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Ecdysozoa () is a group of protostome animals, including Arthropoda ( insects, chelicerata, crustaceans, and myriapods), Nematoda, and several smaller
phyla Phyla, the plural of ''phylum'', may refer to: * Phylum, a biological taxon between Kingdom and Class * by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another Phyl ...
. They were first defined by Aguinaldo ''et al.'' in 1997, based mainly on
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 ...
s constructed using 18S ribosomal RNA genes. A large study in 2008 by Dunn ''et al.'' strongly supported the Ecdysozoa as a
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, ...
, that is, a group consisting of a common ancestor and all its descendants. The group is also supported by morphological characters, and includes all animals that grow by
ecdysis Ecdysis is the moulting of the cuticle in many invertebrates of the clade Ecdysozoa. Since the cuticle of these animals typically forms a largely inelastic exoskeleton, it is shed during growth and a new, larger covering is formed. The remna ...
, moulting their
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
. The group was initially contested by a significant minority of biologists. Some argued for groupings based on more traditional taxonomic techniques, while others contested the interpretation of the molecular data.


Etymology

The name ''Ecdysozoa'' stems etymologically .


Characteristics

The most notable characteristic shared by ecdysozoans is a three-layered
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
(four in Tardigrada) composed of organic material, which is periodically molted as the animal grows. This process of molting is called
ecdysis Ecdysis is the moulting of the cuticle in many invertebrates of the clade Ecdysozoa. Since the cuticle of these animals typically forms a largely inelastic exoskeleton, it is shed during growth and a new, larger covering is formed. The remna ...
, and gives the group its name. The ecdysozoans lack locomotory
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 produce mostly amoeboid sperm, and their
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 ...
s do not undergo spiral cleavage as in most other protostomes. Ancestrally, the group exhibited sclerotized teeth within the foregut, and a ring of spines around the mouth opening, though these features have been secondarily lost in certain groups. A respiratory and circulatory system is only present in onychophorans and arthropods (often absent in smaller arthropods like mites); in the rest of the groups, both systems are missing.


Phylogeny

The Ecdysozoa include the following phyla: Arthropoda, Onychophora, Tardigrada, Kinorhyncha, Priapulida, Loricifera, Nematoda, and Nematomorpha. A few other groups, such as the gastrotrichs, have been considered possible members but lack the main characters of the group, and are now placed elsewhere. The Arthropoda, Onychophora, and Tardigrada have been grouped together as the Panarthropoda because they are distinguished by segmented body plans. Dunn ''et al.'' in 2008 suggested that the tardigrada could be grouped along with the nematodes, leaving Onychophora as the sister group to the arthropods. The non-panarthropod members of Ecdysozoa have been grouped as
Cycloneuralia Cycloneuralia is a clade of ecdysozoan animals including the Scalidophora (Kinorhynchans, Loriciferans, Priapulids) and the Nematoida (nematodes, Nematomorphs). It may be paraphyletic, or may be a sister group to Panarthropoda. Or perhaps Pana ...
but they are more usually considered
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
in representing the primitive condition from which the Panarthropoda evolved. 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. It is indicated when approximately clades radiated into newer clades in millions of years ago (Mya); dashed lines show especially uncertain placements. The phylogenetic tree is based on Nielsen ''et.al.'' with provisional placement of Loricifera from Hiroshi ''et al.''


Older alternative groupings


Articulata hypothesis

The grouping proposed by Aguinaldo ''et al.'' is almost universally accepted, replacing an older hypothesis that Panarthropoda should be classified with Annelida in a group called the Articulata, and that Ecdysozoa are polyphyletic. Nielsen has suggested that a possible solution is to regard Ecdysozoa as a sister-group of Annelida, though later considered them unrelated. Inclusion of the roundworms within the Ecdysozoa was initially contested but since 2003, a broad consensus has formed supporting the Ecdysozoa and in 2011 the
Darwin–Wallace Medal The Darwin–Wallace Medal is a medal awarded by the Linnean Society of London for "major advances in evolutionary biology". Historically, the medals have been awarded every 50 years, beginning in 1908. That year marked 50 years after the joint p ...
was awarded to James Lake for the discovery of the New Animal Phylogeny consisting of the Ecdysozoa, the
Lophotrochozoa Lophotrochozoa (, "crest/wheel animals") is a clade of protostome animals within the Spiralia. The taxon was established as a monophyletic group based on molecular evidence. The clade includes animals like annelids, molluscs, bryozoans, brachi ...
, and the Deuterostomia.


Coelomata hypothesis

Before Aguinaldo's Ecdysozoa proposal, one of the prevailing theories for the evolution of the
bilateral animals 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 ...
was based on the morphology of their body cavities. There were three types, or
grades Grade most commonly refers to: * Grade (education), a measurement of a student's performance * Grade, the number of the year a student has reached in a given educational stage * Grade (slope), the steepness of a slope Grade or grading may also r ...
of organization: the
Acoelomata 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 ...
(no
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 ...
), the
Pseudocoelomata The Aschelminthes (also known as Aeschelminthes, Nemathelminthes, Nematodes), closely associated with the Platyhelminthes, are an obsolete phylum of pseudocoelomate and other similar animals that are no longer considered closely related and have be ...
(partial coelom), and the Eucoelomata (true
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 ...
). Adoutte and coworkers were among the first to strongly support the Ecdysozoa. With the introduction of molecular phylogenetics, the coelomate hypothesis was abandoned, although some molecular, phylogenetic support for the Coelomata continued until as late as 2005.


References


External links


UCMP-Ecdysozoa introduction
* https://web.archive.org/web/20030313115747/http://www.palaeos.com/Kingdoms/Animalia/Ecdysozoa.html * https://web.archive.org/web/20080618211750/http://www.nematodes.org/tardigrades/Tardigrades_and_Ecdysozoa.html * https://web.archive.org/web/20100510081324/http://chuma.cas.usf.edu/~garey/articulata.html * https://web.archive.org/web/20030313065540/http://chuma.cas.usf.edu/~garey/essential.html * http://www.nematomorpha.net {{Taxonbar, from=Q5176 Superphyla Extant Cambrian first appearances