Polydesmida

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Polydesmida
Apheloriavirginiensis.jpg
Apheloria virginiensis
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
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, containing approximately 3,500 species, [2] including all the millipedes reported to produce hydrogen cyanide (HCN). [3] Polydesmids grow and develop through a series of moults, 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 moulting and the addition of segments and legs stop. [4] This mode of development, known as teloanamorphosis, distinguishes this order from most other orders of millipedes, which usually continue to moult as adults, developing through either euanamorphosis or hemianamorphosis. [4] [5]

Contents

Description

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. [6] [7] These keels are produced by the posterior half (metazonite) of each body ring behind the collum. [8] Polydesmids have no eyes, and vary in length from 3 to 130 mm (0.12 to 5.12 in). [9] Many of the larger species show bright coloration patterns which warn predators of their toxic secretions. [10]

Adults usually have 20 segments, [10] counting the collum as the first ring and the telson as the last ring. [11] [4] Juveniles have from 7 to 19 rings. [8] 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. [8] [11] [4]

Variation

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. [11] [12] This millipede is the only species in the infraclass Helminthomorpha without gonopods. [13]

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, Poratia, and Bacillidesmus. [4] In these species, adult females have only 29 pairs of legs, and adult males have only 28 pairs of walking legs. [8] [11] In a few species, including Hexadesmus lateridens , Agenodesmus reticulatus , and Eutynellus flavior , 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). [4] 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. [4] Even more unusual are two species, Ammodesmus congoensis and A. granum , in which adults in each sex can have 18 or 19 segments. [14] [15] [13]

Flat-backed Millipede (Polydesmida) Flat-backed Millipede (Polydesmida).jpg
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 . [4] [11] 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). [16] [4] [17] 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). [4] [17] 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). [18] [11]

Ecology

Polydesmids are very common in leaf litter, where they burrow by levering with the anterior end of the body. [7] Some are preyed on by funnel-web spiders. [19]

Classification

The c. 3500 species of Polydesmida are variously classified into four suborders (names ending in "-idea"), and 29 families, the largest (numerically) including Paradoxosomatidae, Xystodesmidae, and Chelodesmidae. [2]

Dalodesmidea Hoffman, 1980. 2 families
Leptodesmidea Brölemann, 1916. 13 families
Paradoxosomatidea Daday, 1889. [lower-alpha 1] 1 family
Polydesmidea Pocock, 1887. 12 families

Related Research Articles

<span class="mw-page-title-main">Millipede</span> Class of arthropods

Millipedes are a group of arthropods that are characterised by having two pairs of jointed legs on most body segments; they are known scientifically as the class Diplopoda, the name derived from this feature. Each double-legged segment is a result of two single segments fused together. Most millipedes have very elongated cylindrical or flattened bodies with more than 20 segments, while pill millipedes are shorter and can roll into a tight ball. Although the name "millipede" derives from Latin for "thousand feet", no species was known to have 1,000 or more until the discovery in 2020 of Eumillipes persephone, which can have over 1,300 legs. There are approximately 12,000 named species classified into 16 orders and around 140 families, making Diplopoda the largest class of myriapods, an arthropod group which also includes centipedes and other multi-legged creatures.

<i>Harpaphe haydeniana</i> Species of millipede

Harpaphe haydeniana, commonly known as the yellow-spotted millipede, almond-scented millipede or cyanide millipede, is a species of polydesmidan ("flat-backed") millipede found in the moist forests along the Pacific coast of North America, from Southeast Alaska to California, and Thailand .The dark coloration with contrasting yellow-tipped keels warn of its ability to exude toxic hydrogen cyanide as a defense. The cyanide secretions are not dangerous to humans, but can cause irritation and pain if it contacts sensitive areas such as the mouth, eyes, or nose.

<span class="mw-page-title-main">Xystodesmidae</span> Family of millipedes

Xystodesmidae is a family of millipedes. Its members often have very small distributional areas, with many species only known from a single locality. They are found across the northern hemisphere, with peak diversity in the Appalachian Mountains, where one-third of the 300 or so species occur. They are particularly abundant in deciduous broadleaf forests in the Mediterranean Basin, Africa, Asia, Central and North America, and Russia. Information on basic taxonomy is scant for this family; for example, it is estimated that the genus Nannaria contains over 200 species, but only 25 were described as of 2006. By 2022, 78 species in Nannaria have been described.

