Bilateria

The Bilateria are all animals having a bilateral symmetry, i.e. they have a front and a back end, as well as an upside and downside. Radially symmetrical animals like jellyfish have a topside and downside, but no front and back. The bilateralians are a subregnum (a major group) of animals, including the majority of phyla; the most notable exceptions are the sponges and cnidarians. For the most part, Bilateria have bodies that develop from three different germ layers, called the endoderm, mesoderm, and ectoderm. From this they are called triploblastic. Nearly all are bilaterally symmetrical, or approximately so. The most notable exception is the echinoderms, which achieve near-radial symmetry as adults, but are bilaterally symmetrical as larvae.

Except for a few highly reduced forms, the Bilateria have complete digestive tracts with separate mouth and anus. Most Bilateria also have a type of internal body cavity, called a coelom. It was previously thought that acoelomates gave rise to the other group, but there is some evidence now that in the main acoelomate phyla (flatworms and gastrotrichs) the absence could be secondary.

Evolution

The hypothetical last common ancestor of all bilaterians is termed the "Urbilaterian". There is some debate about its appearance. The first evidence of bilateria in the fossil record comes from trace fossils in Ediacaran sediments, and the first bona fide bilaterian fossil is Kimberella, dating to . Earlier fossils are controversial; The fossil Vernanimalcula may be the earliest known bilaterian, but may also represent an infilled bubble. Fossil embryos are known from around the time of Vernanimalcula but none of these have bilaterian affinities.

Phylogeny

There are two or more superphyla (main lineages) of Bilateria. The deuterostomes include the echinoderms, hemichordates, chordates, and possibly a few smaller phyla. The protostomes include most of the rest, such as arthropods, annelids, mollusks, flatworms, and so forth. There are a number of differences, most notably in how the embryo develops. In particular, the first opening of the embryo becomes the mouth in protostomes, and the anus in deuterostomes. Many taxonomists now recognize at least two more superphyla among the protostomes, Ecdysozoa (molting animals) and Lophotrochozoa. Some taxonomists also recognize another protostome superphylum, Platyzoa, while others would include the Platyzoans in Lophotrochozoa or not include them in any superphylum. The arrow worms (Chaetognatha) have proven particularly difficult to classify, with some taxonomists placing them among the deuterostomes and others placing them among the protostomes. The two most recent studies to address the question of chaetognath origins (Helfenbein et al. 2004 and Papillon et al. 2004) support protostome affinities.

A proposed phylogeny of the Bilateria after Nielsen is as follows. An alternate phylogeny suggests a basal group called the ecdysozoa.

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Notes

Other references

Helfenbein, Kevin G., H. Matthew Fourcade, Rohit G. Vanjani, and Jeffrey L. Boore (2004). The mitochondrial genome of Paraspadella gotoi is highly reduced and reveals that chaetognaths are a sister group to protostomes. Proceedings of the National Academy of Sciences of the United States of America 101(29), July 20 2004: 10639-10643.
Papillon, Daniel, Yvan Perez, Xavier Caubit, and Yannick Le Parco (2004). Identification of chaetognaths as protostomes is supported by the analysis of their mitochondrial genome. Molecular Biology and Evolution 21(11), November 2004: 2122-2129.