Мыңаяқтылар: Түрлер, қазбалар және эволюциясы

Class of arthropods

Millipedes (originating from the Latin mille, "thousand", and pes, "foot") 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. Other vernacular names include "thousand legger" or simply "diplopod".

[[File:Millipede order species comparison. png|thumb|left|alt=Diversity|Approximate relative diversity of extant millipede orders, ranging from ca. 3,500 species of Polydesmida to 2 species of Siphoniulida Few species of millipede are at all widespread; they have very poor dispersal abilities, depending as they do on terrestrial locomotion and humid habitats. These factors have favoured genetic isolation and rapid speciation, producing many lineages with restricted ranges. The living members of the Diplopoda are divided into sixteen orders in two subclasses. The basal subclass Penicillata contains a single order, Polyxenida (bristle millipedes). All other millipedes belong to the subclass Chilognatha consisting of two infraclasses: Pentazonia, containing the short bodied pill millipedes, and Helminthomorpha (worm like millipedes), containing the great majority of the species.

Millipedes are among the first animals to have colonised land during the Silurian period. Early forms probably ate mosses and primitive vascular plants. There are two major groups of millipedes whose members are all extinct: the Archipolypoda ("ancient, many legged ones") which contain the oldest known terrestrial animals, and Arthropleuridea, which contain the largest known land invertebrates. Pneumodesmus newmani is the earliest member of the millipedes from the late Wenlock epoch of the late Silurian around , Other early fossils of millipedes are Kampecaris obanensis and Archidesmus sp. from 425 millions years ago in the late Silurian. During the Upper Carboniferous , Arthropleura became the largest known land dwelling invertebrate on record, reaching lengths of at least Millipedes also exhibit the earliest evidence of chemical defence, as some Devonian fossils have defensive gland openings called ozopores.

The diplosegments of millipedes have evolved in conjunction with their burrowing habits, and nearly all millipedes adopt a mainly subterranean lifestyle. They use three main methods of burrowing; bulldozing, wedging and boring. Members of the orders Julida, Spirobolida and Spirostreptida, lower their heads and barge their way into the substrate, the collum leading the way. Flat backed millipedes in the order Polydesmida tend to insert their front end, like a wedge, into a horizontal crevice, and then widen the crack by pushing upwards with their legs, the paranota in this instance constituting the main lifting surface. Boring is used by members of the order Polyzoniida. These have smaller segments at the front and increasingly large ones further back; they propel themselves forward into a crack with their legs, the wedge shaped body widening the gap as they go. Some millipedes have adopted an above ground lifestyle and lost the burrowing habit. This may be because they are too small to have enough leverage to burrow, or because they are too large to make the effort worthwhile, or in some cases because they move relatively fast (for a millipede) and are active predators. feeding on insects, centipedes, earthworms, or snails. Some species have piercing mouth parts that allow them to suck up plant juices.

Due to their lack of speed and their inability to bite or sting, millipedes' primary defence mechanism is to curl into a tight coil – protecting their delicate legs inside an armoured exoskeleton. Many species also emit various foul smelling liquid secretions through microscopic holes called ozopores (the openings of "odoriferous" or "repugnatorial glands"), along the sides of their bodies as a secondary defence. Among the many irritant and toxic chemicals found in these secretions are alkaloids, benzoquinones, phenols, terpenoids, and hydrogen cyanide. Some of these substances are caustic and can burn the exoskeleton of ants and other insect predators, and the skin and eyes of larger predators. Primates such as capuchin monkeys and lemurs have been observed intentionally irritating millipedes in order to rub the chemicals on themselves to repel mosquitoes. Some of these defensive compounds also show antifungal activity. The bristly millipedes (order Polyxenida) lack both an armoured exoskeleton and odiferous glands, and instead are covered in numerous bristles that in at least one species, Polyxenus fasciculatus, detach and entangle ants.