Mite

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Mites
Temporal range: Early Devonian  – Present, 410–0  Ma
Trombidium holosericeum (aka).jpg
Trombidium holosericeum mite (Acariformes)
Varroa destructor, 1 2019-09-06-19.12.07 ZS PMax UDR (48697155713).jpg
Varroa destructor (Parasitiformes)
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Class: Arachnida
Mites are found in two superorders

Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari. However, most recent genetic analyses do not recover the two as each other's closest relative within Arachnida, rendering the group non-monophyletic. Most mites are tiny, less than 1 mm (0.04 in) in length, and have a simple, unsegmented body plan. The small size of most species makes them easily overlooked; some species live in water, many live in soil as decomposers, others live on plants, sometimes creating galls, while others are predators or parasites. This last type includes the commercially destructive Varroa parasite of honey bees, as well as scabies mites of humans. Most species are harmless to humans, but a few are associated with allergies or may transmit diseases.

The scientific discipline devoted to the study of mites is called acarology.

Evolution and taxonomy

The microscopic mite Lorryia formosa (Tydeidae) Yellow mite (Tydeidae) Lorryia formosa 2 edit.jpg
The microscopic mite Lorryia formosa (Tydeidae)

The mites are not a defined taxon, but is used for two distinct groups of arachnids, the Acariformes and the Parasitiformes. The phylogeny of the Acari has been relatively little studied, but molecular information from ribosomal DNA is being extensively used to understand relationships between groups. The 18 S rRNA gene provides information on relationships among phyla and superphyla, while the ITS2, and the 18S ribosomal RNA and 28S ribosomal RNA genes, provide clues at deeper levels. [1]

Taxonomy

Fossil record

Mite, cf Glaesacarus rhombeus, fossilised in Baltic amber, Upper Eocene Milbe cf Glaesacarus rhombeus.jpg
Mite, cf Glaesacarus rhombeus, fossilised in Baltic amber, Upper Eocene

The mite fossil record is sparse, due to their small size and low preservation potential. [5] The oldest fossils of acariform mites are from the Rhynie Chert, Scotland, which dates to the early Devonian, around 410 million years ago [6] [5] while the earliest fossils of Parasitiformes are known from amber specimens dating to the mid-Cretaceous, around 100 million years ago. [5] [7] Most fossil acarids are no older than the Tertiary (up to 65 mya). [8]

Phylogeny

Members of the superorders Opilioacariformes and Acariformes (sometimes known as Actinotrichida) are mites, as well as some of the Parasitiformes (sometimes known as Anactinotrichida). [9] Recent genetic research has suggested that Acari is polyphyletic (of multiple origins). [10] [11] [12] [13] A study using molecular data from the mitochondria and nucleus recovered Acariformes as sister to the Solifugae (camel spiders) and Parasitiformes as sister to the Pseudoscorpionida, with other arachnid orders separating these two groupings on the phylogenetic tree, as shown below. [10]

Arachnida

Palpigradi

Pseudoscorpionida

False scorpions Neobisium sylvaticum 03.png

Parasitiformes

Ixodida (ticks) Ixodes scapularis P1170301a (white background).png

Parasitic mites, inc. Varroa Varroa destructor, 1 2019-09-06-19.12.07 ZS PMax UDR (48697155713) (white background).jpg

Acariformes

Trombidiformes (chiggers, velvet mites, etc) Rote Samtmilbe Namibia.png  

Sarcoptiformes (dust & fur mites, etc) Galumnidae sp.png

Solifugae

Camel spiders Ammotrecha itzaana 4414721993.png

other Arachnids including spiders and scorpions

"Acari"
(mites and ticks)

However, a few phylogenomic studies have found strong support for monophyly of Acari and a sister relationship between Acariformes and Parasitiformes, [14] [15] although this finding has been questioned, with other studies suggesting that this likely represents a long branch attraction artefact. [12]

Anatomy

External

Mites are tiny members of the class Arachnida; most are in the size range 250 to 750 μm (0.01 to 0.03 in) but some are larger and some are no bigger than 100 μm (0.004 in) as adults. The body plan has two regions, a cephalothorax (with no separate head) or prosoma, and an opisthosoma or abdomen. Segmentation has almost entirely been lost and the prosoma and opisthosoma are fused, only the positioning of the limbs indicating the location of the segments. [16]

