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Polydesmida
Polydesmida
Polydesmida
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Not to be confused with Platydesmida or Polyzoniida.

Polydesmida
Apheloria corrugata
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Myriapoda
Class: Diplopoda
Subclass: Chilognatha
Infraclass: Helminthomorpha
Subterclass: Eugnatha
Superorder: Merocheta
Cook, 1895
Order: Polydesmida
Leach, 1815[1]
Suborders
Synonyms

Proterospermatophora Verhoeff, 1900

Polydesmida (from the Greek poly "many" and desmos "bond") is the largest order of millipedes, with more than 5,000 species,[2][3][4][5] including all the millipedes reported to produce hydrogen cyanide (HCN).[6] This order is also the most diverse of the millipede orders in terms of morphology.[7] Millipedes in this order are found in all regions of the world other than Antarctica.[7]

Description

[edit]

Members of the order Polydesmida are also known as flat-backed millipedes, because on most species, each body segment has wide lateral keels known as paranota.[8][9] These keels are produced by the posterior half (metazonite) of each body ring behind the collum.[10] Polydesmids have no eyes, and vary in length from 1.4 to 134 mm.[7] Many of the larger species show bright coloration patterns which warn predators of their toxic secretions.[11]

Adults usually have 20 segments,[11] counting the collum as the first ring and the telson as the last ring.[12][13] Juveniles have from 7 to 19 rings.[10] In species with the usual 20 segments, adult females have 31 pairs of legs, but in adult males, the eighth leg pair (the first leg pair of the 7th ring) is modified into a single pair of gonopods, leaving only 30 pairs of walking legs.[10][12][13]

Variation

[edit]

Many species deviate from the typical body plan. One striking and unique deviation occurs in adult males of the species Aenigmopus alatus, which retain 31 pairs of walking legs and feature no gonopods.[12][14] This millipede is the only species in the infraclass Helminthomorpha without gonopods.[7]

The most common deviation, however, is a reduction in the number of segments. Many species have only 19 segments (including the telson) as adults, including those in the genera Brachydesmus, Macrosternodesmus, and Bacillidesmus.[13] In these species, adult females have only 29 pairs of legs, and adult males have only 28 pairs of walking legs.[10][12] In a few species, including Agenodesmus reticulatus, Deharvengius bedosae, and Doratodesmus hispidus, adults have only 18 segments (including the telson), with a corresponding reduction in the number of leg pairs (27 in the adult female, 26 in the adult male, excluding the gonopods).[13] Still other species exhibit sexual dimorphism in segment number, for example, Prosopodesmus panporus (the usual 20 in adult females, but only 19 in adult males) and Doratodesmus pholeter (19 in adult females; 18 in adult males), with the expected number of leg pairs given the number of segments in each sex.[13] Even more unusual are two species, Ammodesmus congoensis and A. granum, in which adults in each sex can have 18 or 19 segments.[15][16][7]

Flat-backed Millipede (Polydesmida)
Flat-backed Millipede (Polydesmida)

A few species deviate by having more than the usual number of segments, including those in the cave-dwelling genus Devillea.[13][12] For example, in the species D. tuberculata, adult females have 22 segments and adult males have 21 (including the telson), with a corresponding increase in the number of leg pairs (35 in adult females and 32 in adult males, excluding the gonopods).[17][13][18] Some species in this genus also exhibit variation in segment number within the same sex, for example, in D. subterranea, adult males can have as few as 19 segments or as many as 23 (including the telson).[13][18] The most extreme outlier in segment number among polydesmids, however, is a cave-dwelling species discovered in Brazil, Dobrodesmus mirabilis, with adult males found to have 40 segments (including the telson).[4][12]

Development

[edit]

Polydesmids grow and develop through a series of molts, adding segments until they reach a fixed number in the adult stage, which is usually the same for a given sex in a given species, at which point the molting and the addition of segments and legs stop.[13] This mode of development, known as teloanamorphosis, distinguishes this order from most other orders of millipedes. Millipedes in other orders usually continue to molt as adults, developing through either euanamorphosis or hemianamorphosis.[13][19]

Millipedes in this order typically develop through a series of eight stages, hatching with only 7 segments (including the telson) and 3 pairs of legs, then molting seven times. These eight stages feature 7, 9, 12, 15, 17, 18, 19, then finally 20 segments. The adults then reproduce and die without another molt.[13] Species with fewer than the usual 20 segments as adults go through the same stages observed in other polydesmids until reaching maturity, which occurs one molt earlier for 19 segments or two molts earlier for 18 segments.[13][4] Species with more than the usual 20 segments are thought to add these extra segments through additional molts, adding one segment per molt.[13][4]

