The subphylum Hexapoda (from
Greek
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Anything of, from, or related to Greece, a country in Southern Europe:
*Greeks, an ethnic group.
*Greek language, a branch of the Indo-European language family.
**Proto-Greek language, the assumed last common ancestor ...
for 'six legs') comprises most species of
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 and includes the
insect
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 ...
s as well as three much smaller groups of wingless arthropods:
Collembola
Springtails (Collembola) form the largest of the three lineages of modern hexapods that are no longer considered insects (the other two are the Protura and Diplura). Although the three orders are sometimes grouped together in a class called Ento ...
,
Protura
The Protura, or proturans, and sometimes nicknamed coneheads, are very small (0.6-1.5mm long), soil-dwelling animals, so inconspicuous they were not noticed until the 20th century. The Protura constitute an order of hexapods that were previously ...
, and
Diplura
The order Diplura ("two-pronged bristletails") is one of three orders of non-insect hexapods within the class Entognatha (alongside Collembola (springtails) and Protura). The name "diplura", or "two tails", refers to the characteristic pair of ...
(all of these were once considered insects). The Collembola (or springtails) are very abundant in terrestrial environments. ''Hexapods'' are named for their most distinctive feature: a consolidated thorax with three pairs of legs (six legs). Most other arthropods have more than three pairs of legs. Most recent studies have recovered Hexapoda as a subgroup of
Crustacea.
Morphology
Hexapods have bodies ranging in length from 0.5 mm to over 300 mm which are divided into an anterior head,
thorax
The thorax or chest is a part of the anatomy of humans, 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 divisions of the cre ...
, and posterior abdomen. The
head
A head is the part of an organism which usually includes the ears, brain, forehead, cheeks, chin, eyes, nose, and mouth, each of which aid in various sensory functions such as sight, hearing, smell, and taste. Some very simple animals may ...
is composed of a presegmental ''acron'' that usually bears eyes (absent in Protura and Diplura), followed by six segments, all closely fused together, with the following appendages:
:Segment I. None
:Segment II.
Antennae (sensory), absent in
Protura
The Protura, or proturans, and sometimes nicknamed coneheads, are very small (0.6-1.5mm long), soil-dwelling animals, so inconspicuous they were not noticed until the 20th century. The Protura constitute an order of hexapods that were previously ...
:Segment III. None
:Segment IV.
Mandible
In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
s (crushing jaws)
:Segment V.
Maxillae
The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
(chewing jaws)
:Segment VI.
Labium (lower lip)
The mouth lies between the fourth and fifth segments and is covered by a projection from the sixth, called the
labrum (upper lip). In true insects (class Insecta) the mouthparts are exposed or
ectognathous, while in other groups they are enveloped or
endognathous. Similar appendages are found on the heads of
Myriapoda
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 ...
and
Crustacea, although these have secondary
antennae.
The thorax is composed of three segments, each of which bears a single pair of legs. As is typical of arthropods adapted to life on land, each leg has only a single walking branch composed of five segments, without the gill branches found in some other arthropods and with gill on the abdominal segments of some immature aquatic insects. In most insects the second and third thoracic segments also support wings. It has been suggested that these may be homologous to the gill branches of crustaceans, or they may have developed from extensions of the segments themselves.
The abdomen follow epimorphic development, where all segments are already present at the end of embryonic development in all the hexapod groups except for Protura, which has an anamorphic development where the hatched juveniles has an incomplete complement of segments, and goes through a post-embryonic segment addition with each molting before the final adult number of segments is reached. All true insects have eleven segments (often reduced in number in many insect species), but in Protura there are twelve, and in
Collembola
Springtails (Collembola) form the largest of the three lineages of modern hexapods that are no longer considered insects (the other two are the Protura and Diplura). Although the three orders are sometimes grouped together in a class called Ento ...
only six (sometimes reduced to only four). The appendages on the abdomen are extremely reduced, restricted to the external genitalia and sometimes a pair of sensory
cerci on the last segment.
Evolution and relationships
The
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, ...
have traditionally been considered the closest relatives of the hexapods, based on morphological similarity. These were then considered subclasses of a subphylum called
Uniramia
Uniramia (''uni'' – one, ''ramus'' – branch, i.e. single-branches) is a group within the arthropods. In the past this group included the Onychophora, which are now considered a separate category. The group is currently used in a narrower sen ...
or Atelocerata. In the first decade of the 21st century, however, this was called into question, and it appears the hexapods' closest relatives may be the
crustacean
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 ...
s.
The non-insect hexapods have variously been considered a single evolutionary line, typically treated as Class
Entognatha
The Entognatha are a class of wingless and ametabolous arthropods, which, together with the insects, makes up the subphylum Hexapoda. Their mouthparts are entognathous, meaning that they are retracted within the head, unlike the insects. Entog ...
, or as several lines with different relationships with the Class Insecta. In particular, the Diplura may be more closely related to the Insecta than to the
Collembola
Springtails (Collembola) form the largest of the three lineages of modern hexapods that are no longer considered insects (the other two are the Protura and Diplura). Although the three orders are sometimes grouped together in a class called Ento ...
(springtails) or the
Protura
The Protura, or proturans, and sometimes nicknamed coneheads, are very small (0.6-1.5mm long), soil-dwelling animals, so inconspicuous they were not noticed until the 20th century. The Protura constitute an order of hexapods that were previously ...
. There is also some evidence suggesting that the hexapod groups may not share a common origin, and in particular that the Collembola belong elsewhere.
Molecular analysis suggests that the hexapods diverged from their sister group, the
Anostraca
Anostraca is one of the four orders of crustaceans in the class Branchiopoda; its members are referred to as fairy shrimp. They live in vernal pools and hypersaline lakes across the world, and they have even been found in deserts, ice-covered m ...
(fairy shrimps), at around the start of the
Silurian
The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleozo ...
period , coinciding with the appearance of
vascular plants
Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They al ...
on land.
The
cladogram
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
below follows the work of Bernhard Misof et al. (2014)
and shows the relationships between the extant orders of Hexapoda:
The following
cladogram
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
is given by Kjer et al. (2016):
An incomplete possible insect fossil, ''
Strudiella devonica,'' has been recovered from the
Devonian
The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
period. This fossil may help to fill the
arthropod gap from 385 million to 325 million years ago, although some researchers oppose this view and suggest that the fossil may instead represent a decomposed crustacean or other non-insect.
References
External links
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{{Taxonbar, from=Q105146
Animal subphyla
Extant Early Devonian first appearances
Pancrustacea