Protostomia () is the

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, ...

of animals once thought to be characterized by the formation of the organism's mouth before its anus during

embryonic development 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 sperm ...

. This nature has since been discovered to be extremely variable among Protostomia's members, although the reverse is typically true of its sister clade,

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 ...

. Well known examples of protostomes are

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 ...

mollusc 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 esti ...

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 ...

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

nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...

s. They are also called schizocoelomates since

schizocoely Schizocoely (adjective forms: schizocoelous or schizocoelic) is a process by which some animal embryos develop. The schizocoely mechanism occurs when secondary body cavities ( coeloms) are formed by splitting a solid mass of mesodermal embryonic ...

typically occurs in them. Together with the Deuterostomia and

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 ...

, these form the clade

Bilateria The Bilateria or bilaterians are animals with bilateral symmetry 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 ...

, animals 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 ...

, anteroposterior axis and

three 3 is a number, numeral, and glyph. 3, three, or III may also refer to: * AD 3, the third year of the AD era * 3 BC, the third year before the AD era * March, the third month Books * '' Three of Them'' (Russian: ', literally, "three"), a 1901 ...

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 ...

Protostomy

In animals at least as complex as

earthworms An earthworm is a terrestrial invertebrate that belongs to the phylum Annelida. They exhibit a tube-within-a-tube body plan; they are externally segmented with corresponding internal segmentation; and they usually have setae on all segments. Th ...

, the first phase in gut development involves 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 ...

forming a dent on one side (the blastopore) which deepens to become its digestive tube (the

archenteron The primary gut that forms during gastrulation in a developing embryo is known as the archenteron, the gastrocoel or the primitive digestive tube. It develops into the endoderm and mesoderm of an animal. Formation in sea urchins As primary mesen ...

). In the sister-clade, the deuterostomes (), the original dent becomes the

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 ...

while the gut eventually tunnels through to make another opening, which forms the

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 ...

. The protostomes (from Greek 'first' + 'mouth') were so named because it was once believed that in all cases the embryological dent formed the mouth while the anus was formed later, at the opening made by the other end of the gut. It is now known that the fate of the blastopore among protostomes is extremely variable; while the evolutionary distinction between deuterostomes and protostomes remains valid, the descriptive accuracy of the name ''protostome'' is disputable. Protostome and deuterostome embryos differ in several other ways. Many protostomes (the

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 ...

clade) undergo spiral cleavage during cell division instead of radial cleavage. Spiral cleavage happens because the cells' division planes are angled to the polar major axis, instead of being parallel or perpendicular to it. Another difference is that secondary body cavities (

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 ...

s) generally form by

, where the coelom forms out of a solid mass of embryonic tissue splitting away from the rest, instead of by enterocoelic pouching, where the coelom would otherwise form out of in-folded gut walls.

Evolution

The common ancestor of protostomes and deuterostomes was evidently a worm-like aquatic animal of the

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 ...

. The two clades diverged about 600 million years ago. Protostomes evolved into over a million species alive today, compared to about 60,000 deuterostome species. Protostomes are divided into the Ecdysozoa (e.g.

arthropods 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 ...

nematodes The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a broa ...

) and the

(e.g.

molluscs 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 taxon, extant species of molluscs are recognized. The number of fossil sp ...

annelids 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 ecolo ...

platyhelminths 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 ...

, and

rotifers The rotifers (, from the Latin , "wheel", and , "bearing"), commonly called wheel animals or wheel animalcules, make up a phylum (Rotifera ) of microscopic and near-microscopic pseudocoelomate animals. They were first described by Rev. John H ...

). 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 the protostomes is shown below. The timing of clades radiating into newer clades is given in mya (millions of years ago); less certain placements are indicated with dashed lines.

References

External links

* {{Authority control Ediacaran first appearances Infrakingdoms Taxa named by Karl Grobben

Protostomy

Evolution

See also

References

External links