Wikipedia

Millipede

For other uses, seeMillipede (disambiguation).

Millipedes(originating from theLatinmille,"thousand", andpes,"foot" )[1][2]are a group ofarthropodsthat are characterised by having two pairs of jointedlegson most body segments; they are known scientifically as theclassDiplopoda,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, whilepill millipedesare shorter and can roll into a tight ball. Although the name "millipede" derives fromLatinfor "thousand feet", no species was known to have 1,000 or more until the discovery in 2020 ofEumillipes persephone,which can have over 1,300 legs.[3]There are approximately 12,000 namedspeciesclassified into 16ordersand around 140families,making Diplopoda the largest class ofmyriapods,an arthropod group which also includescentipedesand other multi-legged creatures.

Millipedes
Temporal range:428–0MaLateSiluriantoPresent
An assortment of millipedes (not to scale)
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Myriapoda
Class: Diplopoda
BlainvilleinGervais,1844
Subclasses
Diversity
16 orders, c. 12,000 species

Most millipedes are slow-movingdetritivores,eating decaying leaves and other dead plant matter; however, some eatfungior drink plant fluid. Millipedes are generally harmless to humans, although some can become household or gardenpests.Millipedes can be an unwanted nuisance particularly ingreenhouseswhere they can potentially cause severe damage to emergent seedlings. Most millipedes defend themselves with a variety of chemicals secreted from pores along the body, although the tinybristle millipedesare covered with tufts of detachable bristles. Its primary defence mechanism is to curl into a tight coil, thereby protecting its legs and other vital delicate areas on the body behind a hard exoskeleton.Reproductionin most species is carried out by modified male legs calledgonopods,which transfer packets of sperm to females.

First appearing in theSilurianperiod, millipedes are some of the oldest knownland animals.Some members of prehistoric groups, such asArthropleura,grew to over2 m (6+1⁄2ft); the largest modern species reach maximum lengths of27 to 38 cm (10+1⁄2to 15 in). The longest extant species is the giant African millipede (Archispirostreptus gigas).

Among myriapods, millipedes have traditionally been considered most closely related to the tinypauropods,although somemolecular studieschallenge this relationship. Millipedes canbe distinguishedfrom the somewhat similar but only distantly relatedcentipedes(class Chilopoda), which move rapidly, arevenomous,carnivorous,and have only a single pair of legs on each body segment.

The scientific study of millipedes is known as diplopodology, and a scientist who studies them is called a diplopodologist.

Etymology and names

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The term "millipede" is widespread in popular and scientific literature, but among North American scientists, the term "milliped" (without the terminal e) is also used.[4]Other vernacular names include "thousand-legger" or simply "diplopod".[5]The science of millipede biology and taxonomy is called diplopodology: the study of diplopods.[6]

Classification

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Approximate relative diversity ofextantmillipede orders, ranging from ca. 3,500 species ofPolydesmidato 2 species ofSiphoniulida[7]

Approximately 12,000 millipedespecieshave been described. Estimates of the true number of species on earth range from 15,000[8]to as high as 80,000.[9]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 rapidspeciation,producing many lineages with restricted ranges.[10]

The living members of the Diplopoda are divided into sixteen orders in two subclasses.[7]The basal subclass Penicillata contains a single order, Polyxenida (bristle millipedes).[11]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.[12][13]

Outline of classification

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Further information:List of millipede families

The higher-level classification of millipedes is presented below, based on Shear, 2011,[7]and Shear & Edgecombe, 2010[14](extinct groups). Recent cladistic and molecular studies have challenged the traditional classification schemes above, and in particular the position of the orders Siphoniulida and Polyzoniida is not yet well established.[9]The placement and positions of extinct groups (†) known only from fossils is tentative and not fully resolved.[9][14]After each name is listed theauthor citation:the name of the person who coined the name or defined the group, even if not at the current rank.

Class Diplopodade Blainville in Gervais, 1844

Evolution

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Millipedes are among the first animals to havecolonised landduring theSilurian period.[16]Early forms probably atemossesand primitivevascular plants.There are two major groups of millipedes whose members are all extinct: theArchipolypoda( "ancient, many-legged ones" ) which contain the oldest known terrestrial animals, andArthropleuridea,which contain the largest known land invertebrates.Pneumodesmus newmaniis the earliest member of the millipedes from the lateWenlock epochof the lateSilurianaround428million years ago,[17][18]or earlyLochkovianof the earlyDevonianaround 414 million years ago,[19][20]known from1 cm (1⁄2in) long fragment and has clear evidence ofspiracles(breathing holes) attesting to its air-breathing habits.[14][18][21]Other early fossils of millipedes areKampecaris obanensisandArchidesmussp. from 425 millions years ago in the lateSilurian.[22]During the UpperCarboniferous(340 to 280million years ago),Arthropleurabecame the largest known land-dwelling invertebrate on record, reaching lengths of at least2 m (6+1⁄2ft).[23]Millipedes also exhibit the earliest evidence of chemical defence, as someDevonianfossils have defensive gland openings calledozopores.[14]Millipedes, centipedes, and other terrestrial arthropods attained very large sizes in comparison to modern species in the oxygen-rich environments of the Devonian and Carboniferous periods, and some could grow larger than one metre. As oxygen levels lowered through time, arthropods became smaller.[24]

Living groups

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Octoglena sierra(Colobognatha,Polyzoniida)
Anadenobolus monilicornis(Juliformia,Spirobolida)
Harpaphe haydeniana(Polydesmida)

The history of scientific millipede classification began withCarl Linnaeus,who in his10th edition ofSystema Naturae,1758, named seven species ofJulusas "Insecta Aptera" (wingless insects).[25]In 1802, the French zoologistPierre André Latreilleproposed the name Chilognatha as the first group of what are now the Diplopoda, and in 1840 the German naturalistJohann Friedrich von Brandtproduced the first detailed classification. The name Diplopoda itself was coined in 1844 by the French zoologistHenri Marie Ducrotay de Blainville.From 1890 to 1940, millipede taxonomy was driven by relatively few researchers at any given time, with major contributions byCarl Attems,Karl Wilhelm VerhoeffandRalph Vary Chamberlin,who each described over 1,000 species, as well asOrator F. Cook,Filippo Silvestri,R. I. Pocock,andHenry W. Brölemann.[9]This was a period when the science of diplopodology flourished: rates of species descriptions were on average the highest in history, sometimes exceeding 300 per year.[8]

In 1971, the Dutch biologistC. A. W. Jeekelpublished a comprehensive listing of all known millipede genera and families described between 1758 and 1957 in hisNomenclator Generum et Familiarum Diplopodorum,a work credited as launching the "modern era" of millipede taxonomy.[26][27]In 1980, the American biologistRichard L. Hoffmanpublished a classification of millipedes which recognized the Penicillata, Pentazonia, and Helminthomorpha,[28]and the first phylogenetic analysis of millipede orders using moderncladistic methodswas published in 1984 by Henrik Enghoff of Denmark.[29]A 2003 classification by the American myriapodologist Rowland Shelley is similar to the one originally proposed by Verhoeff, and remains the currently accepted classification scheme (shown below), despite more recent molecular studies proposing conflicting relationships.[9][14]A 2011 summary of millipede family diversity byWilliam A. Shearplaced the orderSiphoniulidawithin the larger group Nematophora.[7]

Diplopoda

Fossil record

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In addition to the 16 living orders, there are 9 extinct orders and one superfamily known only from fossils. The relationship of these to living groups and to each other is controversial. The extinct Arthropleuridea was long considered a distinct myriapod class, although work in the early 21st century established the group as a subclass of millipedes.[30][31][32]Several living orders also appear in the fossil record. Below are two proposed arrangements of fossil millipede groups.[9][14]Extinct groups are indicated with adagger(†). The extinct orderZosterogrammida,a chilognath of uncertain position,[14]is not shown.

