Gonopod

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One of two gonopods of the greenhouse millipede (Oxidus gracilis). Scale bar: 0.2 mm or approximately 1/127 in Oxidus gracilis gonopod SEM.png
One of two gonopods of the greenhouse millipede (Oxidus gracilis). Scale bar: 0.2 mm or approximately 1/127 in

Gonopods are specialized appendages of various arthropods used in reproduction or egg-laying. In males, they facilitate the transfer of sperm from male to female during mating, and thus are a type of intromittent organ. In crustaceans and millipedes, gonopods are modified walking or swimming legs. Gonopods may be highly decorated with elaborate structures which may play roles in sperm competition, and can be used to differentiate and identify closely related species. Gonopods generally occur in one or more pairs, as opposed to the single (un-paired) reproductive organs such as the aedeagus of insects or the penis of harvestmen.

Contents

Insects

Paired gonopods (g) on the male genitalia of Corydalus cornutus Corydalus cornutus genitalia.png
Paired gonopods (g) on the male genitalia of Corydalus cornutus

In insects, gonopods are appendages of the genital segment that may be used in insemination, or that comprise the egg-laying apparatus. [1] [2]

Crustaceans

In male decapod crustaceans, gonopods are modified swimming appendages (pleopods). The anterior two pair of pleopods in males are modified for sperm transferring, with differing degree of morphological diversification. [3]

Millipedes

Gonopod diversity in 20 Chaleponcus species (Spirostreptida, Odontopygidae) from Tanzania Chaleponcus gonopods.png
Gonopod diversity in 20 Chaleponcus species (Spirostreptida, Odontopygidae) from Tanzania

In millipedes, gonopods consist of one or two pairs of often highly modified walking legs in mature males, and are primarily found in members of the subgroup Helminthomorpha—containing most orders and the vast majority of species—where they are located on the seventh body segment consisting of leg pairs 8 and/or 9. [4] Males of the subgroup Pentazonia (which includes the Oniscomorpha (pill millipedes) and Glomeridesmida) lack gonopods but possess enlarged appendages known as telopods at the rear of the body used to firmly hold females during mating. [5] The complex structure of gonopods is a primary method of distinguishing closely related species of millipede, [6] although the terminology used to describe the same structures may vary between authors. [7] The complex morphology of millipede gonopods may be driven by sperm competition or other forms of sexual selection, with some structures serving to scoop out or displace sperm of other males, and others acting to stimulate females into becoming sexually receptive. [5]

Millipede gonopods do not produce sperm directly, but rather gather sperm produced from a gonopore on the base of the third body segment. [8]

Gonopods develop gradually during the growth of an individual. In early developmental stages, all legs are of the walking type, and cannot be used to determine sex. Through successive molts, the walking legs metamorphose into mature gonopods. [8]

