gutter that runs down the upper surface of the arm to the mouth at the centre.


Tube feet of a red cushion sea star (Oreaster reticulatus), Singer Island, Florida.

Though stalked sea lilies were the most abundant crinoid in fossil times, the commonest forms today are the feather stars. Instead of stalks, they have a cluster of curling roots with which they attach themselves to coral or rocks. In places on the Great Barrier Reef, they swarm in huge numbers, covering the floor of the tidal pools with a tufted coarse carpet of brown. When disturbed, however, they can suddenly swim away, writhing their five limbs like Catherine wheels.

The fivefold symmetry and the hydrostatically operated tube feet are such distinctive characteristics that they make other echinoderms very easy to recognise. The starfish and their more sprightly cousins, the brittle stars, both possess them. These creatures appear to be crinoids that have neither stalk nor rootlets and are lying in an inverted position with their mouths on the ground and their five arms outstretched. Sea urchins too are obviously related. They seem to have curled their arms up from the mouth as five ribs and then connected them by more plates to form a sphere.

The sausage-like sea cucumbers that sprawl on sandy patches in the reef are also echinoderms, although in most species their shelly internal skeletons are reduced to tiny structures beneath the skin. Most lie neither face up nor face down, but on their sides. At one end there is an opening called the anus, though the term is not completely appropriate for the animal uses it not only for excretion but also for breathing, sucking water gently in and out over tubules just inside the body. The mouth, placed at the other end, is surrounded by tube feet that have become enlarged into short tentacles. These fumble about in the sand or mud. Particles adhere to them and the sea cucumber slowly curls them back into its mouth and sucks them clean with its fleshy lips.

One highly specialised deep-sea sea cucumber, called a sea-pig, lives in the mud of the deep seabed at depths of up to 5,000 metres. They are rotund little creatures about 15 centimetres long and have large tube-like structures on their underside with which they rootle about in the mud. They have been filmed in the deep sea, assembled in herds, perhaps for reproduction or attracted by the smell of a new source of food drifting down from the surface.

If you pick up a sea cucumber, do so with care, for they have an extravagant way of defending themselves. They simply extrude their internal organs. A slow but unstoppable flood of sticky tubules pours out of the anus, fastening your fingers together in an adhesive tangle of threads. When an inquisitive fish or crab provokes them to such action, it finds itself struggling in a mesh of filaments while the sea cucumber slowly inches itself away on the tube feet that protrude from its underside. Over the next few weeks it will slowly grow itself a new set of entrails.

The echinoderms may seem, from a human point of view, to be a blind alley of no particular importance. Were we to imagine that life was purposive, that everything was part of a planned progression due to culminate in the appearance of the human species or some other creature that might rival us in dominating the world, then the echinoderms could be dismissed as of no consequence. But such trends are clearer in the minds of people than they are in the rocks. The echinoderms appeared early in the history of life. Their hydrostatic mechanisms proved a serviceable and effective basis for building a variety of bodies, but were not susceptible in the end to spectacular development. In the areas that suit them, they are still highly successful. A starfish on the reef can crawl across a clam, fasten its tube feet on either side of its gape and slowly wrench the valves apart to feed on the flesh within. The crown-of-thorns starfish occasionally proliferates to plague proportions and devastates great areas of coral. Crinoids are brought up in trawls from the deep sea several thousand at a time. If it is improbable that any further major developments will come from this stock, it is also unlikely, on the evidence of the last 600 million years, that the group will disappear as long as life remains possible at all in the seas of the world.


Panamic cushion sea stars (Pentaceraster cumingi) group on seafloor, Galapagos Islands.

The third category of creatures on the reef contains those with segmented bodies. In this instance, we do have fossil evidence of even earlier forms than the trilobites found in the Moroccan hills. The Ediacaran deposits in Australia which contain the remains of jellyfish and sea pens also preserve impressions of segmented worms. One species, a 5-centimentre-long animal named Spriggina after Reg Spriggs who first discovered the Ediacara fossils, has a crescent-shaped head and up to forty segments, fringed on either side by leg-like projections. What exactly it was, nobody can agree. No legs have been identified, but this may be a limitation in the process of fossilisation. Some scientists think it may represent a completely extinct group. One widely accepted possibility is that it was a kind of annelid worm related to the bristle worms that are so common on a reef and the earthworms that you can find in your garden.

Annelids have grooves encircling their body that correspond to the internal walls that divide its interior into separate compartments. Each of these is equipped with its own set of organs. On the exterior and on either side, there are leg-like projections sometimes equipped with bristles, and another pair of feathery appendages through which oxygen is absorbed. Within its body, each segment has a pair of tubes opening to the exterior from which waste is secreted. A gut, a large blood vessel and a nerve cord run from front to end through all the segments, linking and coordinating them.