Alternate hypothesis of fossil relationships[9][31]
Diplopoda

Relation to other myriapods

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Pauropods are thought to be the closest relative of millipedes.

Although the relationships of millipede orders are still the subject of debate, the class Diplopoda as a whole is considered amonophyletic groupof arthropods: all millipedes are more closely related to each other than to any other arthropods. Diplopoda is a class within the arthropod subphylumMyriapoda,the myriapods, which includescentipedes(class Chilopoda) as well as the lesser-knownpauropods(class Pauropoda) andsymphylans(class Symphyla). Within myriapods, the closest relatives orsister groupof millipedes has long been considered the pauropods, which also have a collum and diplosegments.[9]

Distinction from centipedes

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The differences between millipedes and centipedes are a common question from the general public.[33]Both groups of myriapods share similarities, such as long, multi-segmented bodies, many legs, a single pair of antennae, and the presence ofpostantennal organs,but have many differences and distinct evolutionary histories, as themost recent common ancestorof centipedes and millipedes lived around 450 to 475 million years ago in the Silurian.[34]The head alone exemplifies the differences; millipedes have short, geniculate (elbowed)antennaefor probing the substrate, a pair of robust mandibles and a single pair of maxillae fused into a lip; centipedes have long, threadlike antennae, a pair of small mandibles, two pairs of maxillae and a pair of large poison claws.[35]

A representative millipede and centipede (not necessarily to scale)
Millipede and centipede differences[33]
Trait Millipedes Centipedes
Legs Two pairs on most body segments; attached to underside of body One pair per body segment; attached to sides of body; last pair extends backwards
Locomotion Generally adapted for burrowing or inhabiting small crevices; slow-moving Generally adapted for running, except for the burrowingsoil centipedes
Feeding Primarily detritivores, some herbivores, few carnivores; no venom Primarily carnivores with front legs modified into venomous fangs
Spiracles On underside of body On the sides or top of body
Reproductive openings Third body segment Last body segment
Reproductive behaviour Male generally inserts spermatophore into female with gonopods Male produces spermatophore that is usually picked up by female

Characteristics

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Representative body types of the Penicillata (top), Pentazonia (middle), and Helminthomorpha (bottom)
Anterior anatomy of a generalized helminthomorph millipede

Millipedes come in a variety of body shapes and sizes, ranging from2 mm (1⁄16in) to around 35 cm (14 in) in length,[36]and can have as few as eleven to over three hundred segments.[37][38]They are generally black or brown in colour, although there are a few brightly coloured species, and some haveaposematiccolouring to warn that they are toxic.[5]Species ofMotyxiaproducecyanideas a chemical defence and arebioluminescent.[39]

Body styles vary greatly between major millipede groups. In thebasalsubclassPenicillata,consisting of the tinybristle millipedes,the exoskeleton is soft and uncalcified, and is covered in prominentsetaeor bristles. All other millipedes, belonging to the subclass Chilognatha, have a hardened exoskeleton. The chilognaths are in turn divided into two infraclasses: thePentazonia,containing relatively short-bodied groups such aspill millipedes,and the Helminthomorpha ( "worm-like" millipedes), which contains the vast majority of species, with long, many-segmented bodies.[12][13]

They have also lost the gene that codes for theJHAMTlenzyme, which is responsible for catalysing the last step of the production of a juvenile hormone that regulates the development and reproduction in other arthropods like crustaceans, centipedes and insects.[40]

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The head of a millipede is typically rounded above and flattened below and bears a pair of largemandiblesin front of a plate-like structure called agnathochilarium( "jaw lip" ).[9]The head contains a single pair ofantennaewith seven or eight segments and a group of sensory cones at the tip.[9]Many orders also possess a pair of sensory organs known as theTömösváry organs,shaped as small oval rings posterior and lateral to the base of the antennae. Their function is unknown,[9]but they also occur in somecentipedes,and are possibly used to measure humidity or light levels in the surrounding environment.[41]

Millipede eyes consist of several simple flat-lensedocelliarranged in a group or patch on each side of the head. These patches are also called ocular fields or ocellaria. Many species of millipedes, including the entire ordersPolydesmida,Siphoniulida,Glomeridesmida,SiphonophoridaandPlatydesmida,and cave-dwelling millipedes such asCauseyellaandTrichopetalum,had ancestors that could see but have subsequently lost their eyes and are blind.[36]

Body

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Paranota ofpolydesmidan(left) andplatydesmidanmillipedes

Millipede bodies may be flattened or cylindrical, and are composed of numerousmetameric segments,each with anexoskeletonconsisting of fourchitinousplates: a single plate above (thetergite), one at each side (pleurites), and a plate on the underside (sternite) where the legs attach. In many millipedes, such as Merocheta and Juliformia, these plates are fused to varying degrees, sometimes forming a single cylindrical ring. The plates are typically hard, impregnated with calcium salts.[37]Because they can't close their permanently open spiracles and most species lack a waxy cuticle, millipedes are susceptible to water loss and with a few exceptions must spend most of their time in moist or humid environments.[42]

The first segment behind the head is legless and known as a collum (from the Latin for neck or collar). The second, third, and fourth body segments bear a single pair of legs each and are known as "haplosegments" (the three haplosegments are sometimes referred to as a "thorax"[18]). The remaining segments, from the fifth to the posterior, are properly known as diplosegments or double segments, formed by the fusion of two embryonic segments. Each diplosegment bears two pairs of legs, rather than just one as in centipedes. In some millipedes, the last few segments may be legless. The terms "segment" or "body ring" are often used interchangeably to refer to both haplo- and diplosegments. The final segment is known as thetelsonand consists of a legless preanal ring, a pair of anal valves (closeable plates around the anus), and a small scale below the anus.[9][37]

Millipedes in several orders have keel-like extensions of the body-wall known asparanota,which can vary widely in shape, size, and texture; modifications include lobes, papillae, ridges, crests, spines and notches.[5]Paranota may allow millipedes to wedge more securely into crevices, protect the legs, or make the millipede more difficult for predators to swallow.[43]

The legs are composed of seven segments, and attach on the underside of the body. The legs of an individual are generally rather similar to each other, although often longer in males than females, and males of some species may have a reduced or enlarged first pair of legs.[44]The most conspicuous leg modifications are involved in reproduction, discussed below. Despite the common name, no millipede was known to have 1,000 legs until 2021: common species have between 34 and 400 legs, and the record is held byEumillipes persephone,with individuals possessing up to 1,306 legs – more than any other creature on Earth.[3][45][46]

A femaleIllacme plenipeswith 618 legs (309 pairs)

Internal organs

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Millipedes breathe through two pairs of spiracles located ventrally on each segment near the base of the legs.[33]Each opens into an internal pouch, and connects to a system oftracheae.The heart runs the entire length of the body, with anaortastretching into the head. The excretory organs are two pairs ofmalpighian tubules,located near the mid-part of the gut. The digestive tract is a simple tube with two pairs ofsalivary glandsto help digest the food.[37]