Gonopod development in Nopoiulus kochii (Julida, Blaniulidae). A: juvenile with walking legs on segment 7 (asterisk). B: subsequent stage with walking legs replaced by gonopod primordia. C: adult stage with gonopods (colored) in place of walking legs. Non-systemic metamorphosis in millipedes- Nopoiulus kochii.png
Gonopod development in Nopoiulus kochii (Julida, Blaniulidae). A: juvenile with walking legs on segment 7 (asterisk). B: subsequent stage with walking legs replaced by gonopod primordia. C: adult stage with gonopods (colored) in place of walking legs.
OrderNumber of pairs of
gonopods/telopods
Leg pair(s) modified! [a] Notes
LP = Leg pair
Image [b]
Callipodida 18thLP9 is reduced
Sinocallipus steineri Sinocallipus steineri gonopods.jpg
Sinocallipus steineri
Chordeumatida 28th & 9thGonopods and other accessory mating legs vary within the four suborders: In Chordeumatidea LP9 sperm-transferring, LP10 is reduced, and LP11 possess coxal glands; in Heterochordeumatidea LP9 sperm-transferring, LP10 reduced, and both LP10 and LP11 with coxal gland; in the Craspedosomatidea and Striariidea, LP8 sperm-transferring, LP9 modified, LP10 and LP11 with coxal glands
Glomerida 2 (telopods)Not involved with sperm transfer, only clasping females
Glomeris troglokabyliana walking legs (A, B) and telopod (C) Glomeris troglokabyliana telopods.png
Glomeris troglokabyliana walking legs (A, B) and telopod (C)
Glomeridesmida 1 (telopods)
Julida 28th & 9thLP9 sperm-transferring
Ommatoiulus chambiensis right anterior (1) and posterior (2) Ommatoiulus chambiensis, gonopod.png
Ommatoiulus chambiensis right anterior (1) and posterior (2)
Platydesmida 29th & 10thLeg-like; LP 9 sperm-transferring
Polydesmida 18th
Desmoxytes lingulata Desmoxytes lingulata gonopods.jpg
Desmoxytes lingulata
Polyxenida 0Indirect fertilization
Polyzoniida 29th & 10thleg-like; LP 9 sperm-transferring
Siphoniulida 18th
Siphonocryptida 29th & 10thleg-like, LP 9 sperm-transferring
Siphonophorida 29th & 10thleg-like; LP 9 sperm-transferring
Illacme plenipes, arrow pointing to anterior gonopods Illacme plenipes gonopods in situ.jpg
Illacme plenipes , arrow pointing to anterior gonopods
Sphaerotheriida 2 (telopods)
Sphaeromimus vatovavy posterior telopods Sphaeromimus vatovavy posterior telopods.jpg
Sphaeromimus vatovavy posterior telopods
Spirobolida 28th & 9thLP9 sperm-transferring, LP8 fused into single structure.
Aphistogoniulus jeekeli anterior (A) and left posterior (B) Aphistogoniulus jeekeli holotype gonopods 03.png
Aphistogoniulus jeekeli anterior (A) and left posterior (B)
Spirostreptida 28th & 9thLP8 sperm-transferring
Stemmiulida 18thLP8 sperm-transferring; LP9 reduced
  1. counting from head
  2. not necessarily representative of order

See also

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.

<span class="mw-page-title-main">Appendage</span> External body part or natural prolongation, that protrudes from an organisms body

An appendage is an external body part or natural prolongation that protrudes from an organism's body such as an arm or a leg. Protrusions from single-celled bacteria and archaea are known as cell-surface appendages or surface appendages. In many kinds of eukaryotic cell the protrusions are known as membrane protrusions or cell appendages.

<span class="mw-page-title-main">Decapod anatomy</span> Entire structure of a decapod crustacean

The decapod is made up of 20 body segments grouped into two main body parts: the cephalothorax and the pleon (abdomen). Each segment may possess one pair of appendages, although in various groups these may be reduced or missing. They are, from head to tail:

The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments are of Latin origin, and may be confused with terms for bones: coxa, trochanter, femur, tibia, tarsus, ischium, metatarsus, carpus, dactylus, patella.

<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. 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">Spirobolida</span> Order of millipedes

Spirobolida is an order of "round-backed" millipedes containing approximately 500 species in 12 families. Its members are distinguished by the presence of a "pronounced suture that runs "vertically down the front of the head". Most of the species live in the tropics, and many are brightly coloured. Mature males have two pairs of modified legs, the gonopods, consisting of the 8th and 9th leg pair: the posterior gonopods are used in sperm-transfer while the anterior gonopods are fused into a single plate-like structure.

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

Glomerida is an order of pill-millipedes found primarily in the Northern Hemisphere. Also known as northern pill millipedes, they superficially resemble pill-bugs or woodlice, and can enroll into a protective ball. They have twelve body segments, 17 to 19 pairs of legs, and males have enlarged rear legs involved in mating. The order includes about 30 genera and at least 280 species, including Glomeris marginata, the common European pill-millipede. The order contains members in Europe, South-east Asia and the Americas from California to Guatemala. Although historically considered closely related with the similar sphaerotheriidans that also enroll, some DNA evidence suggest they may be more closely related to glomeridesmidans, a poorly known order that does not enroll.

<i>Mammamia</i> Genus of millipedes

Mammamia profuga is a species of cave-dwelling millipede in the family Julidae. The only known species of the genus Mammamia, it was described in 2011 from a specimen discovered in a cave in Italy.

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

Polydesmida is the largest order of millipedes, with more than 5,000 species, including all the millipedes reported to produce hydrogen cyanide (HCN). This order is also the most diverse of the millipede orders in terms of morphology. Millipedes in this order are found in all regions of the world other than Antarctica.

