The Bilateria or bilaterians are

animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...

s with

bilateral symmetry Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pla ...

as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly and a back (ventral-dorsal axis). Nearly all are bilaterally symmetrical as adults as well; the most notable exception is the

echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea ...

s, which achieve secondary

pentaradial symmetry Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pla ...

as adults, but are bilaterally symmetrical during embryonic development. Most

s are bilaterians, excluding

sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through t ...

ctenophores Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), a ...

placozoa The Placozoa are a basal form of marine free-living (non-parasitic) multicellular organism. They are the simplest in structure of all animals. Three genera have been found: the classical ''Trichoplax adhaerens'', ''Hoilungia hongkongensis'', an ...

ns and

cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that th ...

ns. For the most part, bilateral embryos are

triploblastic Triploblasty is a condition of the gastrula in which there are three primary germ layers: the ectoderm, mesoderm, and endoderm. Germ cells are set aside in the embryo at the blastula stage, which are incorporated into the gonads during organo ...

, having three

germ layer A germ layer is a primary layer of cells that forms during embryonic development. The three germ layers in vertebrates are particularly pronounced; however, all eumetazoans (animals that are sister taxa to the sponges) produce two or three pr ...

endoderm Endoderm is the innermost of the three primary germ layers in the very early embryo. The other two layers are the ectoderm (outside layer) and mesoderm (middle layer). Cells migrating inward along the archenteron form the inner layer of the gast ...

mesoderm The mesoderm is the middle layer of the three germ layers that develops during gastrulation in the very early development of the embryo of most animals. The outer layer is the ectoderm, and the inner layer is the endoderm.Langman's Medical E ...

, and

ectoderm The ectoderm is one of the three primary germ layers formed in early embryonic development. It is the outermost layer, and is superficial to the mesoderm (the middle layer) and endoderm (the innermost layer). It emerges and originates from t ...

. Except for a few phyla (i.e.

flatworm The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a phylum of relatively simple bilaterian, unsegment ...

s and

gnathostomulid Gnathostomulids, or jaw worms, are a small phylum of nearly microscopic marine animals. They inhabit sand and mud beneath shallow coastal waters and can survive in relatively anoxic environments. They were first recognised and described in 1956. ...

s), bilaterians have complete

digestive tract The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans a ...

s with a separate

mouth In animal anatomy, the mouth, also known as the oral cavity, or in Latin cavum oris, is the opening through which many animals take in food and issue vocal sounds. It is also the cavity lying at the upper end of the alimentary canal, bounded on ...

and

anus The anus (Latin, 'ring' or 'circle') is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, d ...

. Some bilaterians lack body cavities ( acoelomates, i.e.

Platyhelminthes The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek language, Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a Phylum (biology), phylum of relati ...

Gastrotricha The gastrotrichs (phylum Gastrotricha), commonly referred to as hairybellies or hairybacks, are a group of microscopic (0.06-3.0 mm), worm-like, acoelomate animals, and are widely distributed and abundant in freshwater and marine environmen ...

and

Gnathostomulid Gnathostomulids, or jaw worms, are a small phylum of nearly microscopic marine animals. They inhabit sand and mud beneath shallow coastal waters and can survive in relatively anoxic environments. They were first recognised and described in 1956. ...

a), while others display primary body cavities (deriving from the

blastocoel The blastocoel (), also spelled blastocoele and blastocele, and also called cleavage cavity, or segmentation cavity is a fluid-filled or yolk-filled cavity that forms in the blastula during very early embryonic development. At this stage in mammals ...

, as pseudocoeloms) or secondary cavities (that appear ''de novo'', for example the

coelom The coelom (or celom) is the main body cavity in most animals and is positioned inside the body to surround and contain the digestive tract and other organs. In some animals, it is lined with mesothelium. In other animals, such as molluscs, it r ...

