Tagmosis
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biology Biology is the scientific study of life and living organisms. It is a broad natural science that encompasses a wide range of fields and unifying principles that explain the structure, function, growth, History of life, origin, evolution, and ...
, a tagma (Greek: τάγμα, : tagmata – τάγματα - body of soldiers; battalion) is a specialized grouping of multiple segments or metameres into a coherently functional morphological unit. Familiar examples are the head, the
thorax The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of mammals and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main di ...
, and the
abdomen The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
of insects. The segments within a tagma may be either fused (such as in the head of an insect) or so jointed as to be independently moveable (such as in the abdomen of most insects). Usually the term is taken to refer to tagmata in the morphology of members of the
phylum In biology, a phylum (; : phyla) is a level of classification, or taxonomic rank, that is below Kingdom (biology), kingdom and above Class (biology), class. Traditionally, in botany the term division (taxonomy), division has been used instead ...
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 ...
a, but it applies equally validly in other phyla, such as the
Chordata A chordate ( ) is a bilaterian animal belonging to the phylum Chordata ( ). All chordates possess, at some point during their larval or adult stages, five distinctive physical characteristics (Apomorphy and synapomorphy, synapomorphies) th ...
. In a given taxon the names assigned to particular tagmata are in some sense informal and arbitrary; for example, not all the tagmata of species within a given
subphylum In zoological nomenclature, a subphylum is a taxonomic rank below the rank of phylum. The taxonomic rank of " subdivision" in fungi and plant taxonomy is equivalent to "subphylum" in zoological taxonomy. Some plant taxonomists have also used th ...
of the Arthropoda are homologous to those of species in other subphyla; for one thing they do not all comprise corresponding
somite The somites (outdated term: primitive segments) are a set of bilaterally paired blocks of paraxial mesoderm that form in the embryogenesis, embryonic stage of somitogenesis, along the head-to-tail axis in segmentation (biology), segmented animals. ...
s, and for another, not all the tagmata have closely analogous functions or anatomy. In some cases this has led to earlier names for tagmata being more or less successfully superseded. For example, the one-time terms "
cephalothorax The cephalothorax, also called prosoma in some groups, is a tagma of various arthropods, comprising the head and the thorax fused together, as distinct from the abdomen behind. (The terms ''prosoma'' and ''opisthosoma'' are equivalent to ''cepha ...
" and "
abdomen The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
" of the Araneae, though not yet strictly regarded as invalid, are giving way to prosoma and
opisthosoma The opisthosoma is the posterior part of the body in some arthropods, behind the prosoma ( cephalothorax). It is a distinctive feature of the subphylum Chelicerata (arachnids, horseshoe crabs and others). Although it is similar in most respects ...
. The latter two terms carry less of a suggestion of homology with the significantly different tagmata of insects.


Tagmosis

The development of distinct tagmata is believed to be a feature of the evolution of segmented animals, especially
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 ...
s. In the ancestral arthropod, the body was made up of repeated segments, each with similar internal organs and appendages. One evolutionary trend is the grouping together of some segments into larger units, the tagmata. The evolutionary process of grouping is called tagmosis (or tagmatization). The first and simplest stage was a division into two tagmata: an anterior "head" (cephalon) and a posterior "trunk". The head contained the brain and carried sensory and feeding appendages. The trunk bore the appendages responsible for locomotion and respiration (
gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow r ...
s in aquatic species). In almost all modern arthropods, the trunk is further divided into a "thorax" and an "abdomen", with the thorax bearing the main locomotory appendages. In some groups, such as
arachnid Arachnids are arthropods in the Class (biology), class Arachnida () of the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, opiliones, harvestmen, Solifugae, camel spiders, Amblypygi, wh ...
s, the cephalon (head) and thorax are hardly distinct externally and form a single tagma, the "cephalothorax" or "prosoma".
Mite Mites are small arachnids (eight-legged arthropods) of two large orders, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari. However, most recent genetic analyses do not recover the two as eac ...
s appear to have a single tagma with no obvious external signs of either segments or separate tagmata. Tagmosis proceeded differently in different groups of arthropods, so that the tagmata are not derived from corresponding (homologous) segments, even though the same names may be used for the tagmata.
Copepod Copepods (; meaning 'oar-feet') are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (living in the water column), some are benthos, benthic (living on the sedimen ...
s (a kind of
crustacean Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...
) provide an example. The basic copepod body consists of a head, a thorax with six segments, ancestrally each with a swimming leg, and an abdomen with five appendageless segments. Except in parasitic species, the body is divided functionally into two tagmata, that may be called a "prosome" and a "urosome", with an articulation between them allowing the body to flex. Different groups of copepods have the articulation at different places. In the Calanoida, the articulation is between the thoracic and abdominal segments, so that the boundary between the prosome and urosome corresponds to the boundary between thoracic and abdominal segments. However, in the Harpacticoida, the articulation is between the fifth and sixth thoracic segments, so that the sixth thoracic segment is in the urosome (see the diagram). Tagmosis is an extreme form of heteronomy, mediated by ''Hox'' genes and the other developmental genes they influence.


