ANIMAL HISTORY

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Plant and Animal Evolution
Classification Plant Evolution Animal Evolution Homology
The History of Animal Evolution
For many people animals are perhaps the most familiar, and most interesting, of living things. This may be because we are animals ourselves. As such, we have a number of features in common with all the organisms placed in the animal kingdom, and these common features indicate that we have a shared evolutionary history.
All animals and plants are classified as multicellular eukaryotes: their bodies are made up of large numbers of cells, and microscopic inspection of these cells reveals that they contain a nucleus and a number of other organelles. Compared to prokaryotic organisms such as bacteria, plants and animals have a relatively recent evolutionary origin. DNA evidence suggests that the first eukaryotes evolved from prokaryotes, between 2500 and 1000 million years ago. That is, eukaryotes as a taxon date from the Proterozoic Era, the final Era of the Precambrian. Fossils of both simple unicellular and more complex multicellular organisms are found in abundance in rocks from this period of time. In fact, the name "Proterozoic" means "early life".Plants and animals both owe their origins to endosymbiosis, a process where one cell ingests another, but for some reason then fails to digest it. The evidence for this lies in the way their cells function. Both plant and animal rely on structures called mitochondria to release energy in their cells, using aerobic respiration to produce the energy-carrying molecule ATP. There is considerable evidence that mitochondria evolved from free-living aerobic bacteria: they are the size of bacterial cells; they divide independently of the cell by binary fission; they have their own genome in the form of a single circular DNA molecule; their ribosomes are more similar to those of bacteria than to the ribosomes found in the eukaryote cell's cytoplasm; and like chloroplasts they are enclosed by a double membrane as would be expected if they derived from bacterial cells engulfed by another cell.
Like the plants, animals evolved in the sea. And that is where they remained for at least 600 million years. This is because, in the absence of a protective ozone layer, the land was bathed in lethal levels of UV radiation. Once photosynthesis had raised atmospheric oxygen levels high enough, the ozone layer formed, meaning that it was then possible for living things to venture onto the land.
The oldest fossil evidence of multicellular animals, or metazoans, is burrows that appear to have been made by smooth, wormlike organisms. Such trace fossils have been found in rocks from China, Canada, and India, but they tell us little about the animals that made them apart from their basic shape.
The Ediacaran animals
The Cambrian "explosion" and the Burgess Shale
What caused the Cambrian "explosion"?
A foot on the land
The earliest vertebrates
Appearance of the fish
The jawless fish
Colonisation of the land
Problems encountered in the move to land
The evolution of amphibians
What drove amphibian evolution?
Early reptiles and the amniotic egg
The early mammals
Developments in the dinosaur lineage
Further developments in the early mammals
Taking wing: Archaeopteryx and the origin of the birds
The end of the dinosaur age
The appearance of modern mammal groups
The Ediacaran animals
Between 620 and 550 million years ago (during the Vendian Period) relatively large, complex, soft-bodied multicellular animals appear in the fossil record for the first time. While found in several localities around the world, this particular group of animals is generally known as the Ediacaran fauna, after the site in Australia where they were first discovered.
The Ediacaran animals are puzzling in that there is little or no evidence of any skeletal hard parts i.e. they were soft-bodied organisms, and while some of them may have belonged to groups that survive today others don't seem to bear any relationship to animals we know. Although many of the Ediacaran organisms have been compared to modern-day jellyfish or worms, they have also been described as resembling a mattress, with tough outer walls around fluid-filled internal cavities - rather like a sponge.
As a group, Ediacaran animals had a flat, quilted appearance and many showed radial symmetry. They ranged in size form 1cm to >1m, and have been classified into three main groups on the basis of their shape: discoidal, frond-like, or ovate-elongate. The large variety of Ediacaran animals is significant, as it suggests there must have been a lengthy period of evolution prior to their first appearance in the fossil record.
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The Cambrian "explosion" and the Burgess Shale
The Ediacaran animals disappear from the fossil record at the end of the Vendian (544 million years ago). In their place we find representatives of almost all the modern phyla recognised today: sponges, jellyfish and corals, flatworms, molluscs, annelid worms, insects, echinoderms and chordates, plus many "lesser" phyla such as nemertean worms. These "modern" organisms appear relatively quickly in the geological time scale, and their abrupt appearance is often described as the "Cambrian explosion" however, bear in mind that the fossil record of the "explosion" is spread over about 30 million years. I keep taking things out of brackets because it is interesting relevant and memorable
One of the most famous assemblages of Cambrian fossils comes from the Burgess Shale of British Colombia. The rocks of the Burgess Shale were laid down in the middle Cambrian, when the "explosion" had already been underway for several million years. They contain familiar animals such as trilobites, molluscs and echinoderms, but also the first appearance of brachiopods, and some odd animals, e.g. Opabinia, that may have belonged to extinct phyla. Even an early chordate, Pikaia, has been found in this fossil assemblage.
The Burgess Shale fossils are important, not only for their evidence of early variety among animal forms, but also because both soft parts of animals and their hard bodies (i.e. the whole animal) is preserved, and animals that were entirely soft-bodied. Preservation of soft-bodied organisms is rare, and in this case seems to have occurred when the animals were rapidly buried in a mudslide down into deep, anaerobic waters, where there was little bacterial decay. Prior to the discovery of this fossil assemblage, early in the 20th century, there was no evidence of soft-bodied animals from the Cambrian (remember that this is before the Ediacaran fauna were found).