Body plan

Some of the earliest bilaterians were wormlike, and a bilaterian body can be conceptualized as a cylinder with a gut running between two openings, the mouth and the anus. Around the gut it has an internal body cavity, a

or pseudocoelom. Animals with this bilaterally symmetric

body plan A body plan, ( ), or ground plan is a set of morphological features common to many members of a phylum of animals. The vertebrates share one body plan, while invertebrates have many. This term, usually applied to animals, envisages a "blueprin ...

have a head (anterior) end and a tail (posterior) end as well as a back (dorsal) and a belly (ventral); therefore they also have a left side and a right side. Having a front end means that this part of the body encounters stimuli, such as food, favouring

cephalisation Cephalization is an evolutionary trend in which, over many generations, the mouth, sense organs, and nerve ganglia become concentrated at the front end of an animal, producing a head region. This is associated with movement and bilateral symmetr ...

, the development of a head with

sense organ A sense is a biological system used by an organism for sensation, the process of gathering information about the world through the detection of stimuli. (For example, in the human body, the brain which is part of the central nervous system re ...

s and a mouth. The body stretches back from the head, and many bilaterians have a combination of circular

muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of muscl ...

s that constrict the body, making it longer, and an opposing set of longitudinal muscles, that shorten the body; these enable soft-bodied animals with a

hydrostatic skeleton A hydrostatic skeleton, or hydroskeleton, is a flexible skeleton supported by fluid pressure. Hydrostatic skeletons are common among simple invertebrate organisms. While more advanced organisms can be considered hydrostatic, they are sometimes refe ...

to move by

peristalsis Peristalsis ( , ) is a radially symmetrical contraction and relaxation of muscles that propagate in a wave down a tube, in an anterograde direction. Peristalsis is progression of coordinated contraction of involuntary circular muscles, which ...

. Most bilaterians (Nephrozoans) have a gut that extends through the body from mouth to anus, while Xenacoelomorphs have a bag gut with one opening. Many bilaterian phyla have primary

larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...

e which swim with

cilia The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...

and have an apical organ containing sensory cells. However, there are exceptions to each of these characteristics; for example, adult echinoderms are radially symmetric (unlike their larvae), and certain

parasitic worms Parasitic worms, also known as helminths, are large macroparasites; adults can generally be seen with the naked eye. Many are intestinal worms that are soil-transmitted and infect the gastrointestinal tract. Other parasitic worms such as schi ...

have extremely

plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...

body structures.

Evolution

The hypothetical

most recent common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...

of all bilateria is termed the "

Urbilaterian The urbilaterian (from German ur- 'original') is the hypothetical last common ancestor of the bilaterian clade, i.e., all animals having a bilateral symmetry. Appearance Its appearance is a matter of debate, for no representative has been (o ...

". The nature of the first bilaterian is a matter of debate. One side suggests that acoelomates gave rise to the other groups (planuloid-aceloid hypothesis by

Ludwig von Graff Ludwig Graff de Pancsova (2 January 1851 – 6 February 1924), known as Ludwig von Graff, was an Austrian zoologist born in Pancsova. In 1871, he received his medical degree in Vienna, afterwards studying zoology at the University of Graz. In 18 ...

Elie Metchnikoff Elie and Earlsferry is a coastal town and former royal burgh in Fife, and parish, Scotland, situated within the East Neuk beside Chapel Ness on the north coast of the Firth of Forth, eight miles east of Leven. The burgh comprised the linked vi ...

Libbie Hyman Libbie Henrietta Hyman (December 6, 1888 – August 3, 1969), was a U.S. zoologist. She wrote numerous works on invertebrate zoology and the widely used '' A Laboratory Manual for Comparative Vertebrate Anatomy'' (1922, revised in 1942). Life B ...

