Bilateria

The bilateria /ˌbaɪləˈtɪəriə/ or bilaterians are animals with bilateral symmetry as an embryo, i.e.
Bilaterians
having a left and a right side that are mirror images of each other. This also means they have a
Temporal range: Ediacaran–Present,
head and a tail (anterior-posterior axis) as well as a back and a belly (ventral-dorsal axis).[2]
Nearly all are bilaterally symmetrical as adults as well; the most notable exception is the 560–0 Ma[1]
echinoderms, which achieve near-radial symmetry as adults, but are bilaterally symmetrical PreЄ Є O S D C P T J K PgN
during embryonic development.

Most animals are bilaterians, excluding sponges, ctenophores, placozoans and cnidarians. For the
most part, bilateral embryos are triploblastic, having three germ layers: endoderm, mesoderm, and
ectoderm. Except for a few phyla (i.e. flatworms and gnathostomulids), bilaterians have complete
digestive tracts with a separate mouth and anus. Some bilaterians lack body cavities (acoelomates,
i.e. Platyhelminthes, Gastrotricha and Gnathostomulida), while others display primary body
cavities (deriving from the blastocoel, as pseudocoeloms) or secondary cavities (that appear de
novo, for example the coelom).[3][4]

Contents
Body plan Diversity of bilaterians.
Evolution
Scientific classification
Phylogeny
Kingdom: Animalia
Evolutionary origin
Subkingdom: Eumetazoa
See also
Notes Clade: ParaHoxozoa

References Clade: Bilateria

External links Hatschek, 1888

Phyla

Body plan Proarticulata †

Some of the earliest bilaterians Xenacoelomorpha
were wormlike, and a bilaterian Nephrozoa:
body can be conceptualized as a
cylinder with a gut running Superphylum
between two openings, the mouth Deuterostomia
and the anus. Around the gut it has
an internal body cavity, a coelom Chordata
or pseudocoelom.[a] Animals with Saccorhytida †
this bilaterally symmetric body
Ambulacraria (unranked)
Idealised wormlike bilaterian body plan. With a cylindrical body plan have a head (anterior) end and
and a direction of movement the animal has head and tail ends. a tail (posterior) end as well as a Hemichordata
Sense organs and mouth form the basis of the head. Opposed back (dorsal) and a belly (ventral);
circular and longitudinal muscles enable peristaltic motion.
Echinodermata
therefore they also have a left side
and a right side.[6][7] Cambroernida †
Protostomia (unranked)
Having a front end means that this part of the body encounters stimuli, such as food, favouring
cephalisation, the development of a head with sense organs and a mouth.[8] The body stretches Superphylum
back from the head, and many bilaterians have a combination of circular muscles that constrict the
Ecdysozoa
body, making it longer, and an opposing set of longitudinal muscles, that shorten the body;[7]
these enable soft-bodied animals with a hydrostatic skeleton to move by peristalsis.[9] They also Cycloneuralia
have a gut that extends through the basically cylindrical body from mouth to anus. Many (unranked)
bilaterian phyla have primary larvae which swim with cilia and have an apical organ containing
sensory cells. However, there are exceptions to each of these characteristics; for example, adult Scalidophora
echinoderms are radially symmetric (unlike their larvae), and certain parasitic worms have (unranked)
extremely simplified body structures.[6][7]
Kinorhyncha
Evolution Loricifera
The hypothetical most recent common ancestor of all bilateria is termed the "Urbilaterian".[10][11] Priapulida
The nature of the first bilaterian is a matter of debate. One side suggests that acoelomates gave
Nematoida
rise to the other groups (planuloid-aceloid hypothesis by Ludwig von Graff, Elie Metchnikoff,
Libbie Hyman, or Luitfried von Salvini-Plawen), while the other poses that the first bilaterian was (unranked)
a coelomate organism and the main acoelomate phyla (flatworms and gastrotrichs) have lost body
Nematoda
cavities secondarily (the Archicoelomata hypothesis and its variations such as the Gastrea by
Haeckel or Sedgwick, the Bilaterosgastrea by Gösta Jägersten, or the Trochaea by Nielsen). Nematomorpha
Panarthropoda
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 555 million years ago.[12] Earlier (unranked)
fossils are controversial; the fossil Vernanimalcula may be the earliest known bilaterian, but may
Onychophora
also represent an infilled bubble.[13][14] Fossil embryos are known from around the time of
Vernanimalcula (580 million years ago), but none of these have bilaterian affinities.[15] Burrows Tactopoda
believed to have been created by bilaterian life forms have been found in the Tacuarí Formation of
Tardigrada
Uruguay, and are believed to be at least 585 million years old.[16]
Arthropoda