Terminology

Trilobite sections-en.svg, The tagmata of a
trilobite Trilobites (; meaning "three-lobed entities") are extinction, extinct marine arthropods that form the class (biology), class Trilobita. One of the earliest groups of arthropods to appear in the fossil record, trilobites were among the most succ ...
: cephalon, thorax and pygidium File:Tagma.png, Tagmata of an insect: head (cephalon), thorax and abdomen Tagmata and major appendages of spider IMG 7639A.JPG, Tagmata and major appendages of a
spider Spiders (order (biology), order Araneae) are air-breathing arthropods that have eight limbs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude spider silk, silk. They are the largest order of arachnids and ran ...
: cephalothorax or prosoma and abdomen or opisthosoma
The number of tagma and their names vary among
taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
. For example, the extinct
trilobite Trilobites (; meaning "three-lobed entities") are extinction, extinct marine arthropods that form the class (biology), class Trilobita. One of the earliest groups of arthropods to appear in the fossil record, trilobites were among the most succ ...
s had three tagmata: the cephalon (meaning head), the thorax (literally meaning chest, but in this application referring to the mid-portion of the body), and the pygidium (meaning rump). The
Hexapoda The subphylum Hexapoda (from Greek for 'six legs') or hexapods comprises the largest clade of arthropods and includes most of the extant arthropod species. It includes the crown group class Insecta (true insects), as well as the much smaller c ...
, including insects, also have three tagmata, usually termed the head, thorax, and abdomen. The bodies of many arachnids, such as
spider Spiders (order (biology), order Araneae) are air-breathing arthropods that have eight limbs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude spider silk, silk. They are the largest order of arachnids and ran ...
s, have two tagmata, as do the bodies of some
crustacean Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...
s: in both groups the anterior tagma may be called the
cephalothorax The cephalothorax, also called prosoma in some groups, is a tagma of various arthropods, comprising the head and the thorax fused together, as distinct from the abdomen behind. (The terms ''prosoma'' and ''opisthosoma'' are equivalent to ''cepha ...
(meaning head plus chest) or the prosoma or prosome (meaning "fore-part of body"). The posterior tagma may be called the abdomen. In those arachnids that have two tagmata, the abdomen is also called the
opisthosoma The opisthosoma is the posterior part of the body in some arthropods, behind the prosoma ( cephalothorax). It is a distinctive feature of the subphylum Chelicerata (arachnids, horseshoe crabs and others). Although it is similar in most respects ...
. In
crustacea Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...
ns, the posterior tagma is also called the pleon or the urosome (meaning the tail part); alternatively, "pleon" may refer only to the abdominal segments incorporated into the posterior tagma, the thoracic segments in this tagma being called the "pereon".


See also

* Arthropod head problem


References


Bibliography

* *{{Cite book , last1=Ruppert , first1=E.E. , last2=Fox , first2=R.S. , last3=Barnes , first3=R.D. , year=2004 , title=Invertebrate Zoology , edition=7th , publisher=Brooks/Cole , isbn=978-0-03-025982-1 , name-list-style=amp , url-access=registration , url=https://archive.org/details/isbn_9780030259821 Arthropod anatomy