1 Chelicerae, 2 Palps, 3 Salivary glands, 4 Gut, 5 Excretory (Malpighian) tubules, 6 Anus, 7 Ovary or testes, 8 Air-breathing tubes (tracheae), 9 Central ganglion, 10 Legs, 11 Hypostome. Acarine anatomy and morphology.png
1 Chelicerae, 2 Palps, 3 Salivary glands, 4 Gut, 5 Excretory (Malpighian) tubules, 6 Anus, 7 Ovary or testes, 8 Air-breathing tubes (tracheae), 9 Central ganglion, 10 Legs, 11 Hypostome.

At the front of the body is the gnathosoma or capitulum. This is not a head and does not contain the eyes or the brain, but is a retractable feeding apparatus consisting of the chelicerae, the pedipalps and the oral cavity. It is covered above by an extension of the body carapace and is connected to the body by a flexible section of cuticle. Two-segmented chelicerae is the ancestral condition in Acariformes, but in more derived groups they are single-segmented. And three-segmented chelicerae is the ancestral condition in Parasitiformes, but has been reduced to just two segments in more derived groups. [18] The pedipalps differ between taxa depending on diet; in some species the appendages resemble legs while in others they are modified into chelicerae-like structures. The oral cavity connects posteriorly to the mouth and pharynx. [16]

Most mites have four pairs of legs (two pairs in Eriophyoidea [19] ), each with six segments, which may be modified for swimming or other purposes. The dorsal surface of the body is clad in hardened tergites and the ventral surface by hardened sclerites; sometimes these form transverse ridges. The gonopore (genital opening) is located on the ventral surface between the fourth pair of legs. Some species have one to five median or lateral eyes but many species are blind, and slit and pit sense organs are common. Both body and limbs bear setae (bristles) which may be simple, flattened, club-shaped or sensory. Mites are usually some shade of brown, but some species are red, orange, black or green, or some combination of these colours. [16]

Many mites have stigmata (openings used in respiration). In some mites, the stigmata are associated with peritremes: paired, tubular, elaborated extensions of the tracheal system. The higher taxa of mites are defined by these structures: [20] [21] [22]

Internal

Mite digestive systems have salivary glands that open into the preoral space rather than the foregut. Most species carry two to six pairs of salivary glands that empty at various points into the subcheliceral space. [26] A few mite species lack an anus: they do not defecate during their short lives. [27] The circulatory system consists of a network of sinuses and most mites lacks a heart, movement of fluid being driven by the contraction of body muscles. But ticks, and some of the larger species of mites, have a dorsal, longitudinal heart. [28] Gas exchange is carried out across the body surface, but many species additionally have between one and four pairs of tracheae. The excretory system includes a nephridium and one or two pairs of Malpighian tubules. [29] Several families of mites, such as Tetranychidae, Eriophyidae, Camerobiidae, Cunaxidae, Trombidiidae, Trombiculidae, Erythraeidae and Bdellidae have silk glands used to produce silk for various purposes. Additionally, water mites (Hydrachnidia) produce long thin threads that may be silk. [30]

Reproduction and life cycle

Harvest mite (Trombiculidae) life cycle: the larvae and nymphs resemble small adults, though the larvae have only six legs. Harvest mite cycle.svg
Harvest mite (Trombiculidae) life cycle: the larvae and nymphs resemble small adults, though the larvae have only six legs.

The sexes are separate in mites; males have a pair of testes in the mid-region of the body, each connected to the gonopore by a vas deferens, and in some species there is a chitinous penis; females have a single ovary connected to the gonopore by an oviduct, as well as a seminal receptacle for the storage of sperm. In most mites, sperm is transferred to the female indirectly; the male either deposits a spermatophore on a surface from which it is picked up by the female, or he uses his chelicerae or third pair of legs to insert it into the female's gonopore. In some of the Acariformes, insemination is direct using the male's penis. [16] The spermatophora in all mites are aflagellate. [31]

The eggs are laid in the substrate, or wherever the mite happens to live. They take up to six weeks to hatch, according to species, with the next stage being the six-legged larvae. After three moults, the larvae become nymphs, [32] with eight legs, and after a further three moults, they become adults. Longevity varies between species, but the lifespan of mites is short compared to many other arachnids. [16]

Ecology

Niches

Russet mite, A. anthocoptes, is found on the invasive weed Cirsium arvense, the Canada thistle, across the world. It may be usable as a biological pest control agent for this weed. Rust Mite, Aceria anthocoptes.jpg
Russet mite, A. anthocoptes , is found on the invasive weed Cirsium arvense , the Canada thistle, across the world. It may be usable as a biological pest control agent for this weed.