Tridontomidae is a small family of millipedes. Its members are endemic to Guatemala. These millipedes range from 22 mm to 28 mm in length and are uniformly grayish in color; their legs and antennae are unusually long and slender. This family includes the remarkable species Aenigmopus alatus, in which adult males feature no gonopods. This millipede is the only species in the infraclass Helminthomorpha without gonopods.

<span class="mw-page-title-main">Julida</span> Order of millipedes

Julida is an order of millipedes. Members are mostly small and cylindrical, typically ranging from 10–120 millimetres (0.39–4.72 in) in length. Eyes may be present or absent, and in mature males of many species, the first pair of legs is modified into hook-like structures. Additionally, both pairs of legs on the 7th body segment of males are modified into gonopods.

<i>Orthomorpha</i> Genus of millipedes

Orthomorpha is a genus of millipedes in the family Paradoxosomatidae containing approximately 50 species distributed in Southeast Asia.

<span class="mw-page-title-main">Chordeumatida</span> Order of millipedes

Chordeumatida is a large order of millipedes containing some 1200 species with a nearly worldwide distribution. Also known as sausage millipedes, they grow and develop through a series of moults, 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 moulting and the addition of segments and legs stop. This mode of development, known as teloanamorphosis, distinguishes this order from most other orders of millipedes, which usually continue to moult as adults, developing through either euanamorphosis or hemianamorphosis.

<span class="mw-page-title-main">C. A. W. Jeekel</span> Dutch myriapodologist and entomologist

Casimir Albrecht Willem Jeekel (1922–2010) was a Dutch myriapodologist and entomologist known for his major contributions to the taxonomy of millipedes. His 1971 monograph Nomenclator Generum et Familiarum Diplopodorum is credited as launching the "modern era" of millipede taxonomy, and has been considered the "most important single work ever published on the Diplopoda". He served as director of the Zoological Museum Amsterdam, and authored over 150 works on the taxonomy of millipedes and other myriapods.

<span class="mw-page-title-main">Platyrhacidae</span> Family of millipedes

Platyrhacidae is a family of polydesmidan millipedes distributed in Southeast Asia and tropical Central and South America.

<span class="mw-page-title-main">Haplodesmidae</span> Family of millipedes

Haplodesmidae is a family of millipedes in the order Polydesmida. Species occur in East Asia, Southeast Asia, and Oceania, although some species have been introduced to the New world tropics. Species are small bodied, often with elaborate sculpturing on the tergites, and some species are capable of rolling into a near-complete ball.

<span class="mw-page-title-main">Ammodesmidae</span> Family of millipedes

Ammodesmidae is a family of small millipedes endemic to Africa, containing seven species in two genera. Ammodesmids range from 1.4 to 5.0 mm long with 18 or 19 body segments in both sexes, and are capable of rolling into a tight sphere.

<span class="mw-page-title-main">Metopidiotrichidae</span> Family of millipedes

Metopidiotrichidae is a family of millipedes in the order Chordeumatida. These millipedes range from 4 mm to 17 mm in length. Adult millipedes in this family have 32 segments, not the 30 segments usually found in this order. Adult males in this family often feature a reduced or vestigial leg pair 10 as part of the gonopod complex, in addition to the two leg pairs typically modified into gonopods in this order. There are about 9 genera and at least 70 described species in Metopidiotrichidae.

<i>Brachydesmus</i> Genus of millipedes

Brachydesmus is a genus of millipedes belonging to the family Polydesmidae. Millipedes in this genus are found mainly in Europe. Species in this genus have only 19 segments as adults, one fewer than found in most polydesmid species. Accordingly, adults in this genus have two fewer legs than most polydesmid adults have: Females have only 29 pairs of legs, and males have only 28 pairs of walking legs, excluding one pair of gonopods. Species in this genus arrive at these lower numbers of legs and segments by going through the same stages of teloanamorphosis observed in other polydesmids but reaching maturity one moult earlier.

