Ecdysozoa () is a group of
protostome
Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryogenesis, embryonic development. This nature has since been discovered to be extremely variable among Pro ...
animals,
including
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 ...
a (
insects
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of j ...
,
chelicerata
The subphylum Chelicerata (from New Latin, , ) constitutes one of the major subdivisions of the phylum Arthropoda. It contains the sea spiders, horseshoe crabs, and arachnids (including Opiliones, harvestmen, scorpions, spiders, Solifugae, solif ...
,
crustaceans
Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
, and
myriapods
Myriapods () are the members of subphylum Myriapoda, containing arthropods such as millipedes and centipedes. The group contains about 13,000 species, all of them terrestrial.
The fossil record of myriapods reaches back into the late Silurian, a ...
),
Nematoda
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 several smaller
phyla. 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
Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from ribosomal ...
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
Tardigrades (), known colloquially as water bears or moss piglets, are a phylum of eight-legged segmented micro-animals. They were first described by the German zoologist Johann August Ephraim Goeze in 1773, who called them Kleiner Wasserbä ...
) 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
An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and retracting pseudopo ...
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
In embryology, cleavage is the division of cells in the early development of the embryo, following fertilization. The zygotes of many species undergo rapid cell cycles with no significant overall growth, producing a cluster of cells the same size ...
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
onychophora
Onychophora (from grc, ονυχής, , "claws"; and , , "to carry"), commonly known as velvet worms (due to their velvety texture and somewhat wormlike appearance) or more ambiguously as peripatus (after the first described genus, '' Peripatus ...
ns 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
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 ...
,
Onychophora
Onychophora (from grc, ονυχής, , "claws"; and , , "to carry"), commonly known as velvet worms (due to their velvety texture and somewhat wormlike appearance) or more ambiguously as peripatus (after the first described genus, '' Peripatus ...
,
Tardigrada
Tardigrades (), known colloquially as water bears or moss piglets, are a phylum of eight-legged segmented micro-animals. They were first described by the German zoologist Johann August Ephraim Goeze in 1773, who called them Kleiner Wasserbä ...
,
Kinorhyncha
Kinorhyncha ( grc, κινέω, kīnéō, I move, ' "snout") is a phylum of small marine invertebrates that are widespread in mud or sand at all depths as part of the meiobenthos. They are also called mud dragons. Modern species are or less, ...
,
Priapulida
Priapulida (priapulid worms, from Gr. πριάπος, ''priāpos'' 'Priapus' + Lat. ''-ul-'', diminutive), sometimes referred to as penis worms, is a phylum of unsegmented marine worms. The name of the phylum relates to the Greek god of fertility ...
,
Loricifera
Loricifera (from Latin, '' lorica'', corselet (armour) + ''ferre'', to bear) is a phylum of very small to microscopic marine cycloneuralian sediment-dwelling animals that had been determined to be 37 described species, in 9 genera, bu ...
,
Nematoda
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
Nematomorpha
Nematomorpha (sometimes called Gordiacea, and commonly known as horsehair worms, hairsnakes, or Gordian worms) are a phylum of parasitoid animals superficially similar to nematode worms in morphology, hence the name. Most species range in size fr ...
. A few other groups, such as the
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 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
Panarthropoda is a proposed animal clade containing the extant phyla Arthropoda, Tardigrada (water bears) and Onychophora (velvet worms). Panarthropods also include extinct marine legged worms known as lobopodians ("Lobopodia"), a paraphylet ...
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 P ...
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
Panarthropoda is a proposed animal clade containing the extant phyla Arthropoda, Tardigrada (water bears) and Onychophora (velvet worms). Panarthropods also include extinct marine legged worms known as lobopodians ("Lobopodia"), a paraphylet ...
should be classified with
Annelida in a group called the
Articulata, and that Ecdysozoa are
polyphyletic
A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of converg ...
. 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
roundworm
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 bro ...
s 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
James A. Lake (born August 10, 1941, Kearney, Nebraska) is an American evolutionary biologist and a Distinguished Professor of Molecular, Cell, and Developmental Biology and of Human Genetics at UCLA. Lake is best known for the New Animal Phyloge ...
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 was based on the morphology of their
body cavities
A body cavity is any space or compartment, or potential space, in an animal body. Cavities accommodate organs and other structures; cavities as potential spaces contain fluid.
The two largest human body cavities are the ventral body cavity, and ...
. 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 (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
The coelom (or celom) is the main body cavity in most animals and is positioned inside the Body (biology), body to surround and contain the digestive tract and other Organ (biology), organs. In some animals, it is lined with mesothelium. In other ...
(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