Fossils can only tell us so much. Even the exceptionally well-preserved remains of Ediacara offer no clue about the connection between the segmented worms and the other early groups. However, there is one further category of evidence to be looked at – the larvae. The segmented worms have spherical larvae with a belt of cilia round their middles and a long tuft on top. These are almost identical to the larva of some molluscs, a strong indication that back in time the two groups sprang from common stock. The echinoderms, on the other hand, have a larva that is quite different, with a twist to its structure and winding bands of cilia around it. This group must have separated from the ancestral flatworms at a very early stage indeed, long before the split between the molluscs and the segmented worms. Geneticists, analysing the DNA of each of these groups, now confirm these deductions and reveal that there are two major groupings of bilaterally symmetrical animals. Octopus, crabs and flatworms form one group, while echinoderms, tunicates and all the backboned animals make up the other.

Segmentation may have developed as a way of enabling worms to increase their efficiency as burrowers in mud. A line of separate limbs down each side is clearly a very effective structure for this purpose and it could have been acquired by repeating the simple body unit to form a chain. The change must have taken place long before Ediacaran times, for when those rocks were deposited the fundamental invertebrate divisions were already established The Ediacaran fossils, in Australia where they were first discovered and in Britain, Newfoundland, Namibia and Siberia, now confirm these deductions. Thereafter their history remains virtually invisible for a 100 million years. Only after this vast span do we reach the period, 600 million years ago, represented by the Moroccan deposits and others throughout the world. By that time many organisms had, as we have seen, developed shells from which we can deduce their existence and shape, but not much more.

However, there is one exceptional fossil site dating from only a little later than those of Ediacara that provides far more detailed information about the bodies of animals than can come from mere shells. In the Rocky Mountains of British Columbia, the Burgess Pass crosses a ridge between two high snowy peaks. Close to its crest lies an outcrop of particularly fine-grained shales, and in these have been discovered some of the most perfectly preserved fossils in the world. The shales were laid down about 530 million years ago, close to the beginning of the Cambrian period in a basin of the seafloor at a depth of about 150 metres. It must have been sheltered by a submarine ridge, for there were no currents to disturb the fine sediments on the floor or to bring down oxygenated water from nearer the surface. Few animals lived in those dark stagnant waters. There are no signs of tracks or burrows. Once in a while, however, mud from the ridge above slipped down in a turbid cloud, carrying with it all kinds of small creatures, and dumped them there. Since there was neither oxygen to fuel the processes of decay nor any scavenging animals to feed on the bodies and destroy them, many of the tiny carcasses remained complete as the settling mud particles slowly entombed them, preserving even their softest body parts. Eventually the entire deposit became consolidated into shale. Earth movements elevated and folded great areas of these marine deposits during the building of the Rocky Mountains. Many parts of them were distorted and crushed until most traces of life in them were obliterated. But miraculously, this one small patch survived virtually undamaged.


Velvet worm (Peripatus novaezealandiae). Velvet worms are known as ‘living fossils’, having remained the same for approximately 570 million years.

The range of creatures it contains is far wider than that found in rocks of a similar age at any other site so far discovered. There are the jellyfish that Ediacara would lead us to expect. There are echinoderms, brachiopods, primitive molluscs and half a dozen species of segmented worms – further representatives of the lineage that stretches from the beaches of Ediacara to the Barrier Reef of today.

There are also several creatures which were rather more mysterious. Among the most abundant of these was a strange segmented creature with what seemed to be a line of legs on its underside. It looked rather like a shrimp, though mysteriously none of the species had a head. It was given the name Anomalorcaris: strange shrimp. There were also small disc-shaped fossils marked with lines radiating from its centre that looked somewhat like a tiny slice of pineapple, which was initially thought to be some kind of jellyfish. Perhaps strangest of all, there was an elongated segmented animal that appeared to have seven pairs of spiny stilt-like legs, and seven flexible tentacles along its back, each ending in a tiny mouth. It seemed so strange as to be almost nightmarish, and the researcher who studied it accordingly called it Hallucigenia.

Subsequent work, however, showed that these oddities were not the founder members of some wholly unsuspected animal groups. A very exceptional specimen of Anomalocaris showed that the ‘strange shrimps’ were not complete animals but just the forelimbs belonging to a much bigger creature that used them to grab its prey. And the pineapple slice was eventually shown to have in its centre minute teeth. It was a mouth that belonged to the same animal as the tentacles. Both these pieces of Anomalorcaris’ body apparently had a more heavily strengthened exoskeleton and so regularly became separated from the animal’s more easily decayed body. As for Hallucigenia, further research on other specimens showed that it had been reconstructed in an upside-down position. The spindly legs were in fact protective dorsal spines, and what had been considered tentacles were in reality its legs. It is now thought that it may be the first known member of a strange group called the lobopods which today includes odd little creatures called velvet worms.

