Millipede
Millipedes Late | |
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An assortment of millipedes (not to scale) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Subphylum: | Myriapoda |
Class: | Diplopoda Blainville in Gervais, 1844 |
Subclasses | |
| |
Diversity | |
16 orders, c. 12,000 species |
Millipedes (originating from the
Most millipedes are slow-moving
First appearing in the
Among myriapods, millipedes have traditionally been considered most closely related to the tiny
The scientific study of millipedes is known as diplopodology, and a scientist who studies them is called a diplopodologist.
Etymology and names
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
Approximately 12,000 millipede species have 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 rapid speciation, 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
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 the author citation: the name of the person who coined the name or defined the group, even if not at the current rank.
Class Diplopoda de Blainville in Gervais, 1844
- Subclass PenicillataLatreille, 1831
- Order Polyxenida Verhoeff, 1934
- Subclass †Arthropleuridea (placed in Penicillata by some authors)[14]
- Order †ArthropleuridaWaterlot, 1934
- Order †EoarthropleuridaShear & Selden, 1995
- Order †MicrodecemplicidaWilson & Shear, 2000
- Order †
- Subclass Chilognatha Latreille, 1802
- Order †Zosterogrammida Wilson, 2005 (Chilognatha incertae sedis)[14]
- Infraclass Pentazonia Brandt, 1833
- Order †AmynilyspedidaHoffman, 1969
- Superorder LimacomorphaPocock, 1894
- Order Glomeridesmida Cook, 1895
- Superorder OniscomorphaPocock, 1887
- Order Glomerida Brandt, 1833
- Order Sphaerotheriida Brandt, 1833
- Order †
- Infraclass Helminthomorpha Pocock, 1887
- Superorder †Archipolypoda Scudder, 1882
- Order †Archidesmida Wilson & Anderson 2004
- Order †CowiedesmidaWilson & Anderson 2004
- Order †EuphoberiidaHoffman, 1969
- Order †PalaeosomatidaHannibal & Krzeminski, 2005
- Order †Pleurojulida Schneider & Werneburg, 1998 (possibly sister to Colobognatha)[9]
- Subterclass Colobognatha Brandt, 1834
- Order Platydesmida Cook, 1895
- Order Polyzoniida Cook, 1895
- Order Siphonocryptida Cook, 1895
- Order Siphonophorida Newport, 1844
- Subterclass Eugnatha Attems, 1898
- Superorder Juliformia Attems, 1926
- Order Julida Brandt, 1833
- Order Spirobolida Cook, 1895
- Order Spirostreptida Brandt, 1833
- Superfamily †Xyloiuloidea Cook, 1895 (Sometimes aligned with Spirobolida)[15]
- Superorder Nematophora Verhoeff, 1913
- Order Callipodida Pocock, 1894
- Order Chordeumatida Pocock 1894
- Order StemmiulidaCook, 1895
- Order SiphoniulidaCook, 1895
- Superorder Merocheta Cook, 1895
- Order Polydesmida Pocock, 1887
- Superorder Juliformia Attems, 1926
- Superorder †Archipolypoda Scudder, 1882
- Order †
Evolution
Millipedes are among the first animals to have
Living groups
The history of scientific millipede classification began with
In 1971, the Dutch biologist
Diplopoda |
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Fossil record
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.
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Alternate hypothesis of fossil relationships[9][29] |
Diplopoda |
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Relation to other myriapods
Although the relationships of millipede orders are still the subject of debate, the class Diplopoda as a whole is considered a
Distinction from centipedes
The differences between millipedes and centipedes are a common question from the general public.
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 burrowing soil 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
Millipedes come in a variety of body shapes and sizes, ranging from 2 mm (1⁄16 in) to around 35 cm (14 in) in length,[34] and can have as few as eleven to over three hundred segments.[35][36] They are generally black or brown in colour, although there are a few brightly coloured species, and some have aposematic colouring to warn that they are toxic.[5] Species of Motyxia produce cyanide as a chemical defence and are bioluminescent.[37]
Body styles vary greatly between major millipede groups. In the
They have also lost the gene that codes for the JHAMTl enzyme, 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.[38]
Head
The head of a millipede is typically rounded above and flattened below and bears a pair of large
Millipede eyes consist of several simple flat-lensed ocelli arranged 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 orders Polydesmida, Siphoniulida, Glomeridesmida, Siphonophorida and Platydesmida, and cave-dwelling millipedes such as Causeyella and Trichopetalum, had ancestors that could see but have subsequently lost their eyes and are blind.[34]
Body
Millipede bodies may be flattened or cylindrical, and are composed of numerous
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 the telson and consists of a legless preanal ring, a pair of anal valves (closeable plates around the anus), and a small scale below the anus.[9][35]
Millipedes in several orders have keel-like extensions of the body-wall known as
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.
Internal organs
Millipedes breathe through two pairs of spiracles located ventrally on each segment near the base of the legs.