Reproduction and growth

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Epibolus pulchripesmating; the male is on the right

Millipedes show a diversity of mating styles and structures. In the basal orderPolyxenida(bristle millipedes), mating is indirect: males depositspermatophoresonto webs they secrete with special glands, and the spermatophores are subsequently picked up by females.[33]In all other millipede groups, males possess one or two pairs of modified legs calledgonopodswhich are used to transfer sperm to the female during copulation. The location of the gonopods differs between groups: in males of thePentazoniathey are located at the rear of the body and known as telopods and may also function in grasping females, while in the Helminthomorpha – the vast majority of species – they are located on the seventh body segment.[9]A few species areparthenogenetic,having few, if any, males.[47]

The gonopods ofNipponesmus shirinensisare quite unlike its walking legs.
Left gonopod ofOxidus gracilis.False colourSEMimage, scale bar: 0.2 mm

Gonopods occur in a diversity of shapes and sizes, and in the range from closely resembling walking legs to complex structures quite unlike legs at all. In some groups, the gonopods are kept retracted within the body; in others they project forward parallel to the body. Gonopod morphology is the predominant means of determining species among millipedes: the structures may differ greatly between closely related species but very little within a species.[48]The gonopods develop gradually from walking legs through successive moults until reproductive maturity.[49]

Growth stages ofNemasoma(Nemasomatidae), which reaches reproductive maturity in stage V

The genital openings (gonopores) of both sexes are located on the underside of the third body segment (near the second pair of legs) and may be accompanied in the male by one or twopeneswhich deposit the sperm packets onto the gonopods. In the female, the genital pores open into paired small sacs calledcyphopodsor vulvae, which are covered by small hood-like lids, and are used to store the sperm after copulation.[37]The cyphopod morphology can also be used to identify species. Millipedespermlackflagella,a unique trait among myriapods.[9]

In all except the bristle millipedes, copulation occurs with the two individuals facing one another. Copulation may be preceded by male behaviours such as tapping with antennae, running along the back of the female, offering edible glandular secretions, or in the case of some pill-millipedes,stridulationor "chirping".[50]During copulation in most millipedes, the male positions his seventh segment in front of the female's third segment, and may insert his gonopods to extrude the vulvae before bending his body to deposit sperm onto his gonopods and reinserting the "charged" gonopods into the female.[44]

Females lay from ten to three hundred eggs at a time, depending on species, fertilising them with the stored sperm as they do so. Many species deposit the eggs on moist soil or organic detritus, but some construct nests lined with driedfaeces,and may protect the eggs within silk cocoons.[37]In most species, the female abandons the eggs after they are laid, but some species in the ordersPlatydesmidaandStemmiulidaprovideparental carefor eggs and young.[33]

The young hatch after a few weeks, and typically have only three pairs of legs, followed by up to four legless segments. As they grow, they continuallymoult,adding further segments and legs as they do so, a mode of development known asanamorphosis.[35]Some species moult within specially prepared chambers of soil or silk,[51]and may also shelter in these during wet weather, and most species eat the discarded exoskeleton after moulting. The adult stage, when individuals become reproductively mature, is generally reached in the final moult stage, which varies between species and orders, although some species continue to moult after adulthood. Furthermore, some species alternate between reproductive and non-reproductive stages after maturity, a phenomenon known as periodomorphosis, in which the reproductive structures regress during non-reproductive stages.[47]Millipedes may live from one to ten years, depending on species.[37]

Ecology

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Habitat and distribution

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Millipedes occur on all continents except Antarctica, and occupy almost all terrestrial habitats, ranging as far north as theArctic Circlein Iceland, Norway, and Central Russia, and as far south asSanta Cruz Province, Argentina.[52][53]Typically forest floor dwellers, they live in leaf litter, dead wood, or soil, with a preference for humid conditions. Intemperate zones,millipedes are most abundant in moist deciduous forests, and may reach densities of over 1,000 individuals per square metre. Other habitats include coniferous forests, caves, and alpine ecosystems.[33][53]Deserticolous millipedes, species evolved to live in the desert, likeOrthoporus ornatus,may show adaptations like a waxy epicuticle and the ability of water uptake from unsaturated air.[54]Some species can survive freshwater floods and live submerged underwater for up to 11 months.[55][56]A few species occur near the seashore and can survive in somewhat salty conditions.[47][57]

Burrowing

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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 ordersJulida,SpirobolidaandSpirostreptida,lower their heads and barge their way into the substrate, the collum leading the way. Flat-backed millipedes in the orderPolydesmidatend 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 orderPolyzoniida.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.[5]

Diet

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Most millipedes aredetritivoresand feed on decomposing vegetation, feces, or organic matter mixed with soil. They often play important roles in the breakdown anddecompositionofplant litter:estimates of consumption rates for individual species range from 1 to 11 percent of all leaf litter, depending on species and region, and collectively millipedes may consume nearly all the leaf litter in a region. The leaf litter is fragmented in the millipede gut and excreted as pellets of leaf fragments, algae, fungi, and bacteria, which facilitates decomposition by the microorganisms.[44]Whereearthwormpopulations are low in tropical forests, millipedes play an important role in facilitating microbial decomposition of the leaf litter.[5]Some millipedes are herbivorous, feeding on living plants, and some species can become serious pests of crops. Millipedes in the orderPolyxenidagraze algae from bark, andPlatydesmidafeed on fungi.[9]A few species areomnivorousor inCallipodidaandChordeumatidaoccasionally carnivorous,[58]feeding on insects, centipedes, earthworms, orsnails.[37][59]Some species have piercing mouth parts that allow them to suck up plant juices.[33]Cave dwelling species inJulidae,Blaniulidae,andPolydesmidaehave specialized mouthparts and appears to be filter feeders, filtering small particles from running water inside caves.[60]

Predators and parasites

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ASceliagesbeetle transporting a millipede carcass

Millipedes arepreyed onby a wide range of animals, including variousreptiles,amphibians,birds,mammals,andinsects.[9]Mammalian predators such ascoatisandmeerkatsroll captured millipedes on the ground to deplete and rub off their defensive secretions before consuming their prey,[61]and certainpoison dart frogsare believed to incorporate the toxic compounds of millipedes into their own defences.[62]Several invertebrates have specialised behaviours or structures to feed on millipedes, including larvalglowworm beetles,[63]Probolomyrmexants,[64]chlamydephorid slugs,[65]and predaceous dung beetles of the generaSceliagesandDeltochilum.[66][67]A large subfamily ofassassin bugs,theEctrichodiinaewith over 600 species, has specialised in preying upon millipedes.[68]Parasitesof millipedes includenematodes,phaeomyiid flies,andacanthocephalans.[9]Nearly 30 fungal species of the orderLaboulbenialeshave been found growing externally on millipedes, but some species may becommensalrather than parasitic.[69]

Defence mechanisms

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Further information:Anti-predator adaptation
Juliformandoniscomorphmillipedes curled in a defensive coil

Due to their lack of speed and their inability to bite or sting, millipedes' primarydefence mechanismis to curl into a tight coil – protecting their delicate legs inside an armoured exoskeleton.[70]