The great variety of creatures in the Burgess Shales is a reminder of how incomplete our knowledge of all fossil faunas actually is. The ancient seas contained many more kinds of animals than we can ever know. In this one site, conditions allowed a uniquely large proportion to be preserved, but even this is only a hint of what must have once existed.

The Burgess Shales also contain superbly preserved examples of trilobites like those in the Moroccan limestones. Their body armour was constructed partly of calcium carbonate and strengthened by a horny substance called chitin, a material that forms the external skeletons of insects. But chitin, unlike skin, does not expand, so any animal with such an external chitinous skeleton has to shed it regularly if it is to grow – as indeed insects do today. Many of the trilobite fossils we find are in fact these empty suits of armour. Sometimes they are concentrated in great drifts, having been sorted by sea currents, as shells sometimes are when they are swept up on beaches today. The underwater avalanches in the Burgess Shales Basin, however, swept down not just discarded armour but living trilobites and buried them. Mud particles filtered into the animals’ bodies and preserved the finest details of their anatomy. So in them we can still see the paired jointed legs that are attached to each body segment, the feathery gill associated with each leg, two feelers at the front of the head, and the gut running the entire length of the body. Even the muscle fibres along the back, which enabled the animal to roll itself up into a ball, are still recognisable in some exceptional specimens.

Trilobites, as far as we know, were the first creatures on earth to develop high-definition eyes. They are mosaics, a cluster of separate components, each with its own lens of crystalline calcite orientated in the precise position in which it transmits light most efficiently, much like the eyes of today’s insects. One eye may contain 15,000 elements, and would have given its owner an almost hemispherical field of view. Late in the dynasty, some species developed an even more sophisticated kind of eye and one that has never been paralleled by any other animal. Here the components are fewer but larger. Their lenses are much thicker and it is thought that these species lived where there was little light and needed thick lenses to collect and concentrate what light there was. However, the optical properties of a simple calcite lens in contact with water are such that it transmits light in a diffused way and cannot bring it to a sharply focused point. To do this, a two-part lens is needed which has a waved surface at the junction between its two elements. And this is exactly what these trilobites evolved. The lower element of the double lens was formed by chitin and the surface between the two conforms to the mathematical principle that human scientists did not discover until 300 years ago when they tried to correct the spherical aberration of lenses in their newly invented telescopes.

As the trilobites spread through the seas of the world, they diversified into a great number of species. Many seem to have lived on the seafloor, chomping their way through mud. Some colonised the deep seas where there was little light and lost their eyes altogether. Others, to judge from the shape of their limbs, may well have paddled about, legs uppermost, scanning the seafloor below with their large eyes.

In due course, as creatures of many kinds and varying ancestries came to live on the bottom of the seas, the trilobites lost their supremacy. Two hundred and fifty million years ago, their dynasty came to an end. One relation alone survives, the horseshoe crab. It’s a misleading name for it is not a crab and only half its shell bears any resemblance to a horseshoe. Measuring 30 centimetres or so across, it is many times bigger than most known trilobites and its armour no longer shows any signs of segmentation. Its front section is a huge domed shield, on the front of which are two bean-shaped compound eyes. A roughly rectangular plate, hinged to the back of the shield, carries a sharp spike of a tail. But beneath its shell, the animal’s segmentation is clear. It has several pairs of jointed legs with pincers on the end, and behind these there are plate-like gills, large and flat like the leaves of a book.


Tower-eyed trilobite (Erbenochile erbeni) from the Timrahrhart Formation, Morocco.


Horseshoe crab (Limulus polyphemus) group spawning at high tide at sunset, Cape May, New Jersey.

Horseshoe crabs are seldom seen, for they live at considerable depths. Some inhabit Southeast Asian waters, others are found in the seas along the North Atlantic coast of America. Every spring, they migrate towards the coast. Then on three successive nights, when the moon is full and the tides are high, hundreds of thousands emerge from the sea. The females, their huge shells glinting in the moonlight, move towards the shore, dragging smaller males behind them. Sometimes four or five males, in their anxiety to reach a female, cling to one another and form a chain. As she reaches the edge of the water, the female half buries herself in the sand. There she sheds her eggs and the males release sperm. For kilometre after kilometre along the dark beaches, the living tide of horseshoe crabs is so thick that they form a continuous strip, like a causeway of giant cobbles. The breakers sometimes overturn them and they lie in the sand, with their legs waving, their stiff tails slowly swivelling, in an effort to lever themselves right side up. Many fail and are abandoned by the receding tide to die as thousands more swim in the shallows, pressing forward to take their turn.