Trichopolydesmidae is a family of millipedes belonging to the order Polydesmida. This family includes two genera notable for featuring sexual dimorphism in segment number: adult females in these genera have the 20 segments usually found in this order, but adult males have only 19. This family also includes the species Deharvengius bedosae, notable for being among the very few species in this order to feature adults with only 18 segments rather than the 20 segments usually found in polydesmids.

Chordeumatidae is a family of millipedes belonging to the order Chordeumatida. These millipedes range from 7 mm to 18 mm in length and are found in Europe. Adult millipedes in this family have either 28 or 30 segments. This family features distinctive sex-linked modifications to the legs in adults: In the adult female, a legless sternite replaces the third pair of legs, and in the adult male, five pairs of legs are modified in the gonopod complex. These modifications are more extensive than those found in other adult males in this order, which often have only two leg pairs modified into gonopods. With the more extensive modifications to the legs in this family, species with the usual 30 segments feature adult females with only 49 leg pairs and adult males with only 45 pairs of walking legs, and species with only 28 segments feature adult females with only 45 leg pairs and adult males with only 41 pairs of walking legs.

Haaseidae is a family of millipedes belonging to the order Chordeumatida. Adult millipedes in this family range from 4.5 mm to 12 mm in length and have either 28 or 30 segments. For example, Orobainosoma hungaricum orientale, a subspecies of Haasea hungarica, has only 28 segments in adults rather than the 30 usually found in chordeumatidan adults, with only 46 pairs of legs in adult females and only 44 pairs of walking legs in adult males.

Hoffmaneumatidae is a family of millipedes belonging to the order Chordeumatida. This family includes two genera, one (Hoffmaneuma) found in the Russian Far East and the other (Japanoparvus) found in Japan. Millipedes in this family range from 4 mm to 6 mm in length. Adult millipedes in this family have only 28 segments, not the 30 segments usually found in this order. Adult males in this family feature a reduced leg pair 10 in addition to the two pairs normally modified into gonopods in this order. In the adult male of the species Hoffmaneuma exiguum, for example, the gonopod complex derives from all three leg pairs rather than from just the usual two.

Trachygonidae is a family of millipedes belonging to the order Chordeumatida. Adult millipedes in this family have 28 or 30 segments.

Opisthocheiridae is a family of millipedes belonging to the order Chordeumatida. These millipedes range from 5 mm to 16 mm in length and are found from Belgium to Morocco. Adult millipedes in this family have 26 or 30 segments. This family includes the cave-dwelling species Opisthocheiron canayerensis, notable as one of few chordeumatidan species with only 26 segments. The adult female of this species has only 42 pairs of legs, and the adult male has only 40 leg pairs, as one would expect for adult chordeumatidans with four fewer segments than typically found in this order.

Glomeridellidae is a family of millipedes belonging to the order Glomerida. The adult males in this family have two pairs of telopods rather than the single pair found in other families in this order.