Ecology

[edit]

Polydesmids are very common in leaf litter, where they burrow by levering with the anterior end of the body.[9] These millipedes feed on decaying vegetation. Some species are prey for funnel-web spiders.[20]

Classification

[edit]

The species of Polydesmida are variously classified into four suborders (names ending in "-idea"), and 29 families, the largest (numerically) including Paradoxosomatidae, Xystodesmidae, and Chelodesmidae.[21]

Dalodesmidea Hoffman, 1980. 2 families
Leptodesmidea Brölemann, 1916. 13 families
Paradoxosomatidea Daday, 1889.[a] 1 family
Polydesmidea Pocock, 1887. 12 families
Representative diversity of Polydesmida

References

[edit]
[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Polydesmida

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from Grokipedia
Polydesmida is the largest order of s in the class Diplopoda, comprising approximately 5,500 described that account for nearly half of all known millipede diversity, and is characterized by flattened, wedge-shaped bodies with distinctive lateral extensions of the tergites called paranota, which often bear spines or keels for defense. These lack eyes, have short antennae with eight articles, and typically exhibit 20 body rings in adults, adapting them for life in moist, terrestrial environments such as leaf litter and . Polydesmida are cosmopolitan in distribution, with the highest diversity in tropical and subtropical regions, though they occur worldwide in forests, gardens, and under decaying wood. Taxonomically, Polydesmida belongs to the subclass Chilognatha and superorder Merocheta (or sometimes proposed within an "eighth gonopod clade" alongside orders like Callipodida and Chordeumatida), encompassing over 30 families, more than 500 genera, and a vast array of morphological variations in paranota shape and gonopod structure that aid in species identification. The order's evolutionary history traces back to ancient lineages, with phylogenetic analyses placing it variably as sister to groups like Colobognatha or Julida based on genetic data. Notable families include Xystodesmidae and Polydesmidae, the latter predominantly Holarctic with over 60 genera and nearly 300 species. Ecologically, polydesmid millipedes play key roles as decomposers in forest ecosystems, feeding primarily on decaying plant matter and fungi while contributing to nutrient cycling. They are uniquely equipped with chemical defenses, including cyanogenic compounds that produce (HCN) upon threat, alongside like cresols and , and , making them unpalatable to predators. Some species exhibit specialized adaptations, such as semi-aquatic habits allowing submersion tolerance or arboricolous lifestyles on bark, and a few are troglobitic cave-dwellers. Their abundance in regions like underscores their ecological significance, though habitat loss poses ongoing threats to many populations.

Taxonomy and Overview

Classification and Phylogeny

Polydesmida is an order of millipedes within the subclass Chilognatha and class Diplopoda, encompassing approximately 5,500 described species distributed across over 500 genera and about 30 families as of 2025. This order represents the largest and most diverse group of millipedes, characterized by their keeled or flat-backed body form, and is placed in the superorder Merocheta (also known as Merochaeta). The species are classified into four suborders—Dalodesmidea, Leptodesmidea, Paradoxosomatidea, and Polydesmidea—with the latter two being particularly species-rich. Key families include Paradoxosomatidae, the largest with over 1,000 species primarily in tropical regions; Xystodesmidae, with over 500 species mainly in North America; and Chelodesmidae, noted for its diversity in the Neotropics. Phylogenetically, Polydesmida occupies a basal position among eugnathan millipedes, supported by both molecular and morphological analyses that highlight its early divergence within the subclass. Molecular phylogenomics, including multi-locus studies, recover polydesmidans as to other major eugnathan lineages such as , with morphological traits like the gnathochilarium and reduced tracheal systems reinforcing this placement. The order's is well-established, though internal relationships among suborders remain under revision through integrative approaches combining and . The taxonomic history of Polydesmida dates to its establishment as an order by in 1815, initially based on European species with distinctive body keels. Major revisions occurred in the , notably by Carl Attems, who expanded the in his 1937 monograph on myriapods, and Richard L. Hoffman, whose 1976 and 1999 works refined family-level boundaries and synonymies. Contemporary updates incorporate integrative , blending morphological, molecular, and ecological data to address cryptic diversity, particularly in cyanogenic clades that produce as a defense. Recent developments have accelerated species discovery, with hundreds of new taxa described since 2020, driven by targeted surveys in biodiversity hotspots. Notable examples include seven new species of Xystodesmus (Xystodesmidae) from southwestern in 2025, expanding the genus's known range in , and two new species of Inodesmus (Haplodesmidae) from Colombian oak forests in 2025, marking the first records of that family in the country. Earlier contributions include the genus Nagaxytes, established in 2018 for Southeast Asian species with extended paranota, highlighting ongoing refinements in suborder Paradoxosomatidea. These additions underscore the emphasis on cyanogenic lineages, which dominate phylogenetic studies of the order. Evolutionary insights portray Polydesmida as an ancient lineage, with fossil records from the suggesting origins in the . The group's is evident in its diversification across humid tropical and subtropical environments, where ecological specialization in leaf litter and niches has driven , particularly in the suborders Paradoxosomatidea and Polydesmidea. This radiation is supported by estimates placing key divergences in the late to early .