Phylogeny Spiralia (unranked)

There are two main lineages, superphyla, of Bilateria. The deuterostomes include the Chaetognatha
echinoderms, hemichordates, chordates, and a few smaller phyla. The protostomes include most
Mesozoa (unranked)
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 Monoblastozoa
embryo becomes the mouth in protostomes, and the anus in deuterostomes. Many taxonomists
now recognize at least two more superphyla among the protostomes, Ecdysozoa[17] (molting Superphylum
animals) and Spiralia.[17][18][19][20] The arrow worms (Chaetognatha) have proven difficult to Platyzoa
classify; recent studies place them in the gnathifera.[21][22][23]
Acanthocephala
A modern (2011) consensus phylogenetic tree for Bilateria is shown below, although the positions Cycliophora
of certain clades are still controversial (dashed lines) and the tree has changed considerably
Gastrotricha
between 2000 and 2010.[24][23][25][26][27] The following tree depicts the traditional protostome–
deuterostome dichotomy: Gnathostomulida
Micrognathozoa
Platyhelminthes
Rotifera
Superphylum
Lophotrochozoa

Annelida
Brachiopoda
Bryozoa
Dicyemida
Entoprocta
Hyolitha †
Mollusca
Nemertea
Orthonectida
Phoronida

Synonyms

Triploblasts Lankester, 1873

Planulozoa
680 mya
Cnidaria

Bilateria
†Proarticulata
Xenacoelomorpha
Xenoturbellida

Acoelomorpha
Nemertodermatida
 Acoela

Nephrozoa
650 mya Cephalochordata

Chordata Urochordata
Olfactores

Craniata/Vertebrata

Echinodermata

Deuterostomia
Ambulacraria
Hemichordata

†Cambroernida

†Saccorhytus coronarius

†Vetulocystids

†Vetulicolians

Nematoda
Nematoida

Nematomorpha

Loricifera

Onychophora

Panarthropoda Tardigrada
Ecdysozoa Tactopoda
>529 mya
Arthropoda

Scalidophora

Priapulida

Protostomia
610 mya

Kinorhyncha

Spiralia
Rotifera and allies
Gnathifera

Chaetognatha

Platytrochozoa
 580 mya
Platyhelminthes and allies

Mollusca

Lophotrochozoa
550 mya
Annelida and allies

†Kimberella

A different hypothesis is that the Ambulacraria are sister to Xenacoelomorpha together forming the Xenambulacraria. The phylogenetic tree
shown below depicts this proposal. Nephrozoa is shown here sans Ambulacraria. Also Deuterostomes was proposed to be obsolete
altogether.[28][29][30] It is indicated when approximately clades radiated into newer clades in millions of years ago (Mya).[31] While the below
tree depicts a chordates as a sister group to protostomia according to analyses by Philippe et al., the authors nonetheless caution that "the
support values are very low, meaning there is no solid evidence to refute the traditional protostome and deuterostome dichotomy." [32]
ParaHoxozoa
680 mya
Placozoa

Cnidaria

Bilateria
†Proarticulata
Xenacoelomorpha
Xenoturbellida

Acoelomorpha
Nemertodermatida

Acoela

Xenambulacraria Echinodermata

Ambulacraria
Hemichordata

†Cambroernida

Nephrozoa
650 mya Cephalochordata

Chordata Urochordata
Olfactores

Craniata/Vertebrata

Protostomia Ecdysozoa
610 mya
>529 mya Nematoda
Nematoida

Nematomorpha

Loricifera

Panarthropoda
Onychophora

Tactopoda
 Tardigrada

Arthropoda

Priapulida

Scalidophora

Kinorhyncha

Rotifera and allies

Gnathifera

Chaetognatha

Spiralia Platyhelminthes and allies

Mollusca
Platytrochozoa
580 mya
Lophotrochozoa
550 mya
Annelida and allies

†Kimberella

†Saccorhytus coronarius

†Vetulocystids

†Vetulicolians

Evolutionary origin
The original bilaterian is hypothesized to have been a bottom dwelling worm with a single body opening.[5] It may have resembled the planula
larvae of some cnidaria, which have some bilateral symmetry.[33]

See also
Embryological origins of the mouth and anus

Notes
a. The earliest Bilateria may have had only a single opening, and no coelom.[5]

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External links
Tree of Life web project — Bilateria (http://tolweb.org/tree?group=Bilateria&contgroup=Animals)
University of California Museum of Paleontology — Systematics of the Metazoa (http://www.ucmp.berkeley.edu/phyla/metazoa
sy.html)

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