Friday, March 14, 2008

Origin and fossil record

Origin and fossil record
Animals are generally considered to have evolved from a flagellated eukaryote. Their closest known living relatives are the choanoflagellates, collared flagellates that have a morphology similar to the choanocytes of certain sponges. Molecular studies place animals in a supergroup called the opisthokonts, which also include the choanoflagellates, fungi and a few small parasitic protists. The name comes from the posterior location of the flagellum in motile cells, such as most animal spermatozoa, whereas other eukaryotes tend to have anterior flagella.

The first fossils that might represent animals appear towards the end of the Precambrian, around 610 million years ago, and are known as the Ediacaran or Vendian biota. These are difficult to relate to later fossils, however. Some may represent precursors of modern phyla, but they may be separate groups, and it is possible they are not really animals at all. Aside from them, most known animal phyla make a more or less simultaneous appearance during the Cambrian period, about 542 million years ago. It is still disputed whether this event, called the Cambrian explosion, represents a rapid divergence between different groups or a change in conditions that made fossilization possible. However some paleontologists and geologists would suggest that animals appeared much earlier than previously thought, possibly even as early as 1 billion years ago. Trace fossils such as tracks and burrows found in Tonian era strata in India indicate the presence of triploblastic worm like metazoans roughly as large (about 5mm wide) and complex as earthworms.[6] In addition during the beginning of the Tonian period around 1 billion years ago (roughly the same time that the trace fossils previously discussed in this article date back to) there was a decrease in Stromatolite diversity which may indicate the appearance of grazing animals during this time as Stromatolites also increased in diversity shortly after the end-Ordovician and end-Permian rendered large amounts of grazing marine animals extinct and decreased shortly after their populations recovered. However some other scientists doubt that these fossils are authentic and have suggested these trace fossils are just the result of natural processes such as erosion.[citation needed]


Groups of animals
The sponges (Porifera) diverged from other animals early. As mentioned above, they lack the complex organization found in most other phyla. Their cells are differentiated, but in most cases not organized into distinct tissues. Sponges are sessile and typically feed by drawing in water through pores. Archaeocyatha, which have fused skeletons, may represent sponges or a separate phylum.

Among the eumetazoan phyla, two are radially symmetric and have digestive chambers with a single opening, which serves as both the mouth and the anus. These are the Cnidaria, which include sea anemones, corals, and jellyfish, and the Ctenophora or comb jellies. Both have distinct tissues, but they are not organized into organs. There are only two main germ layers, the ectoderm and endoderm, with only scattered cells between them. As such, these animals are sometimes called diploblastic. The tiny Placozoans are similar, but they do not have a permanent digestive chamber.

The remaining animals form a monophyletic group called the Bilateria. For the most part, they are bilaterally symmetric, and often have a specialized head with feeding and sensory organs. The body is triploblastic, i.e. all three germ layers are well-developed, and tissues form distinct organs. The digestive chamber has two openings, a mouth and an anus, and there is also an internal body cavity called a coelom or pseudocoelom. There are exceptions to each of these characteristics, however - for instance adult echinoderms are radially symmetric, and certain parasitic worms have extremely simplified body structures.

Genetic studies have considerably changed our understanding of the relationships within the Bilateria. Most appear to belong to four major lineages: Deuterostomes, Ecdysozoa, Platyzoa or Lophotrochozoa. In addition to these, there are a few small groups of bilaterians with relatively similar structure that appear to have diverged before these major groups. These include the Acoelomorpha, Rhombozoa, and Orthonectida. The Myxozoa, single-celled parasites that were originally considered Protozoa, are now believed to have developed from the Bilateria as well.
Deuterostomes

Superb Fairy-wren, Malurus cyaneusDeuterostomes differ from the other Bilateria, called protostomes, in several ways. In both cases there is a complete digestive tract. However, in protostomes the initial opening (the archenteron) develops into the mouth, and an anus forms separately. In deuterostomes this is reversed. In most protostomes cells simply fill in the interior of the gastrula to form the mesoderm, called schizocoelous development, but in deuterostomes it forms through invagination of the endoderm, called enterocoelic pouching. Deuterostomes also have a dorsal, rather than a ventral, nerve chord and their embryos undergo different cleavage.

All this suggests the deuterostomes and protostomes are separate, monophyletic lineages. The main phyla of deuterostomes are the Echinodermata and Chordata. The former are radially symmetric and exclusively marine, such as starfish, sea urchins, and sea cucumbers. The latter are dominated by the vertebrates, animals with backbones. These include fish, amphibians, reptiles, birds, and mammals.

In addition to these, the deuterostomes also include the Hemichordata or acorn worms. Although they are not especially prominent today, the important fossil graptolites may belong to this group.

The Chaetognatha or arrow worms may also be deuterostomes, but more recent studies suggest protostome affinities.

Ecdysozoa

Yellow-winged Darter, Sympetrum flaveolumThe Ecdysozoa are protostomes, named after the common trait of growth by moulting or ecdysis. The largest animal phylum belongs here, the Arthropoda, including insects, spiders, crabs, and their kin. All these organisms have a body divided into repeating segments, typically with paired appendages. Two smaller phyla, the Onychophora and Tardigrada, are close relatives of the arthropods and share these traits.

The ecdysozoans also include the Nematoda or roundworms, the second largest animal phylum. Roundworms are typically microscopic, and occur in nearly every environment where there is water. A number are important parasites. Smaller phyla related to them are the Nematomorpha or horsehair worms, which are invisible to the unaided eye, and the Kinorhyncha, Priapulida, and Loricifera. These groups have a reduced coelom, called a pseudocoelom.

The remaining two groups of protostomes are sometimes grouped together as the Spiralia, since in both embryos develop with spiral cleavage.


Platyzoa

Bedford's flatworm, Pseudobiceros bedfordiThe Platyzoa include the phylum Platyhelminthes, the flatworms. These were originally considered some of the most primitive Bilateria, but it now appears they developed from more complex ancestors.[7]

A number of parasites are included in this group, such as the flukes and tapeworms. Flatworms are acoelomates, or lack a body cavity, as do their closest relatives, the microscopic Gastrotricha.[8]

The other platyzoan phyla are microscopic and pseudocoelomate. The most prominent are the Rotifera or rotifers, which are common in aqueous environments. They also include the Acanthocephala or spiny-headed worms, the Gnathostomulida, Micrognathozoa, and possibly the Cycliophora.[9] These groups share the presence of complex jaws, from which they are called the Gnathifera.

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