Bilateria

Taxobox
name = Bilateria
fossil_range = Ediacaran - Recent

---

image_width = 225px
image_caption = A Bowfin , "Amia calva"
regnum = Animalia
subregnum = Eumetazoa
unranked_phylum = Bilateria
unranked_kingdom_authority = Hatschek, 1888
subdivision_ranks = Phyla
subdivision =
* Orthonectida
* Rhombozoa
* Acoelomorpha
* Chaetognatha
* Superphylum Deuterostomia
** Chordata
** Hemichordata
** Echinodermata
** Xenoturbellida
** Vetulicolia †
* "Protostomia (unranked)"
** Superphylum Ecdysozoa
*** Kinorhyncha
*** Loricifera
*** Priapulida
*** Nematoda
*** Nematomorpha
*** Lobopodia †
*** Onychophora
*** Tardigrada
*** Arthropoda
** Superphylum Platyzoa
*** Platyhelminthes
*** Gastrotricha
*** Rotifera
*** Acanthocephala
*** Gnathostomulida
*** Micrognathozoa
*** Cycliophora
** Superphylum Lophotrochozoa
*** Sipuncula
*** Hyolitha †
*** Nemertea
*** Phoronida
*** Bryozoa
*** Entoprocta
*** Brachiopoda
*** Mollusca
*** Annelida
*** Echiura

The Bilateria (pronEng|ˌbaɪləˈtɪəriə) 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". [Knoll, Andrew H. and Sean B. Carroll. (1999) Early Animal Evolution: Emerging Views from Comparative Biology and Geology. "Science". 25 June 1999: Vol. 284. no. 5423, pp. 2129 - 2137. Found at [http://cas.bellarmine.edu/tietjen/Ecology/early_animal_evolution.htm] - URL retrieved November 15, 2006] [Balavoine, Guillaume, & Adoutte, Andre. 2003. The segmented Urbilateria: A testable scenario. "Integrative & Comparative Biology" 43: 137-147. Found at [http://icb.oxfordjournals.org/cgi/content/full/43/1/137] - URL retrieved November 15, 2006] 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 Ma|555. [For refs see "Ediacara biota"] Earlier fossils are controversial; The fossil "Vernanimalcula" may be the earliest known bilaterian, but may also represent an infilled bubble. [For refs see "Vernanimalcula"] Fossil embryos are known from around the time of "Vernanimalcula" (Ma|580), but none of these have bilaterian affinities. [For refs see Fossil embryos]

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 [ [http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/I/Invertebrates.html The Invertebrate Animals] ] 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. [Nielsen, C. 2001. Animal Evolution: Interrelationships of the Living Phyla. Second Edition. Oxford University Press, Oxford.] An alternate phylogeny suggests a basal group called the ecdysozoa.

clade| style=font-size:75%;line-height:75%
1=clade
label1=Protostomia |1=clade |label1=Spiralia |1=clade |label1=Schizocoelia |1=clade |1=Sipuncula |label2=Articulata |2=clade |1=Mollusca |label2=Euarticulata |2=clade |1=Annelida |label2=Panarthropoda |2=clade |1=Onychophora |2=clade |1=Tardigrada |2=Arthropoda |label2=Bryozoa |2=clade |1=Ectoprocta |2=Entoprocta |label3=Parenchymia |3=clade |1=Platyhelminthes |2=Nemertini |label4=Gnathifera |4=clade |1=Chaetognatha |2=clade |1=Gnathostomulida |2=Rotifera |label2=Cycloneuralia |2=clade |1=Gastrotricha |label2=Introverta |2=clade |1=Nematoda |2=Nematomorpha |label3=Cephalorhyncha |3=clade |1=Priapulida |2=clade |1=Kinorhyncha |2=Loricifera |label2=Deuterostomia |2=clade |1=clade |1=Phoronida |2=Brachiopoda |2=clade |1=Echinodermata |2=Pterobranchia |3=clade |1=Enteropneusta |2=Chordata

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.

External links

* [http://tolweb.org/tree?group=Bilateria&contgroup=Animals Tree of Life web project - Bilateria]
* [http://www.ucmp.berkeley.edu/phyla/metazoasy.html University of California Museum of Paleontology - Systematics of the Metazoa]