Distribution and Diversity

Polydesmida exhibit a , occurring on all continents except and being absent from extreme desert environments, with their presence documented across a wide array of temperate, subtropical, and tropical habitats worldwide. The order's range spans from boreal forests in and to humid rainforests in the Neotropics and Indo-Malaya, reflecting adaptations to diverse climatic conditions but with a clear preference for moist, vegetated areas. Highest species diversity is concentrated in tropical regions, particularly , , and , where environmental stability and habitat complexity support elevated richness. Biogeographic patterns within Polydesmida suggest pantropical origins, with subsequent radiations leading to temperate extensions, such as in where the family Xystodesmidae dominates forest assemblages. Island endemism is prominent in isolated systems like and the , where unique evolutionary trajectories have produced regionally restricted lineages, including chelodesmid millipedes in the . These patterns underscore the order's historical dispersal via vicariance and overwater colonization, with tropical hotspots serving as centers of . The order comprises approximately 5,500 described , with estimates suggesting significantly more undescribed taxa, making it the most diverse order. Biodiversity hotspots include , home to numerous of dragon millipedes in the genus Desmoxytes (), and the Colombian , where Cryptodesmidae exhibit high local amid Andean cloud forests. The majority of are endemic to . Recent discoveries, such as 18 new miniature in the Polydesmidae from in 2021, illustrate ongoing revelations of cryptic diversity in temperate zones. Threats to Polydesmida diversity primarily stem from tropical loss due to , , and , which disproportionately affect undescribed species in biodiversity hotspots. Most species remain unassessed, with conservation statuses classified as under IUCN criteria, underscoring the urgent need for expanded surveys and protection to mitigate extinction risks in vulnerable ecosystems.

Morphology

General Body Structure

Polydesmida exhibit a distinctive body form that ranges from cylindrical to dorsoventrally flattened, primarily due to the presence of paranota, which are lateral keels or wing-like extensions on the dorsal surface of most body segments. These paranota serve protective and possibly display functions, contributing to the order's characteristic flat-backed appearance in many species, while in others they are reduced, resulting in a more cylindrical shape. The body comprises a head followed by typically 20 rings: the legless collum (ring 1), three thoracic rings (2–4) each bearing one pair of legs, 14 diplosegments (rings 5–18) each with two pairs of legs, a legless anal ring (19), and the (20), which includes the preanal ring and anal plates. The head is eyeless, lacking ocelli, and relies on other sensory structures for navigation in dark or humid environments. A pair of antennae, each with eight segments, protrudes from the head and plays a key role in chemosensation and mechanoreception, enabling detection of food, mates, and environmental cues. The exoskeleton is hardened by calcium carbonate deposits, particularly in the tergites, providing structural rigidity while allowing flexibility for movement. Coloration is often aposematic, featuring bold patterns in red, yellow, orange, or black to signal toxicity to predators, a trait linked to their chemical defenses. Body length varies from approximately 3 mm to over 130 mm, with most species falling between 5 and 50 mm. Internally, Polydesmida possess a simple tubular gut adapted for detritivory, consisting of , , and regions that facilitate the breakdown of decaying plant material and fungi through microbial . Repugnatorial glands, arranged segmentally, produce (HCN) from cyanogenic precursors, which can be released as a defensive ; these glands open via ozopores on the paranota or body sides. In males, the eighth leg pair is modified into gonopods for transfer, resulting in 30 pairs of walking legs, while females have 31 pairs.