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 arealkaloids,benzoquinones,phenols,terpenoids,andhydrogen cyanide.[71][72][73][74][75]Some of these substances arecausticand can burn the exoskeleton ofantsand other insect predators, and the skin and eyes of larger predators. Primates such ascapuchin monkeysandlemurshave been observed intentionally irritating millipedes in order to rub the chemicals on themselves to repelmosquitoes.[76][77][78]Some of these defensive compounds also show antifungal activity.[79]

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.[80]

Other inter-species interactions

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Psammodesmus bryophoruscamouflaged withsymbioticmosses

Some millipedes formmutualistic relationshipswith organisms of other species, in which both species benefit from the interaction, orcommensal relationships,in which only one species benefits while the other is unaffected. Several species form close relationships with ants, a relationship known asmyrmecophily,especially within the familyPyrgodesmidae(Polydesmida), which contains "obligate myrmecophiles", species which have only been found in ant colonies. More species are "facultative myrmecophiles", non-exclusively associated with ants, including many species of Polyxenida that have been found in ant nests around the world.[81]

Many millipede species have commensal relationships withmitesof the ordersMesostigmataandAstigmata.Many of these mites are believed to bephoreticrather than parasitic, which means that they use the millipede host as a means of dispersal.[82][83]

A novel interaction between millipedes and mosses was described in 2011, in which individuals of the newly discoveredPsammodesmus bryophoruswas found to have up to ten species living on its dorsal surface, in what may providecamouflagefor the millipede and increased dispersal for the mosses.[84][85]

Interactions with humans

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Giant fire millipede (Aphistogoniulus corallipes), Madagascar

Millipedes generally have little impact on human economic or social well-being, especially in comparison with insects, although locally they can be a nuisance or agriculturalpest.Millipedes do not bite, and their defensive secretions are mostly harmless to humans — usually causing only minor discolouration on the skin — but the secretions of some tropical species may cause pain, itching, localerythema,edema,blisters,eczema,and occasionally cracked skin.[86][87][88][89]Eye exposures to these secretions causes general irritation and potentially more severe effects such asconjunctivitisandkeratitis.[90]This is calledmillipede burn.First aidconsists of flushing the area thoroughly with water; further treatment is aimed at relieving the local effects.

Spotted snake millipedes can be agricultural pests.

Some millipedes are considered household pests, includingXenobolus carnifexwhich can infestthatched roofsin India,[91]andOmmatoiulus moreleti,which periodically invades homes in Australia. Other species exhibit periodicalswarming behaviour,which can result in home invasions,[92]crop damage,[93]and train delays when the tracks become slippery with the crushed remains of hundreds of millipedes.[44][94][95]Some millipedes can cause significant damage to crops: thespotted snake millipede(Blaniulus guttulatus) is a pest ofsugar beetsand other root crops, and as a result is one of the few millipedes with acommon name.[47]

Some of the larger millipedes in the orders Spirobolida, Spirostreptida, andSphaerotheriidaare popular as pets.[96]Some species commonly sold or kept include species ofArchispirostreptus,Aphistogoniulus,Narceus,andOrthoporus.[97]

Flat millipede found in the Mount Cameroon Forest

Millipedes appear infolkloreandtraditional medicinearound the world. Some cultures associate millipede activity with coming rains.[98]In Zambia, smashed millipede pulp is used to treat wounds, and theBafia peopleof Cameroon use millipede juice to treat earache.[98]In certain HimalayanBhotiyatribes, dry millipede smoke is used to treathaemorrhoids.[99]Native people inMalaysiause millipede secretions in poison-tipped arrows.[98]The secretions ofSpirobolus bungiihave been observed to inhibit division of human cancer cells.[100]The only recorded usage of millipedes as food by humans comes from theBobo peopleofBurkina FasoinWest Africa,who consume boiled, dried millipedes belonging to the familiesGomphodesmidaeandSpirostreptidae[101]: 341 [102]to which they add tomato sauce.[102]

Millipedes have also inspired and played roles in scientific research. In 1963, a walking vehicle with 36 legs was designed, said to have been inspired by a study of millipede locomotion.[103] Experimental robots have had the same inspiration,[104][105]in particular when heavy loads are needed to be carried in tight areas involving turns and curves.[106]In biology, some authors have advocated millipedes asmodel organismsfor the study of arthropod physiology and the developmental processes controlling the number and shape of body segments.[44]

Similar tovermicompost,millipedes can be used to convert plant matter into compost in what has been named millicomposting, which improves the quality of the compost.[107][108]

