| Colobognatha | |
|---|---|
 | |
| Octoglena sierra (Polyzoniida) | |
Scientific classification  | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Arthropoda |
| Subphylum: | Myriapoda |
| Class: | Diplopoda |
| Subclass: | Chilognatha |
| Infraclass: | Helminthomorpha |
| Subterclass: | Colobognatha Brandt, 1834 |
| Orders | |
Colobognatha is a clade (formally considered a subterclass) of helminthomorph millipedes containing four orders: Platydesmida, Polyzoniida, Siphonocryptida, and Siphonophorida. [1]
The Colobognatha are united by several shared traits (synapomorphies). Males have two pairs of simple, leg-like gonopods, consisting of the rear leg pair of body segment 7 and the foreword-most leg pair of segment 8. They possess tubular defensive glands that open on the sides of the body, and lack a palp-like structure on their gnathochilaria. Other traits found in colobognathans, but not exclusively, include a distinctly narrow head, lack of Tömösváry organs, and no more than two pairs of ocelli. [2] [3]
Millipedes are a group of arthropods that are characterised by having two pairs of jointed legs on most body segments; they are known scientifically as the class Diplopoda, the name derived from this feature. Each double-legged segment is a result of two single segments fused together. Most millipedes have very elongated cylindrical or flattened bodies with more than 20 segments, while pill millipedes are shorter and can roll into a tight ball. Although the name "millipede" derives from Latin for "thousand feet", no species was known to have 1,000 or more until the discovery in 2020 of Eumillipes persephone, which can have over 1,300 legs. There are approximately 12,000 named species classified into 16 orders and around 140 families, making Diplopoda the largest class of myriapods, an arthropod group which also includes centipedes and other multi-legged creatures.
Pill millipedes are any members of two living orders of millipedes, often grouped together into a single superorder, Oniscomorpha. The name Oniscomorpha refers to the millipedes' resemblance to certain woodlice (Oniscidea), also called pillbugs or "roly-polies". However, millipedes and woodlice are not closely related ; rather, this is a case of convergent evolution.
Gonopods are specialized appendages of various arthropods used in reproduction or egg-laying. In males, they facilitate the transfer of sperm from male to female during mating, and thus are a type of intromittent organ. In crustaceans and millipedes, gonopods are modified walking or swimming legs. Gonopods may be highly decorated with elaborate structures which may play roles in sperm competition, and can be used to differentiate and identify closely related species. Gonopods generally occur in one or more pairs, as opposed to the single (un-paired) reproductive organs such as the aedeagus of insects or the penis of harvestmen.
Spirobolida is an order of "round-backed" millipedes containing approximately 500 species in 12 families. Its members are distinguished by the presence of a "pronounced suture that runs "vertically down the front of the head". Most of the species live in the tropics, and many are brightly coloured. Mature males have two pairs of modified legs, the gonopods, consisting of the 8th and 9th leg pair: the posterior gonopods are used in sperm-transfer while the anterior gonopods are fused into a single plate-like structure.
Glomerida is an order of pill-millipedes found primarily in the Northern Hemisphere. Also known as northern pill millipedes, they superficially resemble pill-bugs or woodlice, and can enroll into a protective ball. They have twelve body segments, 17 to 19 pairs of legs, and males have enlarged rear legs involved in mating. The order includes about 30 genera and at least 280 species, including Glomeris marginata, the common European pill-millipede. The order contains members in Europe, South-east Asia and the Americas from California to Guatemala. Although historically considered closely related with the similar sphaerotheriidans that also enroll, some DNA evidence suggest they may be more closely related to glomeridesmidans, a poorly known order that does not enroll.
Spirostreptida is an order of long, cylindrical millipedes. There are approximately 1000 described species, making Spirostreptida the third largest order of millipedes after Polydesmida and Chordeumatida.
Polydesmida is the largest order of millipedes, containing approximately 3,500 species, including all the millipedes reported to produce hydrogen cyanide (HCN). Polydesmids grow and develop through a series of moults, adding segments until they reach a fixed number in the adult stage, which is usually the same for a given sex in a given species, at which point the moulting and the addition of segments and legs stop. This mode of development, known as teloanamorphosis, distinguishes this order from most other orders of millipedes, which usually continue to moult as adults, developing through either euanamorphosis or hemianamorphosis.
Julida is an order of millipedes. Members are mostly small and cylindrical, typically ranging from 10–120 millimetres (0.39–4.72 in) in length. Eyes may be present or absent, and in mature males of many species, the first pair of legs is modified into hook-like structures. Additionally, both pairs of legs on the 7th body segment of males are modified into gonopods.
Polyxenida is an order of millipedes readily distinguished by a unique body plan consisting of a soft, non-calcified body ornamented with tufts of bristles – traits that have inspired the common names "bristly millipedes" or "pincushion millipedes". There are at least 86 species in four families worldwide, and are the only living members of the subclass Penicillata.
Polyzoniida is an order of millipedes in the subterclass Colobognatha containing three families and at least 74 described species.
Platydesmida is an order of millipedes containing two families and over 60 species. Some species practice paternal care, in which males guard the eggs.
Siphonophorida is an order of millipedes containing two families and over 100 species.
Callipodida is an order of millipedes containing around 130 species, many characterized by crests or ridges.
Pentazonia is a taxonomic infraclass of millipedes containing the pill-millipedes (Oniscomorpha) which can roll into a ball and the order Glomeridesmida which cannot. Defining traits (apomorphies) include divided sternites, a labrum with single median tooth, and an enlarged pygidium on the hind-most body segment. Pentazonia is in the dominant millipede subclass Chilognatha which have a calcified exoskeleton and modified sperm-transferring legs in males. In contrast to the Helminthomorpha – the other Chilognathan infraclass, the sperm-transferring legs are located on posterior body segments and known as telopods. Pentazonians are relatively short-bodied, with between 13 and 21 body segments. The Pentazonia contains one extinct order, Amynilyspedida, often referred to the Oniscomorpha.
Chordeumatida is a large order of millipedes containing some 1200 species with a nearly worldwide distribution. Also known as sausage millipedes, they grow and develop through a series of moults, adding segments until they reach a fixed number in the adult stage, which is usually the same for a given sex in a given species, at which point the moulting and the addition of segments and legs stop. This mode of development, known as teloanamorphosis, distinguishes this order from most other orders of millipedes, which usually continue to moult as adults, developing through either euanamorphosis or hemianamorphosis.
Siphoniulus is a poorly known genus of millipede containing only two living species: S. alba from Indonesia, and S. neotropicus from Mexico and Guatemala. An additional two fossil species are known from Cretaceous amber. Siphoniulus species are the only members of the family Siphoniulidae and order Siphoniulida, making Siphoniulida the smallest millipede order. Few specimens are known, and their classification is contentious, although most recent studies place them as basal members of the Helminthomorpha.
Stemmiulida is an order of millipedes consisting of approximately 130 species, reaching up to 50 mm in length. It contains a single family, Stemmiulidae.
Juliformia is a taxonomic superorder of millipedes containing three living orders: Julida, Spirobolida, and Spirostreptida, and the extinct group Xyloiuloidea known only from fossils.
Pyrgodesmidae is a family of flat-backed millipedes in the order Polydesmida. The family contains over 200 species distributed in tropics around the world. Some species are found only in ant colonies, and are considered obligate myrmecophiles.
Lophoproctidae is a family of millipedes in the order Polyxenida containing approximately 65 species in 6 genera.
Orders of the class Diplopoda (millipedes) | |||||||||||||||||||||||||||||||
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