This scene must have been enacted every spring for several hundred million years. When it began, the land was without life of any kind, and on such beaches the eggs were safe from sea-dwelling marauders. Perhaps this is why the horseshoe crabs developed the habit. Today beaches are not quite so safe, for hordes of gulls and small wading birds congregate to share the prodigious feast. But many of the fertilised eggs remain buried deep among the sand grains where they will stay for a month until, once more, high water reaches this part of the beach, stirring the sand and releasing the larvae to swim freely in the sea.

Although the trilobites were so successful, they were by no means the only armoured creatures to develop from the segmented worms. So did a group that must have been among the most alarming of all marine monsters – the sea scorpions, called scientifically the Eurypterids. Some grew to a length of two metres and were the largest arthropods ever known to have existed. However, in spite of their appearance and huge claws, many of them were filter feeders. Presumably, their fearsome claws were used in fights between one another rather than in subduing prey. Like the trilobites, they disappeared at the end of the Permian period.

One group related to the trilobites did however survive and today is extremely successful. They differed in one seemingly trivial but nonetheless diagnostic characteristic. They have not one but two pairs of antennae on their heads. They lived alongside the trilobites, comparatively unobtrusively for hundreds of millions of years, and then, when the trilobite dynasty came to an end, it was they who took over. They are the crustaceans. Today there are about 35,000 species of crustacean – seven times as many as there are of birds. Most prowl among the rocks and reefs – crabs, shrimps, prawns and lobsters. Some – the barnacles – have taken up a static life. Others – the krill which forms the food of whales – swim in vast shoals.


Robber crab (Birgus latro) climbing coconut tree, Aldabra Seychelles.

An external skeleton is highly versatile; it serves the tiny water flea as well as it does the giant Japanese spider crab that measures over three metres from claw to claw. Each crustacean species modifies the shape of its many paired legs for particular purposes. Those at the front may become pincers or claws; those in the middle, paddles, walking legs or tweezers. Some have feathery branches, gills through which oxygen is absorbed from the water. Others develop attachments so that they can carry eggs.

The limbs, which are tubular and jointed, are operated by internal muscles. These extend from the end of one section, along its length, to a prong from the next section which projects across the joint. When the muscle contracts between these two attachment points, the limb hinges. Such joints can only move in one plane, but crustaceans deal with that limitation by grouping two or three on a limb, sometimes close together, each working in a different plane so that the free end of the limb can move in a complete circle.

The external shell, however, gives the crustaceans the same problem as it gave the trilobites. It will not expand, and since it completely encloses their bodies, the only way they can grow is to shed it periodically. As the time for the moult approaches, the animal absorbs much of the calcium carbonate from its shell into its blood. It secretes a new, soft and wrinkled skin beneath the shell. The outgrown armour splits at the back and the animal pulls itself out, leaving the shell more or less complete, like a translucent ghost of its former self. Now, because the animal’s skin is soft, it must hide, but it grows fast and swells its body by absorbing water and stretching out the wrinkles of its new carapace. Gradually this hardens so the animal can again venture into a hostile world.

The hermit crab partly avoids this complicated and hazardous process by having a shell-less hind part and protecting it with a discarded mollusc shell, slipping into a new one in a minute or so whenever it has the need.

The external skeleton has one incidental quality which has had momentous results. Mechanically, it works almost as well on land as it does in water, so that, providing a creature can find a way of breathing, there is little to prevent it walking straight out of the sea and up the beach. Many crustaceans, indeed, have done so – sand shrimps and beach hoppers stay quite close to the sea; and pill bugs and penny sows have colonised moist ground throughout the land.

The most spectacular of all these land-living crustaceans is the robber crab. It is found on islands in the Indian Ocean and the western parts of the Pacific. At the back of its main carapace, at the junction with the first segment of its abdomen, there is an opening to an air chamber lined with moist puckered skin through which the animal absorbs oxygen. This monster is so big it can embrace the trunk of a palm tree between its outstretched legs. It climbs with ease, and once in the palm’s crest, cuts down with its gigantic pincers the young coconuts on which it feeds. It has to return to the sea to lay its eggs, but otherwise it is entirely at home on land.

Other descendants of the marine invertebrates have also left the water. Among the molluscs there are the snails and the shell-less slugs, but these emerged from water relatively recently in the group’s history. The first to make the move to land were probably descendants of the segmented worms, the millipedes. Their droppings have been found fossilised in the rocks of Shropshire. They were followed by pioneers which recent DNA studies show to have been crustaceans. And some of these made such a success of life in their new surroundings that they eventually gave rise to the most numerous and diverse group of all land animals – the insects.


Life on Earth

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