Mites occupy a wide range of ecological niches. For example, Oribatida mites are important decomposers in many habitats. They eat a wide variety of material including living and dead plant and fungal material, lichens and carrion; some are predatory, though no oribatid mites are parasitic. [34] Mites are among the most diverse and successful of all invertebrate groups. They have exploited a wide array of habitats, and because of their small size go largely unnoticed. They are found in freshwater (e.g. the water mites or Hydrachnidia [35] ) and saltwater (most Halacaridae [36] ), in the soil, in forests, pastures, agricultural crops, ornamental plants, thermal springs and caves. They inhabit organic debris of all kinds and are extremely numerous in leaf litter. They feed on animals, plants and fungi and some are parasites of plants and animals. [37] Some 48,200 species of mites have been described, [38] but there may be a million or more species as yet undescribed. [16] The tropical species Archegozetes longisetosus is one of the strongest animals in the world, relative to its mass (100 μg): It lifts up to 1,182 times its own weight, over five times more than would be expected of such a minute animal. [39] A mite also holds a speed record: for its length, Paratarsotomus macropalpis is the fastest animal on Earth. [40]

The mites living in soil consist of a range of taxa. Oribatida and Prostigmata are more numerous in soil than Mesostigmata, and have more soil-dwelling species. [41] When soil is affected by an ecological disturbance such as agriculture, most mites (Astigmata, Mesostigmata and Prostigmata) recolonise it within a few months, whereas Oribatida take multiple years. [42]

Parasitism

Many mites are parasitic on plants and animals. One family of mites, Pyroglyphidae, or nest mites, live primarily in the nests of birds and other animals. These mites are largely parasitic and consume blood, skin and keratin. Dust mites, which feed mostly on dead skin and hair shed from humans instead of consuming them from the organism directly, evolved from these parasitic ancestors. [43] Ticks are a prominent group of mites that are parasitic on vertebrates, mostly mammal and birds, feeding on blood with specialised mouthparts. [44]

Parasitic mites sometimes infest insects. Varroa destructor attaches to the body of honey bees, and Acarapis woodi (family Tarsonemidae) lives in their tracheae. Hundreds of species are associated with other bees, mostly poorly described. They attach to bees in a variety of ways. For example, Trigona corvina workers have been found with mites attached to the outer face of their hind tibiae. [45] Some are thought to be parasites, while others are beneficial symbionts. Mites also parasitize some ant species, such as Eciton burchellii . [46] Most larvae of Parasitengona are ectoparasites of arthropods, while later life stages in this group tend to shift to being predators. [47]

Lime nail galls on Tilia x europaea, caused by the mite Eriophyes tiliae Eriophyes tiliae tiliae.JPG
Lime nail galls on Tilia × europaea , caused by the mite Eriophyes tiliae

Plant pests include the so-called spider mites (family Tetranychidae), thread-footed mites (family Tarsonemidae), and the gall mites (family Eriophyidae). [48] Among the species that attack animals are members of the sarcoptic mange mites (family Sarcoptidae), which burrow under the skin. Demodex mites (family Demodecidae) are parasites that live in or near the hair follicles of mammals, including humans. [49]

Dispersal

Being unable to fly, mites need some other means of dispersal. On a small scale, walking is used to access other suitable locations in the immediate vicinity. Some species mount to a high point and adopt a dispersal posture and get carried away by the wind, while others waft a thread of silk aloft to balloon to a new position. [50]

Parasitic mites use their hosts to disperse, and spread from host to host by direct contact. Another strategy is phoresy; the mite, often equipped with suitable claspers or suckers, grips onto an insect or other animal, and gets transported to another place. A phoretic mite is just a hitch-hiker and does not feed during the time it is carried by its temporary host. These travelling mites are mostly species that reproduce rapidly and are quick to colonise new habitats. [50]

Relationship with humans

Public health worker Stefania Lanzia using a scabies mite to publicise scabies, an often overlooked condition especially among the elderly Stefania Lanza and her soft toy Scabies Mite.jpg
Public health worker Stefania Lanzia using a scabies mite to publicise scabies, an often overlooked condition especially among the elderly

Mites are tiny and apart from those that are of economic concern to humans, little studied. The majority are beneficial, living in the soil or aqueous environments and assisting in the decomposition of decaying organic material, as part of the carbon cycle. [37]

Two species live on humans, namely Demodex folliculorum and Demodex brevis ; both are frequently referred to as eyelash mites.