Segmental and Appendage Variations

While most Polydesmida exhibit a typical with 20 segments (including the collum and ), notable variations occur across , with adult segment counts ranging from 18 to 21 rings in many families. This range often shows sex-specific differences, such as females having one more segment than males in certain taxa, reflecting modifications associated with reproductive structures. Extreme deviations include the Brazilian cave-dwelling Dobrodesmus mirabilis, discovered in 2016, which possesses 40 segments in adult males, far exceeding the norm and accompanied by 79 pairs of legs, highlighting evolutionary adaptations in isolated subterranean environments. Such supernumerary segmentation underscores the plasticity in polydesmidan trunk development, potentially linked to habitat-specific selective pressures. Appendage diversity in Polydesmida is particularly evident in the male gonopods, derived from the eighth leg pair, which display intricate morphologies crucial for species recognition during . These structures vary widely in branching patterns, lengths, and ornamentation even within , serving as primary diagnostic traits for and enabling precise mate discrimination among closely related . A rare exception is Aenigmopus alatus, a Central American in the Tridontomidae, where males lack gonopods entirely and retain 31 pairs of unmodified walking legs, an anomaly unique among helminthomorph millipedes that may imply alternative reproductive mechanisms. Paranota, the lateral extensions of the dorsal tergites often functioning as keels, exhibit significant morphological diversity, ranging from simple flanges to elaborate, spine-like crests that enhance body width and possibly aid in locomotion or defense. In the Southeast Asian genus Desmoxytes, known as dragon millipedes, paranota can form striking, ornate projections, such as the long, thin, spiniform types in cave-adapted like D. spinissima, which may facilitate navigation in confined spaces. Some families display in paranota, with males featuring more pronounced or differently shaped extensions compared to females, contributing to mate attraction or isolation. Miniaturization represents another key variation, with some Holarctic Polydesmidae species attaining lengths under 2 mm, such as certain members of oligotypic genera measuring 1.4–5.0 mm and possessing only 18 or 19 segments. These diminutive forms, often found in litter, contrast with elongated cave-adapted taxa in families like , where troglobitic evolve longer bodies, extended legs, and antennae to traverse dark, narrow habitats, as seen in elongated forms from Chinese karst caves. Intraspecific variation further diversifies Polydesmida morphology, with color morphs—such as pale grayish versus dark brown patterns in Xystodesmus species—influenced by environmental factors like substrate and humidity, potentially serving adaptive roles in or warning signaling. Similarly, keel (paranota) shapes show plasticity, with broader or more rounded forms in humid dwellers versus narrower ones in drier microhabitats, reflecting responses to local ecological conditions.

Development and Reproduction

Life Cycle Stages

Polydesmida exhibit teloanamorphic development, a post-embryonic growth pattern in which segment addition ceases after a fixed number of stadia, culminating in the form with no subsequent molts. This process typically involves 8 stadia, during which the progressively adds pairs of segments and legs from the posterior end until reaching the definitive segment count, which varies by species but often totals around 19–20 leg-bearing diplosegments. occurs in Stadium I with 7 body rings, including 3 pairs of walking legs on the initial diplosegments, followed by the collum and a few apodous rings. Subsequent stadia build on this foundation: Stadium II adds the first additional segment pair, Stadium III adds another, and so on, with segments incorporated behind the until Stadium VIII, when the final adult morphology is achieved. In species like Poratia salvator, development progresses through 7 post-hatching stages (equivalent to 8 including the hatching ), with juveniles in early stadia showing high mortality rates (up to 54% between stages) as they molt to incorporate new segments. Growth occurs via gradual size increases across molts, triggered by environmental cues such as , temperature, and photoperiod, which regulate the timing of ; maturity is typically reached after 1–2 years in natural conditions. Juveniles in initial stadia possess reduced paranota (lateral projections on the tergites) that are shorter and less developed compared to adults, along with simpler appendages lacking full . The transition to adult form happens during the final molt, where paranota expand fully, appendages mature, and gonopods form in males, marking the end of segment addition. In the wild, Polydesmida spans 2–5 years, with individuals capable of iteroparity, producing multiple broods over successive reproductive seasons after reaching maturity. For instance, in Polydesmus angustus, late-season cohorts extend their life cycle to 2 years to align with optimal breeding windows, supporting repeated reproduction.

Reproductive Strategies

Polydesmida primarily reproduce sexually through indirect sperm transfer, where males use their specialized gonopods—modified legs on the seventh body ring—to collect from the gonopores on the second ring and insert it into the female's cyphopods on the third ring, often in the form of spermatophores for precise . This mechanism ensures targeted deposition, with the gonopods locking into the female's reproductive openings during copulation, which typically lasts several minutes to hours depending on species and density. Mating in Polydesmida involves chemical and behavioral cues, including pheromone attraction to draw partners, particularly during humid seasons when activity peaks from June to September in temperate regions. In some species, males engage in precopulatory struggles or with rivals, involving coiling, pushing, and coercion to secure mating access, while mate-guarding post-copulation can last days to prevent remating. Females exhibit ranging from 7–24 eggs per , which are buried in nests constructed from fecal material and ; incubation periods last 10–21 days before hatching. is rare in Polydesmida but documented in certain populations, such as the thelytokous form in Poratia salvator, where unfertilized eggs develop into females, potentially aiding survival in isolated habitats. Sex ratios in Polydesmida populations are typically near 1:1, though some exhibit slight bias during breeding seasons; females are generally larger than males, reflecting sexual size dimorphism that supports higher egg production, with maintained through obligatory in most species.