References

  1. synonym: Strongylosomatidea Brölemann, 1916
  1. Robert Mesibov (2005). "A new genus of millipede (Diplopoda: Polydesmida: Dalodesmidae) from Tasmania with a pseudo-articulated gonopod telopodite" (PDF). Zootaxa . 1064: 39–49. doi:10.11646/zootaxa.1064.1.4.
  2. 1 2 Shear, W (2011). Zhang, Z.-Q. (ed.). Class Diplopoda de Blainville in Gervais, 1844. In: Animal biodiversity : an outline of higher-level classification and survey of taxonomic richness (PDF). Zootaxa. pp. 159–164. ISBN   978-1-86977-850-7.
  3. Thomas Eisner (2005). "Vinegaroons and other wizards". For Love of Insects. Harvard University Press. pp. 44–73. ISBN   978-0-674-01827-3.
  4. 1 2 3 4 5 6 7 8 9 10 Enghoff, Henrik; Dohle, Wolfgang; Blower, J. Gordon (1993). "Anamorphosis in Millipedes (Diplopoda) — The Present State of Knowledge with Some Developmental and Phylogenetic Considerations". Zoological Journal of the Linnean Society. 109 (2): 103–234. doi:10.1111/j.1096-3642.1993.tb00305.x.
  5. Fusco, Giuseppe (December 2005). "Trunk segment numbers and sequential segmentation in myriapods". Evolution & Development. 7 (6): 608–617. doi:10.1111/j.1525-142X.2005.05064.x. PMID   16336414. S2CID   21401688 . Retrieved 25 August 2020.
  6. Henen, Derek; Brown, Jeff. Millipedes of Ohio (PDF). Ohio Division of Wildlife. pp. 3, 44.
  7. 1 2 Colin Little (1983). "Onychophorans and myriapods". The Colonisation of Land: Origins and Adaptations of Terrestrial Animals. Cambridge University Press. pp. 127–145. ISBN   978-0-521-25218-8.
  8. 1 2 3 4 Blower, J. Gordon (1985). Millipedes : keys and notes for the identification of the species. Linnean Society of London, Estuarine and Brackish-water Sciences Association. London: Published for the Linnean Society of London and the Estuarine and Brackish-Water Sciences Association by E.J. Brill. ISBN   90-04-07698-0. OCLC   13439686.
  9. William H. Robinson (2005). "Other arthropods in the urban environment". Handbook of Urban Insects and Arachnids. Cambridge University Press. pp. 389–440. ISBN   978-0-521-81253-5.
  10. 1 2 Shelley, Rowland M. (1999). "Centipedes and Millipedes with Emphasis on North American Fauna". The Kansas School Naturalist. 45 (3): 1–16. Archived from the original on 2016-11-12. Retrieved 2013-10-14.
  11. 1 2 3 4 5 6 Mesibov, Robert. "External Anatomy of Polydesmida: Body plans". myriapodology.org. Retrieved 2022-02-20.
  12. Loomis, H. F.; Hoffman, Richard L. (1962). "A remarkable new family of spined polydesmoid Diplopoda, including a species lacking gonopods in the male sex". Proceedings of the Biological Society of Washington. 75: 145–158. ISSN   0006-324X via Biodiversity Heritage Library.
  13. 1 2 Enghoff, Henrik; Golovatch, Sergei; Short, Megan; Stoev, Pavel; Wesener, Thomas (2015-01-01). "Diplopoda — taxonomic overview". Treatise on Zoology - Anatomy, Taxonomy, Biology. The Myriapoda, Volume 2: 363–453. doi:10.1163/9789004188273_017. ISBN   9789004188273.
  14. VandenSpiegel, Didier; Golovatch, Sergei I. (2015). "A new millipede of the family Ammodesmidae found in central Africa (Diplopoda, Polydesmida)". ZooKeys (483): 1–7. doi: 10.3897/zookeys.483.9150 . PMC   4351444 . PMID   25755621.
  15. VandenSpiegel, Didier; Golovatch, Sergei (2012). "The millipede family Ammodesmidae (Diplopoda, Polydesmida) in western Africa". ZooKeys (221): 1–17. doi: 10.3897/zookeys.221.3739 . PMC   3487632 . PMID   23129978.
  16. Brölemann, H.-W. (1902). "Myriapodes Cavericoles". Annales de la Société Entomologique de France (in French). 71: 448–460 via Biodiversity Heritage Library.
  17. 1 2 Minelli, Alessandro (2015-01-01). "Diplopoda — development". Treatise on Zoology - Anatomy, Taxonomy, Biology. The Myriapoda, Volume 2: 267–302. doi:10.1163/9789004188273_012. ISBN   9789004188273.
  18. Shear, William A.; Ferreira, Rodrigo Lopes; Iniesta, Luiz Felipe Moretti; Marek, Paul (2016-10-25). "A millipede missing link: Dobrodesmidae, a remarkable new polydesmidan millipede family from Brazil with supernumerary rings (Diplopoda, Polydesmida), and the establishment of a new suborder Dobrodesmidea". Zootaxa. 4178 (3): 371–390. doi:10.11646/zootaxa.4178.3.4. ISSN   1175-5334. PMID   27811714 via ResearchGate.
  19. "Polydesmid Millipedes". The Australian Museum. Retrieved 2022-09-11.