Protostomia () is the

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

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

embryonic development In developmental biology, animal embryonic development, also known as animal embryogenesis, is the developmental stage of an animal embryo. Embryonic development starts with the fertilization of an egg cell (ovum) by a sperm, sperm cell (spermat ...

. 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 Deuterostomes (from Ancient Greek, Greek: ) are bilaterian animals of the superphylum Deuterostomia (), typically characterized by their anus forming before the mouth during embryogenesis, embryonic development. Deuterostomia comprises three Phyl ...

. Well-known examples of protostomes are

arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...

mollusc Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...

annelid The annelids (), also known as the segmented worms, are animals that comprise the phylum Annelida (; ). The phylum contains over 22,000 extant species, including ragworms, earthworms, and leeches. The species exist in and have adapted to vario ...

flatworm Platyhelminthes (from the Greek language, Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") is a Phylum (biology), phylum of relatively simple bilaterian, Segmentation (biology), ...

s and

nematode The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (h ...

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 ti ...

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 (physica ...

, these form the clade

Bilateria Bilateria () is a large clade of animals characterised by bilateral symmetry during embryonic development. This means their body plans are laid around a longitudinal axis with a front (or "head") and a rear (or "tail") end, as well as a left� ...

, 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, the face of a human being has a plane of symme ...

, anteroposterior axis and three

germ layer A germ layer is a primary layer of cell (biology), 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 tw ...

Protostomy

In animals at least as complex as

earthworms An earthworm is a soil-dwelling terrestrial animal, terrestrial invertebrate that belongs to the phylum Annelida. The term is the common name for the largest members of the class (biology), class (or subclass (biology), subclass, depending on ...

, the first phase in gut development involves the

embryo An embryo ( ) is the initial stage of development for 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 sp ...

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

archenteron The archenteron, also called the gastrocoel, is the internal cavity formed in the gastrulation stage in early embryonic development that becomes the cavity of the primitive gut. Formation in sea urchins As primary mesenchyme cells detach fro ...

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

anus In mammals, invertebrates and most fish, the anus (: anuses or ani; from Latin, 'ring' or 'circle') is the external body orifice at the ''exit'' end of the digestive tract (bowel), i.e. the opposite end from the mouth. Its function is to facil ...

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

mouth A mouth also referred to as the oral is the body orifice through which many animals ingest food and animal communication#Auditory, vocalize. The body cavity immediately behind the mouth opening, known as the oral cavity (or in Latin), is also t ...

. 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. Secondary body cavities (

coelom The coelom (or celom) is the main body cavity in many 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, i ...

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 ( ) is a geological period of the Neoproterozoic geologic era, Era that spans 96 million years from the end of the Cryogenian Period at 635 Million years ago, Mya to the beginning of the Cambrian Period at 538.8 Mya. It is the last ...

. The two clades diverged about 600 million years ago. Protostomes evolved into over a million species alive today, compared to ca. 73,000 deuterostome species. Protostomes are divided into the

Ecdysozoa Ecdysozoa () is a group of protostome animals, including Arthropoda (insects, chelicerates (including arachnids), crustaceans, and myriapods), Nematoda, and several smaller phylum (biology), phyla. The grouping of these animal phyla into a single ...

(e.g.

arthropods Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...

nematodes The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (he ...

) and the

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

(e.g.

molluscs Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...

annelids The annelids (), also known as the segmented worms, are animals that comprise the phylum Annelida (; ). The phylum contains over 22,000 extant species, including ragworms, earthworms, and leeches. The species exist in and have adapted to vario ...

, platyhelminths, and

rotifers The rotifers (, from Latin 'wheel' and 'bearing'), sometimes 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 Harris ...

). A modern consensus

phylogenetic tree A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA. In ...

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

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

Protostomy

Evolution

See also

References

External links