Medical significance

The majority of mite species are harmless to humans and domestic animals, but a few species can colonize mammals directly, acting as vectors for disease transmission, and causing or contributing to allergenic diseases. Mites which colonize human skin are the cause of several types of itchy skin rashes, such as gamasoidosis, [51] rodent mite dermatitis, [52] grain itch, [53] grocer's itch, [53] and scabies; Sarcoptes scabiei is a parasitic mite responsible for scabies, which is one of the three most common skin disorders in children. [54] Demodex mites, which are common cause of mange in dogs and other domesticated animals, [49] have also been implicated in the human skin disease rosacea, although the mechanism by which demodex contributes to the disease is unclear. [55] Ticks are well known for carrying diseases, such as Lyme disease [56] and Rocky Mountain spotted fever. [57]

Mites and their eggs, drawn by Robert Hooke, Micrographia, 1665 Robert Hooke, Micrographia, mites; eggs Wellcome L0000180.jpg
Mites and their eggs, drawn by Robert Hooke, Micrographia , 1665

Chiggers are known primarily for their itchy bite, but they can also spread disease in some limited circumstances, such as scrub typhus. [58] The house-mouse mite is the only known vector of the disease rickettsialpox. [59] House dust mites, found in warm and humid places such as beds, cause several forms of allergic diseases, including hay fever, asthma and eczema, and are known to aggravate atopic dermatitis. [60]

Among domestic animals, sheep are affected by the mite Psoroptes ovis which lives on the skin, causing hypersensitivity and inflammation. [61] Hay mites are a suspected reservoir for scrapie, a prion disease of sheep. [62]

In beekeeping

The mite Varroa destructor is a serious pest of honey bees, contributing to colony collapse disorder in commercial hives. This organism is an obligate external parasite, able to reproduce only in bee colonies. It directly weakens its host by sucking up the bee's fat, and can spread RNA viruses including deformed wing virus. Heavy infestation causes the death of a colony, generally over the winter. Since 2006, more than 10 million beehives have been lost. [63] [64]

Biological pest control

Various mites prey on other invertebrates and can be used to control their populations. Phytoseiidae, especially members of Amblyseius , Metaseiulus , and Phytoseiulus , are used to control pests such as spider mites. [65] Among the Laelapidae, Gaeolaelaps aculeifer and Stratiolaelaps scimitus are used to control fungus gnats, poultry red mites and various soil pests. [66]

In culture

Mites were first observed under the microscope by the English polymath Robert Hooke. In his 1665 book Micrographia , he stated that far from being spontaneously generated from dirt, they were "very prettily shap'd Insects". [67] In 1898, Arthur Conan Doyle wrote a satirical poem, "A Parable", with the conceit of some cheese mites disputing the origin of the round cheddar cheese in which they all lived. [68] The world's first science documentary featured cheese mites, seen under the microscope; the short film was shown in London's Alhambra music hall in 1903, causing a boom in the sales of simple microscopes. [67]

See also

Related Research Articles

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

Arachnida is a class of joint-legged arthropods, in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegaroons.

<span class="mw-page-title-main">Tick</span> Order of arachnids in the arthropod phylum

Ticks are parasitic arachnids that are part of the mite superorder Parasitiformes. Adult ticks are approximately 3 to 5 mm in length depending on age, sex, species, and "fullness". Ticks are external parasites, living by feeding on the blood of mammals, birds, and sometimes reptiles and amphibians. The timing of the origin of ticks is uncertain, though the oldest known tick fossils are from the Cretaceous period, around 100 million years old. Ticks are widely distributed around the world, especially in warm, humid climates.