Ecology and Behavior

Habitats and Feeding

Polydesmida millipedes predominantly inhabit moist microhabitats that provide high humidity and , such as , layers, and under loose bark on trees or logs. In temperate regions, like those in the family Xystodesmidae thrive in humus-rich deciduous or mixed forests, including the U.S. Midwest and woodlands, where they into or riparian zones with coniferous cover. Tropical species exhibit greater vertical stratification, occupying not only the but also extending into canopy layers of rainforests, where decaying epiphytes and accumulate. As detritivores, Polydesmida primarily consume decomposing organic material, including decaying wood, leaf , and fungi, contributing to nutrient cycling in their ecosystems. Some species engage in mycophagy, selectively feeding on fungal hyphae or spores within , while gut microbial symbionts assist in breaking down and for nutrient extraction. For example, temperate Xystodesmidae in forests target lichens, alongside fungi in bark and litter habitats. Foraging in Polydesmida is characterized by slow, deliberate crawling, often nocturnal to avoid and predation, with activity peaking during periods of high such as after rainfall or in the . Seasonal patterns vary by region; temperate populations show fall abundance in moist , while tropical forms maintain year-round activity in consistently humid environments. Microhabitat adaptations include lifestyles on the surface for many species, contrasted with geophilous burrowing in soil, and specialized troglomorphic forms in caves, such as cavernicolous Polydesmidae in Vietnamese systems, which rely on limited detrital inputs in dark, stable subterranean niches.

Defenses and Interactions

Polydesmida employ sophisticated chemical defenses primarily through repugnatorial glands located on their lateral body segments, which eject noxious secretions when threatened. These glands store cyanogenic precursors such as mandelonitrile or , which, upon release, are enzymatically degraded by hydroxynitrile lyase to produce hydrogen cyanide (HCN) and . The biosynthesis pathway derives from aromatic like , with potentially aiding in maintaining acidity for precursor stability, particularly in immature stages. HCN is highly toxic, proving fatal to arthropods and small vertebrates in confined spaces, while benzaldehyde contributes irritating odors and tastes that deter a broad spectrum of predators, including , beetles, spiders, and vertebrates. Some species also incorporate quinones or , enhancing overall repellency against and microbial threats. In addition to chemical mechanisms, Polydesmida utilize behavioral defenses to evade predation. Many species coil their bodies into tight spirals or balls, shielding vulnerable undersides and directing secretions outward from the exposed dorsum. This is often accompanied by thanatosis, where individuals feign death by remaining immobile, reducing attractiveness to visually hunting predators. Aposematic coloration, featuring bold patterns of red, yellow, or black, serves as a visual warning signal in several families like Xystodesmidae and Chelodesmidae, unpalatability and promoting avoidance learning in predators. Polydesmida serve as primary prey for various predators in forest litter and soil ecosystems, influencing co-evolutionary dynamics. They are commonly consumed by centipedes, ground-dwelling spiders (including Atypidae funnel-web spiders), birds, and amphibians, with predation often occurring in moist leaf habitats where these millipedes . Specialized predators like phengodid larvae circumvent chemical defenses by injecting digestive fluids to subdue and consume them. These interactions have driven adaptations in both millipedes and predators, such as enhanced tolerance in litter arthropods. Ecological interactions of Polydesmida extend to mutualistic, competitive, and parasitic relationships. They engage in mutualism with fungi by dispersing spores through fecal pellets, which aids fungal propagation while millipedes benefit from nutrient-rich . As detritivores, they compete with other invertebrates like isopods and earthworms for decomposing , influencing rates. is relatively rare but documented, with nematodes of the orders Rhigonematomorpha and Oxyuridomorpha inhabiting their hindguts, such as Rhigonema naylae in xystodesmid hosts. In human contexts, Polydesmida contribute to by enhancing nutrient cycling and microbial activity through detritivory, supporting ecosystem fertility. Certain , such as Oxidus gracilis (Paradoxosomatidae), act as minor pests in greenhouses, damaging seedlings and requiring management in controlled environments.

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