<i>Dromia personata</i> Species of crab

Dromia personata, also known as the sponge crab or sleepy crab, is a species of crab found in the North Sea, the Mediterranean Sea, and connecting parts of the northeastern Atlantic Ocean. Like most other epibenthic crustaceans, the biomass of this species is especially dense in the Mediterranean continental shelf. It mainly resides from the lower shore to a depth of 50 meters (164 ft), often in caves. Occasionally, they are found living in depths as low as 110 meters (360 ft). They serve as prey for octopus, starfish, and other fish. Their last two pairs of legs are positioned dorsally, and are used to hold a sponge in place as camouflage.

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

Xystodesmidae is a family of millipedes in the suborder Leptodesmidea within the order Polydesmida. The family Xystodesmidae was created by the American biologist Orator F. Cook in 1895 and named after the genus Xystodesmus. This family includes more than 390 known species distributed among 62 genera. Many species, however, remain undescribed: 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.

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

Polyxenida is an order of millipedes readily distinguished by a unique body plan consisting of a soft, non-calcified body ornamented with tufts of bristles. These features have inspired the common names bristly millipedes or pincushion millipedes. This order includes about 148 species in four families worldwide, which represent the only living members of the subclass Penicillata.

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

Chordeumatida is a large order of millipedes containing more than 1,400 species. Also known as sausage millipedes, they are found nearly worldwide. Chordeumatida is the largest order in the superorder Nematophora, a group also known as spinning millipedes because their telsons feature spinnerets used to build nests of silk. These millipedes produce this silk to create chambers in which to molt or to lay their eggs.

<i>Brachycybe</i> Genus of millipedes

Brachycybe (Greek for "short head") is a genus of social millipedes with species in the United States and East Asia. Species in this genus are often referred to as "feather millipedes" due to their shape.

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

Haplodesmidae is a family of millipedes in the order Polydesmida. This family includes about 70 species. Species occur in East Asia, Southeast Asia, and Oceania, although some species have been introduced to the New world tropics.

<span class="mw-page-title-main">Juliformia</span> Suborder of millipedes

Juliformia is a taxonomic superorder of millipedes containing three living orders: Julida, Spirobolida, and Spirostreptida, and the extinct group Xyloiuloidea known only from fossils.

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

Metopidiotrichidae is a family of millipedes in the order Chordeumatida. This family includes more than 70 species. These millipedes are found in Indochina, Australia, and on Pacific islands from New Zealand to Japan.

Agenodesmus reticulatus is a species of millipede in the family Fuhrmannodesmidae, which some authorities consider a junior synonym of Trichopolydesmidae. This millipede is among the very few species in the order Polydesmida to feature adults with only 18 segments rather than the 20 segments usually found in this order. This species is notable as the first polydesmidan millipede discovered with only 18 segments in adults, the smallest number recorded in the order Polydesmida. Before the discovery of A. reticulatus, polydesmidans were known to have only 19 or 20 segments in adults.

Prosopodesmus panporus is a species of flat-backed millipede in the family Haplodesmidae. Also known as the hothouse millipede, this species was first discovered in hothouses for tropical plants in England, where it is well established, but is probably native to Australia. This species features a unique distribution of ozopores, which appear on all segments with two pairs of legs. The species P. panporus is named for this complete series of ozopores and is the only species in the order Polydesmida with this distribution. This millipede is also notable for exhibiting sexual dimorphism in segment number: Whereas adult females of this species feature the 20 segments usually observed in the order Polydesmida, the adult males of this species feature only 19 segments.

References

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  3. Bauer, Raymond T. (1986). "Phylogenetic Trends in Sperm Transfer and Storage Complexity in Decapod Crustaceans" (PDF). Journal of Crustacean Biology. 6 (3): 313–325. Bibcode:1986JCBio...6..313B. doi:10.2307/1548173. JSTOR   1548173.
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  6. Mesibov, Robert. "Gonopods". External Anatomy of Polydesmida. Archived from the original on 9 July 2017. Retrieved 27 October 2013.
  7. Car, Catherine; Harvey, Mark (2013). "A review of the Western Australian keeled millipede genus Boreohesperus (Diplopoda, Polydesmida, Paradoxosomatidae)". ZooKeys (290): 1–19. Bibcode:2013ZooK..290....1C. doi: 10.3897/zookeys.290.5114 . PMC   3677291 . PMID   23794855.
  8. 1 2 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 . PMC   3170261 . PMID   21859471.