<span class="mw-page-title-main">Mange</span> Type of skin disease caused by parasitic mites

Mange is a type of skin disease caused by parasitic mites. Because various species of mites also infect plants, birds and reptiles, the term "mange", or colloquially "the mange", suggesting poor condition of the skin and fur due to the infection, is sometimes reserved for pathological mite-infestation of nonhuman mammals. Thus, mange includes mite-associated skin disease in domestic mammals, in livestock, and in wild mammals. Severe mange caused by mites has been observed in wild bears. Since mites belong to the arachnid subclass Acari, another term for mite infestation is acariasis.

<i>Varroa</i> Genus of mites

Varroa is a genus of parasitic mesostigmatan mites associated with honey bees, placed in its own family, Varroidae. The genus was named for Marcus Terentius Varro, a Roman scholar and beekeeper. The condition of a honeybee colony being infested with Varroa mites is called varroosis.

Acariasis is an infestation with mites.

<span class="mw-page-title-main">Acariformes</span> Superorder of mite

The Acariformes, also known as the Actinotrichida, are the more diverse of the two superorders of mites. Over 32,000 described species are found in 351 families, with an estimated total of 440,000 to 929,000 species, including undescribed species.

<span class="mw-page-title-main">Parasitiformes</span> Superorder of arachnids

Parasitiformes are a superorder of Arachnids, constituting one of the two major groups of mites, alongside Acariformes. Parasitiformes has, at times, been classified at the rank of order or suborder.

<span class="mw-page-title-main">Opilioacaridae</span> Order of mites

Opilioacaridae is the sole family of mites in the order Opilioacarida, made up of about 13 genera. The mites of this family are rare, large mites, and are widely considered primitive, as they retain six pairs of eyes, and abdominal segmentation. They have historically been considered separate from other mites belonging to Acariformes and Parasitiformes, but are now generally considered a subgroup of Parasitiformes based on molecular phylogenetics.

<span class="mw-page-title-main">Oribatida</span> Order of mites

Oribatida, also known as oribatid mites, moss mites or beetle mites, are an order of mites, in the "chewing Acariformes" clade Sarcoptiformes. They range in size from 0.2 to 1.4 millimetres. There are currently 12,000 species that have been identified, but researchers estimate that there may be anywhere from 60,000 to 120,000 total species. Oribatid mites are by far the most prevalent of all arthropods in forest soils, and are essential for breaking down organic detritus and distributing fungi.

<span class="mw-page-title-main">Astigmatina</span> Group of mites

Astigmatina is a clade of mites in the superorder Acariformes. Astigmata has been ranked as an order or suborder in the past, but was lowered to the unranked clade Astigmatina of the clade Desmonomatides in the order Sarcoptiformes. Astigmatina is now made up of the two groups Acaridia and Psoroptidia, which have been suborders of the order Astigmata in the past. Astigmatina contains about 10 superfamilies and 76 families under Acaridia and Psoroptidia.

<span class="mw-page-title-main">Eriophyoidea</span> Superfamily of mites

Eriophyoidea are a superfamily of herbivorous mites. All post-embryonic instars lack the third and fourth pairs of legs, and the respiratory system is also absent.

<span class="mw-page-title-main">Prostigmata</span> Suborder of mites

Prostigmata is a suborder of mites belonging to the order Trombidiformes, which contains the "sucking" members of the "true mites" (Acariformes).

<i>Ornithodoros</i> Genus of arachnids in the soft-bodied tick family, Argasidae.

Ornithodoros is a genus in the soft-bodied tick family, Argasidae.

<span class="mw-page-title-main">Hydrachnidia</span> Group of mites

Hydrachnidia, also known as "water mites", Hydrachnidiae, Hydracarina or Hydrachnellae, are among the most abundant and diverse groups of benthic arthropods, composed of 6,000 described species from 57 families. As water mites of Africa, Asia, and South America have not been well-studied, the numbers are likely to be far greater. Other taxa of parasitengone mites include species with semi-aquatic habits, but only the Hydracarina are properly subaquatic. Water mites follow the general Parasitengona life cycle: active larva, inactive (calyptostasic) protonymph, active deutonymph, inactive tritonymph and active adult. Usually, larvae are parasites, while deutonymphs and adults are predators.