References

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  1. ^"Millipede | Etymology of millipede by etymonline".
  2. ^"Millipede".
  3. ^abMarek, Paul E.; Buzatto, Bruno A.; Shear, William A.; Means, Jackson C.; Black, Dennis G.; Harvey, Mark S.; Rodriguez, Juanita (2021)."The first true millipede—1306 legs long".Scientific Reports.11(1): 23126.Bibcode:2021NatSR..1123126M.doi:10.1038/s41598-021-02447-0.PMC8677783.PMID34916527.
  4. ^Hoffman, Richard L.(1990). "Diplopoda". In Dindal, Daniel L. (ed.).Soil Biology Guide.John Wiley & Sons. p. 835.ISBN978-0-471-04551-9.
    Hoffman, Richard L. (2000)."Milliped or Millipede?"(PDF).Bulletin of the British Myriapod Group.16:36–37.Archived(PDF)from the original on 2015-02-21.Retrieved2015-02-21.
  5. ^abcdeRuppert, Edward E.; Fox, Richard, S.; Barnes, Robert D. (2004).Invertebrate Zoology, 7th edition.Cengage Learning. pp. 711–717.ISBN978-81-315-0104-7.{{cite book}}:CS1 maint: multiple names: authors list (link)
  6. ^Cockburn, Harry (2021-12-21)."Millipedes 'as big as cars' once roamed Northern England, fossil discovery reveals".The Independent.Retrieved2022-03-12.
  7. ^abcdShear, W.(2011)."Class Diplopoda de Blainville in Gervais, 1844. In: Zhang, Z.-Q. (Ed.) Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness"(PDF).Zootaxa.3148:159–164.doi:10.11646/zootaxa.3148.1.32.Archived(PDF)from the original on 2019-07-25.Retrieved2013-10-14.
  8. ^abBrewer, Michael S.; Sierwald, Petra; Bond, Jason E. (2012)."Millipede taxonomy after 250 years: Classification and taxonomic practices in a mega-diverse yet understudied arthropod group".PLOS ONE.7(5): e37240.Bibcode:2012PLoSO...737240B.doi:10.1371/journal.pone.0037240.PMC3352885.PMID22615951.
  9. ^abcdefghijklmnopqrSierwald, Petra; Bond, Jason E. (2007). "Current status of the myriapod class Diplopoda (Millipedes): Taxonomic diversity and phylogeny".Annual Review of Entomology.52(1): 401–420.doi:10.1146/annurev.ento.52.111805.090210.PMID17163800.
  10. ^Barker, G.M. (2004).Natural Enemies of Terrestrial Molluscs.CABI. pp. 405–406.ISBN978-0-85199-061-3.
  11. ^"Bristle Millipedes (Subclass Penicillata)".iNaturalist.Retrieved2023-03-07.
  12. ^abBueno-Villegas, Julián; Sierwald, Petra; Bond, Jason E."Diplopoda"(PDF).In Bousquets, J. L.; Morrone, J. J. (eds.).Biodiversidad, taxonomia y biogeografia de artropodos de Mexico.pp. 569–599.Archived(PDF)from the original on 2016-12-20.Retrieved2016-12-18.
  13. ^abShelley, Rowland M."Millipedes".University of Tennessee: Entomology and Plant Pathology.Archivedfrom the original on 16 August 2016.Retrieved17 July2016.
  14. ^abcdefghiShear, William A.;Edgecombe, Gregory D. (2010). "The geological record and phylogeny of the Myriapoda".Arthropod Structure & Development.39(2–3): 174–190.Bibcode:2010ArtSD..39..174S.doi:10.1016/j.asd.2009.11.002.PMID19944188.
  15. ^Hoffman, R. L. (1963). "New genera and species of Upper Paleozoic Diplopoda".Journal of Paleontology.37(1): 167–174.JSTOR1301419.
  16. ^Garwood, Russell; Edgecombe, Gregory (2011)."Early terrestrial animals, evolution and uncertainty".Evolution: Education and Outreach.4(3): 489–501.doi:10.1007/s12052-011-0357-y.
  17. ^Wellman, C.H.; Lopes, G.; McKellar, Z.; Hartley, A. (2023)."Age of the basal 'Lower Old Red Sandstone' Stonehaven Group of Scotland: The oldest reported air-breathing land animal is Silurian (late Wenlock) in age".Journal of the Geological Society.181.The Geological Society of London.doi:10.1144/jgs2023-138.hdl:2164/22754.ISSN0016-7649.
  18. ^abcWilson, Heather M.; Anderson, Lyall I. (2004). "Morphology and taxonomy of Paleozoic millipedes (Diplopoda: Chilognatha: Archipolypoda) from Scotland".Journal of Paleontology.78(1): 169–184.Bibcode:2004JPal...78..169W.doi:10.1666/0022-3360(2004)078<0169:MATOPM>2.0.CO;2.S2CID131201588.
  19. ^Suarez, Stephanie E.; Brookfield, Michael E.; Catlos, Elizabeth J.; Stöckli, Daniel F. (2017-06-28)."A U-Pb zircon age constraint on the oldest-recorded air-breathing land animal".PLOS ONE.12(6): e0179262.Bibcode:2017PLoSO..1279262S.doi:10.1371/journal.pone.0179262.ISSN1932-6203.PMC5489152.PMID28658320.
  20. ^Brookfield, M. E.; Catlos, E. J.; Garza, H. (2024-07-07)."The oldest 'millipede'-plant association? Age, paleoenvironments and sources of the Silurian lake sediments at Kerrera, Argyll and Bute, Scotland".Historical Biology:1–13.doi:10.1080/08912963.2024.2367554.ISSN0891-2963.
  21. ^"Fossil millipede found to be oldest land creature".CNN.Reuters. Jan 27, 2004.Archivedfrom the original on March 4, 2016.RetrievedDecember 24,2021.
  22. ^Brookfield, M. E.; Catlos, E. J.; Suarez, S. E. (2021-10-03)."Myriapod divergence times differ between molecular clock and fossil evidence: U/Pb zircon ages of the earliest fossil millipede-bearing sediments and their significance".Historical Biology.33(10): 2014–2018.Bibcode:2021HBio...33.2014B.doi:10.1080/08912963.2020.1762593.ISSN0891-2963.S2CID238220137.
  23. ^Sues, Hans-Dieter (15 January 2011)."Largest Land-Dwelling" Bug "of All Time".National Geographic.Archived fromthe originalon 4 March 2016.Retrieved25 February2016.
  24. ^Lockley, M. G.; Meyer, Christian (2013)."The Tradition of Tracking Dinosaurs in Europe".Dinosaur Tracks and Other Fossil Footprints of Europe.Columbia University Press.pp. 25–52.ISBN978-0-231-50460-7.Archivedfrom the original on 2020-06-02.Retrieved2015-10-27.
  25. ^Caroli Linnaei (1758).Systema naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis.Vol. v.1. Impensis Direct. Laurentii Salvii. pp. 639–640.Archivedfrom the original on 2018-07-12.Retrieved2018-02-20.
  26. ^Shelley, R. M. (2007)."Taxonomy of extant Diplopoda (Millipeds) in the modern era: Perspectives for future advancements and observations on the global diplopod community (Arthropoda: Diplopoda)"(PDF).Zootaxa.1668:343–362.doi:10.11646/zootaxa.1668.1.18.Archived(PDF)from the original on 2014-12-26.Retrieved2014-11-11.
  27. ^Shelley, Rowland M.; Sierwald, Petra; Kiser, Selena B.; Golovatch, Sergei I. (2000).Nomenclator generum et familiarum Diplopodorum II: a list of the genus and family-group names in the class Diplopoda from 1958 through 1999.Sofia, Bulgaria: Pensoft. p. 5.ISBN978-954-642-107-4.
  28. ^Hoffman, Richard L. (1980).Classification of the Diplopoda.Geneva, Switzerland: Muséum d'Historie Naturelle. pp. 1–237.
