The subphylum Hexapoda (from

Greek Greek may refer to: Greece 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 segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...

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 ...

s as well as three much smaller groups of wingless arthropods: Collembola,

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 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

: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 (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 ...

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 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 can ...

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) or the

. 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 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 ...

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 ...

below follows the work of Bernhard Misof et al. (2014) and shows the relationships between the extant orders of Hexapoda: The following

is given by Kjer et al. (2016): An incomplete possible insect fossil, '' Strudiella devonica,'' has been recovered from the Devonian 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

* * * {{Taxonbar, from=Q105146 Animal subphyla Extant Early Devonian first appearances Pancrustacea