These fossils also provide good evidence of predatory animals (e.g. Anomalocaris ), and therefore of complex predator-prey relationships. They also give insights into how evolution might have progressed relatively early in the history of multicellular animals, and in fact some authors view the Cambrian as a period of extreme "experimentation" and diversity.
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What caused the Cambrian "explosion"?
The cause of the proliferation of animal forms in the Cambrian is a matter of considerable debate among scientists. Some point to the increase in atmospheric oxygen levels that began around 2000 million years ago, supporting a higher metabolic rate and allowing the evolution of larger organisms and more complex body structures. Changed ocean chemistry would have played a part here, allowing for the first time the development of hard body parts such as teeth and supporting skeletons based on calcium carbonate (CaCO3), and also supporting higher levels of primary production as a result of increased concentrations of phosphates and nitrates. The mass extinction that marked the end of the Vendian period would have opened up ecological niches that the new animals exploited, as would habitat changes wrought by continental drift.
Genetic factors were also crucial. Recent research suggests that the period prior to the Cambrian explosion saw the gradual evolution of a "genetic tool kit" of genes (the homeobox or "hox" genes ) that govern developmental processes. Once assembled, this genetic tool kit enabled an unprecedented period of evolutionary experimentation -- and competition. Many forms seen in the fossil record of the Cambrian disappeared without trace. Future evolutionary change was then limited to acting on the body plans that remained in existence.
Recently many scientists have begun to question whether the Cambrian explosion was a real event, or a reflection of the patchiness of this ancient fossil record. Genetic data suggest that multicellular animals evolved around 1000 million years ago; this is supported by fossil embryos from rocks in China that date back 600 million years. These embryos are more complex than those of simple organisms such as sponges and jellyfish, which suggests that multicellular animals must have evolved much further back in time. In addition, trilobites were a very diverse group even early in the Cambrian, and some scientists suggest that this indicates that the arthropod group must have had a much earlier evolutionary origin.
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A foot on the land
Whatever their origins, animals may have ventured onto land early in the Cambrian. Previously scientists believed that animals did not begin to colonise the land until the Silurian (440 - 410 million years ago). However, the 2002 discovery of the footprints of animals that scuttled about on sand dunes about 530 million years ago has changed this view. These animals were arthropods, and resembled centipedes about the size of crayfish. They probably didn't live on land, instead coming ashore to mate or evade predators. At this time the only land plants appear to have resembled mosses .
The earliest vertebrates
Animals continued to diversify in the Ordovician seas (505 - 440 million years ago). They were mostly invertebrates, including graptolites , which were stick-like branching colonies of tiny animals, together with brachiopods, trilobites, cephalopods, corals, crinoids and conodonts. We now place the conodonts with the chordates, but for a long time they were known only by their tiny, but very common, teeth.
In terms of number of species invertebrates were by far the most common Ordovician animals - as they still are today. However, members of another taxon were also evolving in the Ordovician seas. These were the fish.
Appearance of the fish
Like the conodonts, fish are members of the chordate phylum because they display certain defining characteristics: a dorsal stiffening rod called the notochord, a dorsal nerve cord, pharyngeal gill slits and a tail that extends beyond the anus. However, fish are placed in the subphylum Vertebrata , because they also show the development of skeletal features such as a backbone, skull, and limb bones.
Not all the modern groups of fish were represented in the Ordovician oceans. At this time only the jawless fish had evolved from a chordate ancestor. The sharks and their relatives and two extinct groups, the placoderms (which had bony plates covering their heads) and the acanthodians (the first known jawed vertebrates, with a skeleton of cartilage) made their appearance in the Silurian. However, neither the sharks nor the agnathans became common until the Devonian. The other two living lineages, the ray-finned (e.g. carp and kahawai) and the lobe-finned fish (e.g. lungfish and the coelacanth), evolved during the Devonian period.
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The jawless fish
Agnathans, or jawless fish, were the earliest fish: an excellent fossil of Haikouichthys ercaicunensis dates back about 530 million years, to the Cambrian. Previously the earliest-known agnathans were dated to around 480 million years ago. Agnathans have traditionally been placed with the vertebrates due to the presence of a skull, although the modern forms such as hagfish lack a vertebral column. The earliest agnathans were Ostracoderms. They were bottom-feeders and were almost entirely covered in armour plates. When the sharks and bony fish began to evolve, around 450 million years ago, most ostracoderms became extinct. Only the lineage that produced the modern hagfish and lampreys survived.

Colonisation of the land
Fish continued to evolve during the Silurian period (440 - 410 million years ago). At the same time some groups of plants and animals took a major step as they colonised the land for the first time. We are not sure why this advance occurred, but it was probably the result of competition in the marine ecosystems, plus the opportunity to escape predators and the availability of new terrestrial niches.
Arthropods, which had ventured temporarily onto land 100 million years earlier, were the first animals to become more permanent colonists. Fossil footprints made in the sandy flats surrounding temporary lakes dating back about 420 million years have been found in Western Australia.
The arthropods were pre-adapted to life on land. By the time they moved ashore, they had already evolved lighter bodies and slim, strong legs that could support them against the pull of gravity. Their hard outer exoskeletons provided protection and would help to retain water, although the development of a waxy, waterproof cuticle was necessary for efficient water conservation.
Spiders, centipedes and mites were among the earliest land animals. Some of them were giants: the largest was Slimonia, the size of a man and a relative of the scorpions. This animal was still too big and too heavy and the walking legs too small to venture onto land for any length of time and so they lived in marginal marine (deltaic) environments