  29. ^Enghoff, H. (1984). "Phylogeny of millipedes – a cladistic analysis".Journal of Zoological Systematics and Evolutionary Research.22(1): 8–26.doi:10.1111/j.1439-0469.1984.tb00559.x.
  30. ^Wilson, Heather M.; Shear, William A. (2000). "Microdecemplicida, a new order of minute arthropleurideans (Arthropoda: Myriapoda) from the Devonian of New York State, U.S.A".Transactions of the Royal Society of Edinburgh: Earth Sciences.90(4): 351–375.doi:10.1017/S0263593300002674.S2CID129597005.
  31. ^abKraus, O.; Brauckmann, C. (2003). "Fossil giants and surviving dwarfs. Arthropleurida and Pselaphognatha (Atelocerata, Diplopoda): characters, phylogenetic relationships and construction".Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg.40:5–50.
  32. ^Kraus, O. (2005). "On the structure and biology ofArthropleuraspecies (Atelocerata, Diplopoda; Upper Carboniferous/Lower Permian) ".Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg.41:5–23.
  33. ^abcdefgShelley, Rowland M. (1999)."Centipedes and millipedes with emphasis on North American fauna".The Kansas School Naturalist.45(3): 1–16. Archived fromthe originalon 2016-11-12.Retrieved2013-10-14.
  34. ^Brewer, Michael S.; Bond, Jason E. (2013)."Ordinal-level phylogenomics of the arthropod class Diplopoda (Millipedes) based on an analysis of 221 nuclear protein-coding loci generated using next-generation sequence analyses".PLOS ONE.8(11): e79935.Bibcode:2013PLoSO...879935B.doi:10.1371/journal.pone.0079935.PMC3827447.PMID24236165.
  35. ^abBlower, John Gordon (1985).Millipedes: Keys and Notes for the Identification of the Species.Brill Archive. p. 1.ISBN978-90-04-07698-3.
  36. ^abMinelli, Alessandro; Golovatch, Sergei I. (2001)."Myriapods"(PDF).In Levin, Simon A. (ed.).Encyclopedia of Biodiversity.Academic Press. pp. 291–303.ISBN978-0-12-226865-6.Archived fromthe original(PDF)on 2014-02-21.
  37. ^abcdefghBarnes, Robert D. (1982).Invertebrate Zoology.Philadelphia, PA: Holt-Saunders International. pp. 818–825.ISBN978-0-03-056747-6.
  38. ^Marek, Paul E.; Buzatto, Bruno A.; Shear, William A.; Means, Jackson C.; Black, Dennis G.; Harvey, Mark S.; Rodriguez, Juanita (2021-12-16)."The first true millipede—1306 legs long".Scientific Reports.11(1): 23126.Bibcode:2021NatSR..1123126M.doi:10.1038/s41598-021-02447-0.ISSN2045-2322.PMC8677783.PMID34916527.
  39. ^Marek, Paul E.; Moore, Wendy (2015)."Discovery of a glowing millipede in California and the gradual evolution of bioluminescence in Diplopoda".Proceedings of the National Academy of Sciences.112(20): 6419–6424.Bibcode:2015PNAS..112.6419M.doi:10.1073/pnas.1500014112.PMC4443369.PMID25941389.
  40. ^So, Wai Lok; Nong, Wenyan; Xie, Yichun; Baril, Tobias; Ma, Hai-yao; Qu, Zhe; Haimovitz, Jasmine; Swale, Thomas; Gaitan-Espitia, Juan Diego; Lau, Kwok Fai; Tobe, Stephen S.; Bendena, William G.; Kai, Zhen-Peng; Hayward, Alexander; Hui, Jerome H. L. (2022)."Myriapod genomes reveal ancestral horizontal gene transfer and hormonal gene loss in millipedes".Nature Communications.13(1): 3010.Bibcode:2022NatCo..13.3010S.doi:10.1038/s41467-022-30690-0.PMC9151784.PMID35637228.
  41. ^Lewis, J. G. E. (2008).The Biology of Centipedes(Digitally printed 1st paperback version. ed.). Cambridge:Cambridge University Press.pp. 110–111.ISBN978-0-521-03411-1.
  42. ^Capinera, John L., ed. (2008)."Millipedes".Encyclopedia of Entomology.Springer.pp. 2395–2397.ISBN978-1-4020-6242-1.Archivedfrom the original on 2016-05-15.Retrieved2015-10-27.
  43. ^Mesibov, Robert."Paranota".External Anatomy of Polydesmida.Archived fromthe originalon 4 March 2016.Retrieved30 October2013.
  44. ^abcdeHopkin, Stephen P.; Read, Helen J. (1992).The Biology of Millipedes.Oxford University Press.ISBN978-0-19-857699-0.
  45. ^Lu, Donna (16 December 2021)."The first true millipede: new species with more than 1,000 legs discovered in Western Australia".The Guardian.Archivedfrom the original on 16 December 2021.Retrieved16 December2021.
  46. ^Marek, P.; Shear, W.; Bond, J. (2012)."A redescription of the leggiest animal, the millipedeIllacme plenipes,with notes on its natural history and biogeography (Diplopoda, Siphonophorida, Siphonorhinidae) ".ZooKeys(241): 77–112.Bibcode:2012ZooK..241...77M.doi:10.3897/zookeys.241.3831.PMC3559107.PMID23372415.
  47. ^abcdBlower, J. Gordon (1985).Millipedes: Keys and Notes for the Identification of the Species.London: Published for the Linnean Society of London and the Estuarine and Brackish-Water Sciences Association byE.J. Brill.ISBN978-90-04-07698-3.
  48. ^Mesibov, Robert."Gonopods".External Anatomy of Polydesmida.Archived fromthe originalon 9 July 2017.Retrieved27 October2013.
  49. ^Drago, Leandro; Fusco, Giuseppe; Garollo, Elena; Minelli, Alessandro (2011)."Structural aspects of leg-to-gonopod metamorphosis in male helminthomorph millipedes (Diplopoda)".Frontiers in Zoology.8(1): 19.doi:10.1186/1742-9994-8-19.PMC3170261.PMID21859471.
  50. ^Wesener, Thomas; Köhler, Jörn; Fuchs, Stefan; van den Spiegel, Didier (2011). "How to uncoil your partner—" mating songs "in giant pill-millipedes (Diplopoda: Sphaerotheriida)".Naturwissenschaften.98(11): 967–975.Bibcode:2011NW.....98..967W.doi:10.1007/s00114-011-0850-8.PMID21971844.S2CID12005617.
  51. ^Enghoff, Henrik; Akkari, Nesrine (2011)."A callipodidan cocoon (Diplopoda, Callipodida, Schizopetalidae)".International Journal of Myriapodology.5:49–53.doi:10.3897/ijm.5.1995.
  52. ^Shelley, Rowland M.; Golavatch, Sergei I. (2011). "Atlas of myriapod biogeography. I. Indigenous ordinal and supra-ordinal distributions in the Diplopoda: Perspectives on taxon origins and ages, and a hypothesis on the origin and early evolution of the class".Insecta Mundi.158:1–134.
  53. ^abGolovatch, Sergei I.; Kime, R. Desmond (2009)."Millipede (Diplopoda) distributions: a review"(PDF).Soil Organisms.81(3): 565–597. Archived fromthe original(PDF)on 2016-03-03.Retrieved2014-11-19.
  54. ^"Millipede (Diplopoda) distributions: A review"(PDF).Archived(PDF)from the original on 2020-10-08.Retrieved2020-10-06.
  55. ^Adis, Joachim (1986). "An 'aquatic' millipede from a Central Amazonian inundation forest".Oecologia.68(3): 347–349.Bibcode:1986Oecol..68..347A.doi:10.1007/BF01036737.PMID28311777.S2CID11374324.
  56. ^Burrows, F. J.; Hales, D. F.; Beattie, A. J. (1994). "Aquatic millipedes in Australia: a biological Enigma and a conservation saga".Australian Zoologist.29(3–4): 213–216.doi:10.7882/az.1994.007.
