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.
Chilognatha is a subclass of the class Diplopoda, which includes the vast majority of extant millipedes, about 12,000 species.
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.
Arthropleuridea is an extinct subclass of myriapod arthropods that flourished during the Carboniferous period, having first arose during the Silurian, and perishing in the Early Permian. Members are characterized by possessing diplosegement paranotal tergal lobes separated from the body axis by a suture, and by sclerotized plates buttressing the leg insertions. Despite their unique features, recent phylogenetic research suggests Arthropleuridea be included among millipedes in the class Diplopoda. The subclass contains three recognized orders, each with a single genus.
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.
Mammamia profuga is a species of cave-dwelling millipede in the family Julidae. The only known species of the genus Mammamia, it was described in 2011 from a specimen discovered in a cave in Italy.
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.
Xyloiuloidea is an extinct superfamily of millipedes that existed from the Lower Devonian through the Upper Pennsylvanian period in Europe and North America.
Callipodida is an order of millipedes containing around 130 species, many characterized by crests or ridges.
Colobognatha is a clade of helminthomorph millipedes containing four orders: Platydesmida, Polyzoniida, Siphonocryptida, and Siphonophorida.
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.
Floridobolus is a genus of millipedes commonly known as Florida scrub millipedes containing three described species: Floridobolus penneri, F. orini, and F. floydi; the latter two described in 2014. All three species are endemic to Florida scrub habitat of peninsular Florida, and F. penneri is considered a critically imperiled species by NatureServe. Prior to the description of F. orini, the genus was considered the sole member of the family Floridobolidae, named by William T. Keeton in 1959, however studies in 2014 have argued that Floridobolus does not represent a distinct family but rather is a basal member of the family Spirobolidae, representing the subfamily Floridobolinae, and tribe Floridobolini.
Paeromopus paniculus is a species of millipede endemic to the Sierra Nevada mountains in the United States state of California. Reaching up to 16.5 centimeters in length, it is the longest known millipede in North America.
Cowiedesmus is an extinct millipede genus originally described from the middle Silurian of Scotland, and is considered as earliest known land animals alongside Pneumodesmus from same formation. Although a 2017 study suggested that the geological formation it contains actually appears to be from the Early Devonian (Lochkovian) instead, a 2023 study confirmed the age identification of the 2004 study through palynological, palaeobotanical and ziron analyses incorporating newly discovered additional data. Cowiedesmus was about 4 cm (1.6 in) long and characterized by a greatly enlarged pair of legs on the 8th segment which may have been used in clasping females or functioned as gonopods. Coweiedesmus is distinct enough from other living and fossil millipedes to be placed in its own order, Cowiedesmida. The only known species, C. eroticopodus, was described in 2004.
The Archidesmida is an extinct order of millipedes known from fossils from the Devonian period of Europe and North America. Archidesmidans have broad flat keels (paranota) extending from their body segments, and a modified pair of legs on the 8th segment that may have been involved in mating, similar to the gonopods of living millipedes which insert sperm into females. Alternately, the modified legs may have been used to grasp onto partners during mating.
Titanophyllum spiliarum is a species of cave-dwelling millipede in the family Julidae. The only known species of the genus Titanophyllum, it was described in 2011 from specimens discovered in a cave in Greece. It has several unusual characteristics including eyelessness and a small hook on its hind-most body section that may be involved in keeping the animal ‘locked’ when it coils-up defensively.
Simplified cross section (A) and side view (B) of a juliform millipede segment
Male Ommatoiulus moreleti (Julida, Julidae) showing juliform characteristics: large collum, two pair of gonopods, and fused, cylindrical body segments. The hook-shaped first leg pair is unique to Julida.
