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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 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 ...
. 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. 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 ...
s, molluscs,
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 ...
s, flatworms and nematodes. They are also called schizocoelomates since schizocoely typically occurs in them. Together with the Deuterostomia and Xenacoelomorpha, these form the clade Bilateria, animals with bilateral symmetry, anteroposterior axis and three germ layers.


Protostomy

In animals at least as complex as earthworms, the first phase in
gut Gut or guts may refer to: Anatomy * Abdomen or belly, the region of a vertebrate between the chest and pelvis * Abdominal obesity or "a gut", a large deposit of belly fat * Gastrointestinal tract or gut, the system of digestive organs * Ins ...
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 sperm ...
forming a dent on one side (the blastopore) which deepens to become its digestive tube (the archenteron). In the sister-clade, the deuterostomes (), the original dent becomes the anus 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 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. Many protostomes (the Spiralia 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 ( coeloms) generally form by schizocoely, 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 t ...
. 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, nematodes) and the Spiralia (e.g. molluscs, annelids, platyhelminths, and rotifers). A modern consensus phylogenetic tree for the protostomes is shown below. The timing of clades radiating into newer clades is given in
mya Mya may refer to: Brands and product names * Mya (program), an intelligent personal assistant created by Motorola * Mya (TV channel), an Italian Television channel * Midwest Young Artists, a comprehensive youth music program Codes * Burmese ...
(millions of years ago); less certain placements are indicated with dashed lines.


See also

* Embryological origins of the mouth and anus
The Taxonomicon
for
Karl Grobben Karl Grobben (27 August 1854, in Brno – 13 April 1945, in Salzburg) was an Austrian biologist. He graduated from, and later worked at, the University of Vienna, chiefly on molluscs and crustaceans. He was also the editor of a new edition of Car ...


References


External links

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