  57. ^Barber, A. D., ed. (2013)."World Database of Littoral Myriapoda".World Register of Marine Species.Archivedfrom the original on 7 September 2013.Retrieved25 October2013.
  58. ^Minelli, Alessandro (September 29, 2015).Treatise on Zoology - Anatomy, Taxonomy, Biology. The Myriapoda, Volume 2.BRILL.ISBN9789004188273.Archivedfrom the original on October 9, 2020.RetrievedOctober 6,2020– via Google Books.
  59. ^Barker, G. M. (2004)."Millipedes (Diplopoda) and Centipedes (Chilopoda) (Myriapoda) as predators of terrestrial gastropods".In Barker, G. M. (ed.).Natural Enemies of Terrestrial Molluscs.CAB International.pp. 405–426.ISBN978-0-85199-061-3.
  60. ^Worldwide distribution of cave-dwelling Chelodesmidae (Diplopoda, Polydesmida)
  61. ^Weldon, Paul J.; Cranmore, Catherine F.; Chatfield, Jenifer A. (2006). "Prey-rolling behavior of coatis (Nasuaspp.) is elicited by benzoquinones from millipedes ".Naturwissenschaften.93(1): 14–16.Bibcode:2006NW.....93...14W.doi:10.1007/s00114-005-0064-z.PMID16391932.S2CID22200949.
  62. ^Saporito, R. A.; Donnelly, M. A.; Hoffman, R. L.; Garraffo, H. M.; Daly, J. W. (2003)."A siphonotid millipede (Rhinotus) as the source of spiropyrrolizidine oximes of dendrobatid frogs ".Journal of Chemical Ecology.29(12): 2781–2786.Bibcode:2003JCEco..29.2781S.doi:10.1023/B:JOEC.0000008065.28364.a0.PMID14969363.S2CID4094895.Archivedfrom the original on 2021-03-08.Retrieved2019-07-07.
  63. ^Eisner, T.; Eisner, M.; Attygalle, A. B.; Deyrup, M.; Meinwald, J. (1998)."Rendering the inedible edible: circumvention of a millipede's chemical defence by a predaceous beetle larva".Proceedings of the National Academy of Sciences of the United States of America.95(3): 1108–13.Bibcode:1998PNAS...95.1108E.doi:10.1073/pnas.95.3.1108.PMC18689.PMID9448293.
  64. ^Ito, F. (1998)."Colony composition and specialised predation on millipedes in the Enigma tic ponerine ant genusProbolomyrmex(Hymenoptera, Formicidae) "(PDF).Insectes Sociaux.45(1): 79–83.doi:10.1007/s000400050070.S2CID22119946.Archived(PDF)from the original on 2013-11-12.Retrieved2013-11-12.
  65. ^Herbert, D. G. (2000). "Dining on diplopods: remarkable feeding behaviour in chlamydephorid slugs (Mollusca: Gastropoda)".Journal of Zoology.251(1): 1–5.doi:10.1111/j.1469-7998.2000.tb00586.x.
  66. ^Forgie, Shaun A.; Grebennikov, Vasily V.; Scholtz, Clarke H. (2002). "Revision ofSceliagesWestwood, a millipede-eating genus of southern African dung beetles (Coleoptera: Scarabaeidae) ".Invertebrate Systematics.16(6): 931–955.doi:10.1071/IT01025.
  67. ^Larsen, T. H; Lopera, A.; Forsyth, A.; Genier, F. (2009)."From coprophagy to predation: a dung beetle that kills millipedes".Biology Letters.5(2): 152–155.doi:10.1098/rsbl.2008.0654.PMC2665820.PMID19158030.
  68. ^Forthman, M.; Weirauch, C. (2012)."Toxic associations: a review of the predatory behaviors of millipede assassin bugs (Hemiptera: Reduviidae: Ectrichodiinae)"(PDF).European Journal of Entomology.109(2): 147–153.doi:10.14411/eje.2012.019.Archived(PDF)from the original on 2014-07-14.Retrieved2014-07-14.
  69. ^Santamaría, Sergi; Enghoff, Henrik;Reboleira, Ana Sofía P.S.(2018)."New species ofTroglomycesandDiplopodomyces(Laboulbeniales, Ascomycota) from millipedes (Diplopoda) ".European Journal of Taxonomy(429).doi:10.5852/ejt.2018.429.
  70. ^Animals: The International Wildlife Magazine.Nigel-Sitwell. 1964. p. 21.
  71. ^Blum, Murray S.; Woodring, J. Porter (1962). "Secretion of benzaldehyde and hydrogen cyanide by the millipedePachydesmus crassicutis(Wood) ".Science.138(3539): 512–513.Bibcode:1962Sci...138..512B.doi:10.1126/science.138.3539.512.PMID17753947.S2CID40193390.
  72. ^Kuwahara, Yasumasa; Ômura, Hisashi; Tanabe, Tsutomu (2002). "2-Nitroethenylbenzenes as natural products in millipede defense secretions".Naturwissenschaften.89(7): 308–310.Bibcode:2002NW.....89..308K.doi:10.1007/s00114-002-0328-9.PMID12216861.S2CID30068731.
  73. ^Wood, William F. (1974). "Toluquinone and 2-Methoxy-3-methylbenzoquinone from the Defensive Secretions of Three African Millipedes".Annals of the Entomological Society of America.67(6): 988–989.doi:10.1093/aesa/67.6.988.
  74. ^Wood, William F.; Shepherd, Julian; Chong, Berni; Meinwald, Jerrold (1975). "Ubiquinone-0 in the defensive spray of an African Millipede".Nature.253(5493): 625–626.doi:10.1038/253625a0.PMID1113850.S2CID4195891.
  75. ^Wood, William F.; Hanke, Frederick J.; Kubo, Isao; Carroll, Jennifer A.; Crews, Phillip (2000). "Buzonamine, a new alkaloid from the defensive secretion of the millipede, Buzonium crassipes".Biochemical Systematics and Ecology.28(4): 305–312.Bibcode:2000BioSE..28..305W.doi:10.1016/s0305-1978(99)00068-x.PMID10725589.
  76. ^Weldon, Paul J.; Aldich, Jeffrey R.; Klun, Jerome A.; Oliver, James E.; Debboun, Mustapha (2003)."Benzoquinones from millipedes deter mosquitoes and elicit self-anointing in capuchin monkeys (Cebusspp.) ".Naturwissenschaften.90(7): 301–305.Bibcode:2003NW.....90..301W.doi:10.1007/s00114-003-0427-2.PMID12883771.S2CID15161505.Archivedfrom the original on 2021-02-25.Retrieved2018-04-29.
  77. ^Valderrama, Ximena; Robinson, John G.; Attygalle, Athula B.; Eisner, Thomas (2000). "Seasonal anointment with millipedes in a wild primate: a chemical defense against insects".Journal of Chemical Ecology.26(12): 2781–2790.doi:10.1023/A:1026489826714.S2CID25147071.
  78. ^Birkinshaw, Christopher R. (1999). "Use of millipedes by black lemurs to anoint their bodies".Folia Primatologica.70(3): 170–171.doi:10.1159/000021691.PMID10394067.S2CID36036598.
  79. ^Roncadori, R. W.; Duffey, S. S.; Blum, M. S. (1985). "Antifungal activity of defensive secretions of certain millipedes".Mycologia.77(2): 185–191.doi:10.2307/3793067.JSTOR3793067.
  80. ^Eisner, Thomas; Eisner, Maria; Deyrup, Mark (1996)."Millipede defense: use of detachable bristles to entangle ants".Proceedings of the National Academy of Sciences.93(20): 10848–10851.Bibcode:1996PNAS...9310848E.doi:10.1073/pnas.93.20.10848.PMC38244.PMID8855269.
  81. ^Stoev, Pavel; Lapeva-Gjonova, Albena (2005)."Myriapods from ant nests in Bulgaria (Chilopoda, Diplopoda)"(PDF).Peckiana.4:131–142. Archived fromthe original(PDF)on 2016-03-03.Retrieved2014-06-05.
  82. ^Farfan, Monica; Klompen, Hans (2012)."Phoretic mite associates of millipedes (Diplopoda, Julidae) in the northern Atlantic region (North America, Europe)".International Journal of Myriapodology.7:69–91.doi:10.3897/ijm.7.3064.