Reproduction and growth
Millipedes show a diversity of mating styles and structures. In the basal order Polyxenida (bristle millipedes), mating is indirect: males deposit spermatophores onto webs they secrete with special glands, and the spermatophores are subsequently picked up by females.[31] In all other millipede groups, males possess one or two pairs of modified legs called gonopods which are used to transfer sperm to the female during copulation. The location of the gonopods differs between groups: in males of the Pentazonia they 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 are parthenogenetic, having few, if any, males.[45]
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.[46] The gonopods develop gradually from walking legs through successive moults until reproductive maturity.[47]
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 two penes which deposit the sperm packets onto the gonopods. In the female, the genital pores open into paired small sacs called cyphopods or vulvae, which are covered by small hood-like lids, and are used to store the sperm after copulation.[35] The cyphopod morphology can also be used to identify species. Millipede sperm lack flagella, 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, stridulation or "chirping".[48] 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.[42]
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 dried
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 continually moult, adding further segments and legs as they do so, a mode of development known as anamorphosis.[33] Some species moult within specially prepared chambers of soil or silk,[49] 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.[45] Millipedes may live from one to ten years, depending on species.[35]
Ecology
Habitat and distribution
Millipedes occur on all continents except Antarctica, and occupy almost all terrestrial habitats, ranging as far north as the
Burrowing
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.[5]
Diet
Most millipedes are detritivores and feed on decomposing vegetation, feces, or organic matter mixed with soil. They often play important roles in the breakdown and decomposition of plant 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.[42] Where earthworm populations 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 order Polyxenida graze algae from bark, and Platydesmida feed on fungi.[9] A few species are omnivorous or in Callipodida and Chordeumatida occasionally carnivorous,[56] feeding on insects, centipedes, earthworms, or snails.[35][57] Some species have piercing mouth parts that allow them to suck up plant juices.[31] Cave dwelling species in Julidae, Blaniulidae, and Polydesmidae have specialized mouthparts and appears to be filter feeders, filtering small particles from running water inside caves.[58]
Predators and parasites
Millipedes are preyed on by a wide range of animals, including various reptiles, amphibians, birds, mammals, and insects.[9] Mammalian predators such as coatis and meerkats roll captured millipedes on the ground to deplete and rub off their defensive secretions before consuming their prey,[59] and certain poison dart frogs are believed to incorporate the toxic compounds of millipedes into their own defences.[60] Several invertebrates have specialised behaviours or structures to feed on millipedes, including larval glowworm beetles,[61] Probolomyrmex ants,[62] chlamydephorid slugs,[63] and predaceous dung beetles of the genera Sceliages and Deltochilum.[64][65] A large subfamily of assassin bugs, the Ectrichodiinae with over 600 species, has specialised in preying upon millipedes.[66] Parasites of millipedes include nematodes, phaeomyiid flies, and acanthocephalans.[9] Nearly 30 fungal species of the order Laboulbeniales have been found growing externally on millipedes, but some species may be commensal rather than parasitic.[67]
Defence mechanisms
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.[68]
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
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.[78]
Other inter-species interactions
Some millipedes form mutualistic relationships with organisms of other species, in which both species benefit from the interaction, or commensal relationships, in which only one species benefits while the other is unaffected. Several species form close relationships with ants, a relationship known as myrmecophily, especially within the family Pyrgodesmidae (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.[79]
Many millipede species have commensal relationships with
A novel interaction between millipedes and mosses was described in 2011, in which individuals of the newly discovered Psammodesmus bryophorus was found to have up to ten species living on its dorsal surface, in what may provide camouflage for the millipede and increased dispersal for the mosses.[82][83]
Interactions with humans
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 agricultural pest. 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, local erythema, edema, blisters, eczema, and occasionally cracked skin.[84][85][86][87] Eye exposures to these secretions causes general irritation and potentially more severe effects such as conjunctivitis and keratitis.[88] This is called millipede burn. First aid consists of flushing the area thoroughly with water; further treatment is aimed at relieving the local effects.
Some millipedes are considered household pests, including
Some of the larger millipedes in the orders Spirobolida, Spirostreptida, and Sphaerotheriida are popular as pets.[94] Some species commonly sold or kept include species of Archispirostreptus, Aphistogoniulus, Narceus, and Orthoporus.[95]
Millipedes appear in
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.[101] Experimental robots have had the same inspiration,[102][103] in particular when heavy loads are needed to be carried in tight areas involving turns and curves.[104] In biology, some authors have advocated millipedes as model organisms for the study of arthropod physiology and the developmental processes controlling the number and shape of body segments.[42]
Similar to vermicompost, millipedes can be used to convert plant matter into compost in what has been named millicomposting, which improves the quality of the compost.[105][106]
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External links
- Milli-PEET: The Class Diplopoda – The Field Museum, Chicago
- Millipedes of Australia Archived 2018-06-27 at the Wayback Machine
- Diplopoda: Guide to New Zealand Soil Invertebrates – Massey University
- SysMyr, a myriapod taxonomy database Archived 2020-04-25 at the Wayback Machine
- British Myriapod & Isopod Group