, or ), while the other poses that the first bilaterian was a coelomate organism and the main acoelomate phyla (

s and

gastrotrich The gastrotrichs (phylum Gastrotricha), commonly referred to as hairybellies or hairybacks, are a group of microscopic (0.06-3.0 mm), worm-like, acoelomate animals, and are widely distributed and abundant in freshwater and marine environmen ...

s) have lost body cavities secondarily (the Archicoelomata hypothesis and its variations such as the Gastrea by

Haeckel Ernst Heinrich Philipp August Haeckel (; 16 February 1834 – 9 August 1919) was a German zoologist, naturalist, eugenicist, philosopher, physician, professor, marine biologist and artist. He discovered, described and named thousands of new sp ...

or Sedgwick, the Bilaterosgastrea by , or the Trochaea by Nielsen). One hypothesis is that the original bilaterian was a bottom dwelling worm with a single body opening, similar to ''

Xenoturbella ''Xenoturbella'' is a genus of very simple bilaterians up to a few centimeters long. It contains a small number of marine benthic worm-like species. The first known species (''Xenoturbella bocki'') was discovered in 1915 by Sixten Bock, but it wa ...

''. It may have resembled the planula larvae of some cnidaria, which have some bilateral symmetry.

Fossil record

The first evidence of bilateria in the fossil record comes from trace fossils in

Ediacaran The Ediacaran Period ( ) is a geological period that spans 96 million years from the end of the Cryogenian Period 635 million years ago (Mya), to the beginning of the Cambrian Period 538.8 Mya. It marks the end of the Proterozoic Eon, and th ...

sediments, and the first ''bona fide'' bilaterian fossil is ''

Kimberella ''Kimberella'' is an extinct genus of bilaterian known only from rocks of the Ediacaran period. The slug-like organism fed by scratching the microbial surface on which it dwelt in a manner similar to the gastropods, although its affinity with ...

'', dating to . Earlier fossils are controversial; the fossil ''

Vernanimalcula ''Vernanimalcula guizhouena'' is an acritarch dating from ; it was between 0.1 and 0.2 mm across (roughly the width of one or two human hairs). ''Vernanimalcula'' means "small spring animal", referring to its appearance in the fossil record ...

'' may be the earliest known bilaterian, but may also represent an infilled bubble.

Fossil embryos Embryo fossils are the preserved remains of unhatched or unborn organisms. Many fossils of the Doushantuo Formation have been interpreted as embryos; embryos are also common throughout the Cambrian fossil record. Preservation Taphonomic studies ...

are known from around the time of ''Vernanimalcula'' (), but none of these have bilaterian affinities. Burrows believed to have been created by bilaterian life forms have been found in the Tacuarí Formation of Uruguay, and are believed to be at least 585 million years old.

Phylogeny

The Bilateria has traditionally been divided into two main lineages or superphyla. The

deuterostome Deuterostomia (; in Greek) are animals typically characterized by their anus forming before their mouth during embryonic development. The group's sister clade is Protostomia, animals whose digestive tract development is more varied. Some exampl ...

s include the

hemichordate Hemichordata is a phylum which consists of triploblastic, enterocoelomate, and bilaterally symmetrical marine deuterostome animals, generally considered the sister group of the echinoderms. They appear in the Lower or Middle Cambrian and includ ...

chordate A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fiv ...

s, and a few smaller phyla. The

protostome Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryonic development. This nature has since been discovered to be extremely variable among Protostomia's memb ...

s include most of the rest, such as

arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...

annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...

mollusk Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant species of molluscs are recognized. The number of fossil species is e ...

s, and so forth. There are a number of differences, most notably in how the

embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...

develops. In particular, the first opening of the embryo becomes the mouth in protostomes, and the anus in deuterostomes. Many

taxonomists In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon) and these groups are given ...

now recognize at least two more superphyla among the protostomes,

Ecdysozoa Ecdysozoa () is a group of protostome animals, including Arthropoda (insects, chelicerata, crustaceans, and myriapods), Nematoda, and several smaller phyla. They were first defined by Aguinaldo ''et al.'' in 1997, based mainly on phylogenetic tr ...

(molting animals) and

Spiralia The Spiralia are a morphologically diverse clade of protostome animals, including within their number the molluscs, annelids, platyhelminths and other taxa. The term ''Spiralia'' is applied to those phyla that exhibit canonical spiral cleavage, a ...

. The arrow worms (

Chaetognatha The Chaetognatha or chaetognaths (meaning ''bristle-jaws'') are a phylum of predatory marine worms that are a major component of plankton worldwide. Commonly known as arrow worms, about 20% of the known Chaetognatha species are benthic, and ca ...