  83. ^Swafford, Lynn; Bond, Jason E. (2010)."Failure to cospeciate: an unsorted tale of millipedes and mites".Biological Journal of the Linnean Society.101(2): 272–287.doi:10.1111/j.1095-8312.2010.01499.x.
  84. ^Martínez-Torres, Shirley Daniella; Daza, Álvaro Eduardo Flórez; Linares-Castillo, Edgar Leonardo (2011)."Meeting between kingdoms: discovery of a close association between Diplopoda and Bryophyta in a transitional Andean-Pacific forest in Colombia".International Journal of Myriapodology.6:29–36.doi:10.3897/ijm.6.2187.
  85. ^Marshall, Michael (22 September 2011)."Zoologger: Stealth millipede wears living camouflage".New Scientist.Archivedfrom the original on 23 April 2015.Retrieved26 June2016.
  86. ^Mason, G.; Thompson, H.; Fergin, P.; Anderson, R. (1994). "Spot diagnosis: the burning millipede".Medical Journal of Australia.160(11): 718–726.doi:10.5694/j.1326-5377.1994.tb125915.x.PMID8202008.S2CID204065414.
  87. ^Shpall, S.; Frieden, I. (1991). "Mahogany discoloration of the skin due to the defensive secretion of a millipede".Pediatric Dermatology.8(1): 25–27.doi:10.1111/j.1525-1470.1991.tb00834.x.PMID1862020.S2CID1725209.
  88. ^Radford, A. (1976). "Giant millipede burns in Papua New Guinea".Papua New Guinea Medical Journal.18(3): 138–141.PMID1065155.
  89. ^Radford, A. (1975). "Millipede burns in man".Tropical and Geographical Medicine.27(3): 279–287.PMID1103388.
  90. ^Hudson, B.; Parsons, G. (1997). "Giant millipede 'burns' and the eye".Transactions of the Royal Society of Tropical Medicine and Hygiene.91(2): 183–185.doi:10.1016/S0035-9203(97)90217-0.PMID9196764.
  91. ^Alagesan, P.; Muthukrishnan, J. (2005)."Bioenergetics of the household pest,Xenobolus carnifex(Fabricius, 1775) "(PDF).Peckiana.4:3–14. Archived fromthe original(PDF)on 2016-03-03.Retrieved2013-11-12.
  92. ^Enghoff, Henrik; Kebapći, Ümit (2008). "Calyptophyllum longiventre(Verhoeff, 1941) invading houses in Turkey, with the first description of the male (Diplopoda: Julida: Julidae) ".Journal of Natural History.42(31–32): 2143–2150.Bibcode:2008JNatH..42.2143E.doi:10.1080/00222930802196055.S2CID84768307.
  93. ^Ebregt, E.; Struik, P. C.; Odongo, B.; Abidin, P. E. (2005)."Pest damage in sweet potato, groundnut and maize in north-eastern Uganda with special reference to damage by millipedes (Diplopoda)".NJAS - Wageningen Journal of Life Sciences.53(1): 49–69.doi:10.1016/S1573-5214(05)80010-7.S2CID54205396.
  94. ^Niijima, Keiko (2001).ヤケヤスデ đoàn tàu を ngăn める[A millipede outbreak (Oxidus gracilis,Koch) stopped trains].Edaphologia(in Japanese).68(68): 43–46.doi:10.20695/edaphologia.68.0_43.ISSN0389-1445.Archivedfrom the original on 2020-05-10.Retrieved2020-05-10.
  95. ^Peckham, Matt (4 September 2013)."Millipedes – Yes, Millipedes – May Be Responsible for Australian Train Crash".Time Newsfeed.Time Magazine.Archivedfrom the original on 10 November 2013.Retrieved31 October2013.
  96. ^Stoev, Pavel; Zapparoli, Marzio; Golovatch, Sergei; Enghoff, Henrik; Akkari, Nesrine; Barber, Anthony (2010)."Myriapods (Myriapoda). Chapter 7.2.In:Roqueset al.(Eds). Alien terrestrial arthropods of Europe ".BioRisk.4:97–130.doi:10.3897/biorisk.4.51.
  97. ^Lewbart, Gregory A., ed. (2011-09-20).Invertebrate Medicine(2nd ed.).Wiley-Blackwell.p. 255.ISBN978-0-470-96078-3.Archivedfrom the original on 2022-01-02.Retrieved2020-10-19.
  98. ^abcCosta Neto, Eraldo M. (2007)."The perception of Diplopoda (Arthropoda, Myriapoda) by the inhabitants of the county of Pedra Branca, Santa Teresinha, Bahia, Brazil".Acta Biológica Colombiana.12(2): 123–134.Archivedfrom the original on 2013-11-12.Retrieved2013-11-12.
  99. ^Negi, C. S.; Palyal, V. S. (2007)."Traditional uses of animal and animal products in medicine and rituals by the Shoka tribes of district Pithoragarh, Uttaranchal, India"(PDF).Studies on Ethno-Medicine.1(1): 47–54.doi:10.1080/09735070.2007.11886300.S2CID30993906.Archived(PDF)from the original on 2013-11-12.Retrieved2013-11-12.
  100. ^Jiang, T. L.; Feng, G. W.; Shen, J. H.; Li, L. F.; Fu, X. Q. (1981). "Observation of the effect ofSpirobolus bungiiextract on cancer cells ".Journal of Traditional Chinese Medicine.1(1): 34–8.PMID6926686.
  101. ^Halloran, Afton; Flore, Roberto; Vantomme, Paul; Roos, Nanna, eds. (2018).Edible insects in sustainable food systems.Cham: Springer. pp. xvii+479.ISBN978-3-319-74011-9.OCLC1036756200.
  102. ^abEnghoff, Henrik; Manno, Nicola; Tchibozo, Sévérin; List, Manuela; Schwarzinger, Bettina; Schoefberger, Wolfgang; Schwarzinger, Clemens; Paoletti, Maurizio G. (2014)."Millipedes as food for humans: their nutritional and possible antimalarial value: a first report".Evidence-Based Complementary and Alternative Medicine.2014:1–9.doi:10.1155/2014/651768.PMC3945075.PMID24688592.
  103. ^"Canada: Money in muskeg?".New Scientist:198–199. 25 April 1963.ISSN0262-4079.Archivedfrom the original on 30 November 2016.Retrieved14 August2016.
  104. ^Avirovik, Dragan; Butenhoff, Bryan; Priya, Shashank (2014). "Millipede-inspired locomotion through novel U-shaped piezoelectric motors".Smart Materials and Structures.23(3): 037001.Bibcode:2014SMaS...23c7001A.doi:10.1088/0964-1726/23/3/037001.S2CID109489932.
  105. ^Wakimoto, Shuichi; Suzumori, Koichi; Kanda, Takefumi (2006)."A bio-mimetic amphibious soft cord robot".Nihon Kikai Gakkai Ronbunshu, C Hen/Transactions of the Japan Society of Mechanical Engineers, Part C(in Japanese and English).72(2): 471–477.Archivedfrom the original on 2014-11-29.Retrieved2014-11-18.
  106. ^Beattie, Andrew; Ehrlich, Paul (2001).Wild Solutions: How Biodiversity is Money in the Bank(2nd ed.). New Haven: Yale University Press. pp. 192–194.ISBN978-0-300-10506-3.Archivedfrom the original on 2020-06-02.Retrieved2015-10-27.
  107. ^De Sousa Antunes, Luiz Fernando; Spolador Fernandes, Letícia; De Sousa Vaz, André Felipe; Santos Reis De Andrade Da Silva, Maura; Dos Santos Ferreira, Talita; Teles Dos Santos, Dieini Melissa; Fernandes Correia, Maria Elizabeth (2022)."Millicomposting: Sustainable technique for obtaining organic compost for the cultivation of broccoli seedlings".Cleaner Engineering and Technology.7:100442.Bibcode:2022CEngT...700442D.doi:10.1016/j.clet.2022.100442.S2CID246794375.
  108. ^Millicompost: an alternate biocompost for forest nurseries
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