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 being 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 ball. Although the name "millipede" derives from the Latin for "thousand feet", no known species has 1,000; the record of 750 legs belongs to Illacme plenipes. 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.
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.
Harpaphe haydeniana, commonly known as the yellow-spotted millipede, almond-scented millipede or cyanide millipede, is a species of polydesmidan ("flat-backed") millipede found in the moist forests along the Pacific coast of North America, from Southeast Alaska to California. The dark coloration with contrasting yellow-tipped keels warn of its ability to exude toxic hydrogen cyanide as a defense. Despite the various common names given the species, the coloration pattern, cyanide defense, and associated almond scent occur in other flat-backed millipedes around the world.
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).
Paradoxosomatidae, the only family in the suborder Paradoxosomatidea, is a family of flat-backed millipedes in the order Polydesmida. Containing nearly 200 genera and 975 species as of 2013, it is one of the largest families of millipedes. Paradoxosomatids occur on all continents except Antarctica, and can generally be distinguished by dorsal grooves on most body segments and a dumb-bell shaped gonopod aperture. Notable groups within the Paradoxosomatidae include the dragon millipedes of Southeast Asia, and the widely introduced Greenhouse Millipede Oxidus gracilis.
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.
Siphonophorida is an order of millipedes containing two families and over 100 species. This order includes the millipede with the most legs, Illacme plenipes.
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.
Dalodesmidae is a family of millipedes in the order Polydesmida, containing at least 250 species found in the Southern Hemisphere.
Colobognatha is a clade of helminthomorph millipedes containing four orders: Platydesmida, Polyzoniida, Siphonocryptida, and Siphonophorida.
Sinocallipus is a genus of predominantly cave-dwelling millipedes in the order Callipodida. Five of the six known species are found in limestone caves on the Indochinese Peninsula from Vietnam to southern China, and it is the only callipodidan genus entirely confined to the tropics. Individuals range from 40–70 mm (1.6–2.8 in) long and possess 55 to 70 segments. Sinocallipus is the sole taxon of the callipodidan suborder Sinocallipodidea, and thought to be the most primitive and sister group to all other callipodidans. In contrast to many other millipede groups, the gonopods of Sinocallipus show little variation between species, while other characters such as color, size, and other body parts differ more noticeably between species.
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.
Hiltonius is a genus of cylindrical millipedes in the family Spirobolidae comprising 10 species ranging from the southwestern United States to Guatemala, with most species being found in Mexico. The genus was named by Ralph Vary Chamberlin in 1918, after Professor William A. Hilton of Pomona College who collected the type specimen of H. pulchrus.
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.
Cowiedesmus is an extinct millipede genus known from the middle Silurian of Scotland, and is one of the earliest known land animals. 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.
Juliformia is a taxonomic superorder of millipedes containing three living orders: Julida, Spirobolida, and Spirostreptida, and the extinct group Xyloiuloidea known only from fossils. The species possess long cylindrical bodies with sclerites fused into complete rings. Juliform millipedes possess defensive repugnatorial glands on all body segments except the last few, and are the only known millipedes to produce quinones in their defensive secretions. Juliform males have two pairs of gonopods consisting of the modified 8th and 9th pair of legs: in Julida and Spirobolida the posterior gonopods are primarily involved in sperm-transferring, while in Spirostreptida it is the anterior gonopods. Juliformians also lack Tömösváry organs and have a large collum which overhangs the rear of the head.