) have proven difficult to classify; recent studies place them in the gnathifera. The traditional division of Bilateria into

Deuterostomia Deuterostomia (; in Greek) are animals typically characterized by their anus forming before their mouth during embryonic development. The group's sister clade is Protostomia, animals whose digestive tract development is more varied. Some ex ...

and

Protostomia Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryonic development. This nature has since been discovered to be extremely variable among Protostomia's mem ...

was challenged when new morphological and molecular evidence found support for a sister relationship between the acoelomate taxa,

Acoela Acoela, or the acoels, is an order of small and simple invertebrates in the subphylum Acoelomorpha of phylum Xenacoelomorpha, a deep branching bilaterian group of animals, which resemble flatworms. Historically they were treated as an order o ...

and

Nemertodermatida Nemertodermatida is a class of Acoelomorpha, comprising 18 species of millimetre-sized turbellariform, mostly interstitial worms. Taxonomy The order ''Nemertodermatida'' contains two families with 6 genera. The high level of cryptic diversity ...

(together called

Acoelomorpha Acoelomorpha is a subphylum of very simple and small soft-bodied animals with planula-like features which live in marine or brackish waters. They usually live between grains of sediment, swimming as plankton, or crawling on other organisms, suc ...

), and the remaining bilaterians. The latter clade was called

Nephrozoa Nephrozoa is a major clade of bilaterians, divided into the protostomes and the deuterostomes, containing almost all animal phyla and over a million extant species. Its sister clade is the Xenacoelomorpha. The Ambulacraria (conventionally deut ...

by Jondelius et al. (2002) and

Eubilateria Nephrozoa is a major clade of bilaterians, divided into the protostomes and the deuterostomes, containing almost all animal phyla and over a million extant species. Its Sister group, sister clade is the Xenacoelomorpha. The Ambulacraria (conventi ...

by Baguña and Riutort (2004). The acoelomorph taxa had previously been considered flatworms with secondarily lost characteristics, but the new relationship suggested that the simple acoelomate worm form was the original bilaterian bodyplan and that the coelom, the digestive tract, excretory organs, and nerve cords developed in the Nephrozoa. Subsequently the acoelomorphs were placed in phylum

Xenacoelomorpha Xenacoelomorpha is a small phylum of bilaterian invertebrate animals, consisting of two sister groups: xenoturbellids and acoelomorphs. This new phylum was named in February 2011 and suggested based on morphological synapomorphies (physical ...

, together with the

xenoturbellid ''Xenoturbella'' is a genus of very simple bilaterians up to a few centimeters long. It contains a small number of marine benthic worm-like species. The first known species (''Xenoturbella bocki'') was discovered in 1915 by Sixten Bock, but it ...

s, and the sister relationship between Xenacoelomorpha and Nephrozoa confirmed in phylogenomic analyses. A modern consensus

phylogenetic tree A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...

for Bilateria is shown below, although the positions of certain

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

s are still controversial (dashed lines) and the tree has changed considerably since 2000. A different hypothesis is that the Ambulacraria are sister to Xenacoelomorpha together forming the

Xenambulacraria Xenambulacraria is a proposed clade of animals with bilateral symmetry as an embryo, consisting of the Xenacoelomorpha (i.e., ''Xenoturbella'' and acoelomorphs) and the Ambulacraria (i.e., echinoderms and hemichordates). If confirmed, the cla ...

. The Xenambulacraria may be sister to the Chordata or the Nephrozoa (sans Ambulacraria). The phylogenetic tree shown below depicts the latter proposal. Also the veracity of deuterostomes (without protostomia emerging in it), is under discussion. It is indicated when approximately clades radiated into newer clades in millions of years ago (Mya). While the below tree depicts a

chordates A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These five ...

as a

sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...

protostomia Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryonic development. This nature has since been discovered to be extremely variable among Protostomia's mem ...

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."

Notes

References

External links

Tree of Life web project — Bilateria

{{Authority control Subkingdoms Ediacaran first appearances Taxa named by Berthold Hatschek