<i>Histiostoma</i> Genus of mites

Histiostoma is a genus of mites in the family Histiostomatidae.

<span class="mw-page-title-main">Dermanyssoidea</span> Superfamily of mites

Dermanyssoidea is a superfamily of mites, including most of the mites which parasitise vertebrates.

<span class="mw-page-title-main">Mites of domestic animals</span> Type of parasite of domestic animals

Mites that infest and parasitize domestic animals cause disease and loss of production. Mites are small invertebrates, most of which are free living but some are parasitic. Mites are similar to ticks and both comprise the order Acari in the phylum Arthropoda. Mites are highly varied and their classification is complex; a simple grouping is used in this introductory article. Vernacular terms to describe diseases caused by mites include scab, mange, and scabies. Mites and ticks have substantially different biology from, and are classed separately from, insects. Mites of domestic animals cause important types of skin disease, and some mites infest other organs. Diagnosis of mite infestations can be difficult because of the small size of most mites, but understanding how mites are adapted to feed within the structure of the skin is useful.

<span class="mw-page-title-main">Mites of livestock</span> Small crawling animals related to ticks and spiders

Mites are small crawling animals related to ticks and spiders. Most mites are free-living and harmless. Other mites are parasitic, and those that infest livestock animals cause many diseases that are widespread, reduce production and profit for farmers, and are expensive to control.

References

  1. ↑ Dhooria MS (2016). "Molecular Biology and Acarology". Fundamentals of Applied Acarology. Springer. p. 176. ISBN   978-981-10-1594-6.
  2. ↑ Beaulieu, Frédéric (2011). Zhang, Zhi-Qiang (ed.). "Superorder Parasitiformes: In: Zhang, Z-Q. (ed.) Animal biodiversity: an outline of higher-level classification and survey of taxonomic richness". Zootaxa. 3148. doi:10.11646/zootaxa.3148.1.23. ISBN   978-1-86977-849-1. ISSN   1175-5326.
  3. ↑ Ballesteros JA, Santibáñez López CE, Kováč Ľ, Gavish-Regev E, Sharma PP (December 2019). "Ordered phylogenomic subsampling enables diagnosis of systematic errors in the placement of the enigmatic arachnid order Palpigradi". Proceedings. Biological Sciences. 286 (1917): 20192426. doi:10.1098/rspb.2019.2426. PMC   6939912 . PMID   31847768.
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  6. ↑ Dunlop JA, Garwood RJ (February 2018). "Terrestrial invertebrates in the Rhynie chert ecosystem". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 373 (1739): 20160493. doi:10.1098/rstb.2016.0493. PMC   5745329 . PMID   29254958.
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  8. ↑ de la Fuente J (2003). "The fossil record and the origin of ticks (Acari: Parasitiformes: Ixodida)". Experimental & Applied Acarology. 29 (3–4): 331–344. doi:10.1023/A:1025824702816. PMID   14635818. S2CID   11271627.
  9. ↑ Walter DE, Krantz G, Lindquist E (13 December 1996). "Acari: The mites". Tree of Life Web Project. Retrieved 6 October 2017.
  10. 1 2 Dabert M, Witalinski W, Kazmierski A, Olszanowski Z, Dabert J (July 2010). "Molecular phylogeny of acariform mites (Acari, Arachnida): strong conflict between phylogenetic signal and long-branch attraction artifacts". Molecular Phylogenetics and Evolution. 56 (1): 222–241. doi:10.1016/j.ympev.2009.12.020. PMID   20060051.
  11. ↑ Sanggaard KW, Bechsgaard JS, Fang X, Duan J, Dyrlund TF, Gupta V, et al. (May 2014). "Spider genomes provide insight into composition and evolution of venom and silk". Nature Communications. 5: 3765. Bibcode:2014NatCo...5.3765S. doi:10.1038/ncomms4765. PMC   4273655 . PMID   24801114.
  12. 1 2 Ballesteros JA, Santibáñez-López CE, Baker CM, Benavides LR, Cunha TJ, Gainett G, et al. (February 2022). Teeling E (ed.). "Comprehensive Species Sampling and Sophisticated Algorithmic Approaches Refute the Monophyly of Arachnida". Molecular Biology and Evolution. 39 (2). doi:10.1093/molbev/msac021. PMC   8845124 . PMID   35137183.
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