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Mites
Temporal range: Early Devonian – Present, 410–0 Ma
Trombidium holosericeum mite
Trombidium holosericeum mite (Acariformes)
Varroa destructor (Parasitiformes)
Varroa destructor (Parasitiformes)
Scientific classificationEdit this classification
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Class: Arachnida
Mites are found in two superorders

Mites are small arachnids (eight-legged arthropods) of two large orders, 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 invalid as a clade.[1] 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

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The microscopic mite Lorryia formosa (Tydeidae)

Mites are not a defined taxon, but the term 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.[2]

Taxonomy

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Fossil record

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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.[6] The oldest fossils of acariform mites are from the Rhynie Chert, Scotland, which dates to the early Devonian, around 410 million years ago[7][6] while the earliest fossils of Parasitiformes are known from amber specimens dating to the mid-Cretaceous, around 100 million years ago.[6][8] Most fossil acarids are no older than the Tertiary (up to 65 mya).[9]

Phylogeny

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Members of the superorders Opilioacariformes and Acariformes (sometimes known as Actinotrichida) are mites, as well as some of the Parasitiformes (sometimes known as Anactinotrichida).[10] Recent genetic research has suggested that Acari is polyphyletic (of multiple origins).[11][12][13][14]

Current understanding of probable chelicerate relationships, after Sharma and Gavish-Regev (2025):[1]

Chelicerata

Pycnogonida (sea spiders)

Prosomapoda

Opiliones (harvestmen)

Palpigradi (microwhip scorpions)

Solifugae (camel spiders)

Acariformes

Parasitiformes

Xiphosura (horseshoe crabs)

Ricinulei

Arachnopulmonata
Panscorpiones

Pseudoscorpiones (pseudoscorpions)

Scorpiones (scorpions)

Tetrapulmonata

Araneae (spiders)

Amblypygi (whip-spiders)

Schizomida (shorttailed whipscorpions)

Uropygi (whip scorpions/vinegaroons)

However, a few phylogenomic studies have found strong support for monophyly of Acari and a sister relationship between Acariformes and Parasitiformes,[15][16] although this finding has been questioned, with other studies suggesting that this likely represents a long branch attraction artefact as a result of inadequate sampling.[13][1]

Anatomy

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External

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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.[17]

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[18]

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.[19] 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.[17]

Most mites have four pairs of legs (two pairs in Eriophyoidea[20]), 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.[17]

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:[21][22][23]

  • Oribatida, formerly known as Cryptostigmata (crypto- = hidden), and Endeostigmata (endeo- = internal) lack primary stigmata and peritremes but may have secondary respiratory systems.[24] For example, oribatids in the suborder Brachypylina have stigmata on the ventral plate of the body that are difficult to see (thus the former name Cryptostigmata).[25]
  • Astigmata (a- = without) lack stigmata and respire through their cuticle.[26]
  • Prostigmata (pro- = before/in front) have stigmata at the front of the body, usually on the lateral margins or between the chelicerae. These are associated with peritremes that may be on the prodorsum near the cheliceral bases, or be horn-like and emergent, or form a line or network on the dorsum of the gnathosomal capsule.[22]
  • Opilioacaridae have four pairs of dorsolateral stigmata that are added sequentially during development.[22]
  • The other three orders of Parasitiformes, Holothyrida, Ixodida, and Mesostigmata (meso- = middle), have just one pair of stigmata in the region of the fourth pair of legs. They also have peritremes: in Ixodida these consist of paired encircling plates around the stigmata, while the peritremes in Mesostigmata and Holothyrida are grooves extending from the stigmata anteriorly (sometimes also posteriorly).[23]

Internal

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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.[27] A few mite species lack an anus: they do not defecate during their short lives.[28] The circulatory system consists of a network of sinuses and most mites lack a heart, with movement of fluid being driven by the contraction of body muscles. Ticks, and some of the larger species of mites, have a dorsal, longitudinal heart.[29] 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.[30] 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.[31]

Reproduction and life cycle

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Main article: Life stages of mites
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.[17] The spermatophora in all mites are aflagellate.[32]

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, then may pass through up to six instars: prelarva, larva, protonymph, deutonymph, tritonymph, and adult. These developmental stages may look different or may be omitted depending on the mite group. All mites have an adult stage.[33] Longevity varies between species, but the lifespan of mites is short compared to many other arachnids.[17]

Ecology

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Niches

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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.[34]

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.[35] 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[36]) and saltwater (most Halacaridae[37]), 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.[38] Some 48,200 species of mites have been described,[39] but there may be a million or more species as yet undescribed.[17] 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.[40] A mite also holds a speed record: for its length, Paratarsotomus macropalpis is the fastest animal on Earth.[41]

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.[42] 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.[43]

Parasitism

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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.[44] Ticks are a prominent group of mites that are parasitic on vertebrates, mostly mammal and birds, feeding on blood with specialised mouthparts.[45]

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.[46] Some are thought to be parasites, while others are beneficial symbionts. Mites also parasitize some ant species, such as Eciton burchellii.[47] Most larvae of Parasitengona are ectoparasites of arthropods, while later life stages in this group tend to shift to being predators.[48]

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).[49] 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.[50]

Dispersal

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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.[51]

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.[51]

Relationship with humans

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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.[38]

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

Medical significance

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Further information: Acariasis

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,[52] rodent mite dermatitis,[53] grain itch,[54] grocer's itch,[54] and scabies; Sarcoptes scabiei is a parasitic mite responsible for scabies, which is one of the three most common skin disorders in children.[55] Demodex mites, a common cause of mange in dogs and other domesticated animals,[50] have also been implicated in the human skin disease rosacea; although the mechanism by which demodex contributes to the disease is unclear.[56] Ticks are well known for carrying diseases, such as Lyme disease[57] and Rocky Mountain spotted fever.[58]

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.[59] The house-mouse mite is the only known vector of the disease rickettsialpox.[60] 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.[61]

Among domestic animals, sheep are affected by the mite Psoroptes ovis which lives on the skin, causing hypersensitivity and inflammation.[62] Hay mites are a suspected reservoir for scrapie, a prion disease of sheep.[63]

In beekeeping

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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.[64][65]

Biological pest control

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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.[66] Among the Laelapidae, Gaeolaelaps aculeifer and Stratiolaelaps scimitus are used to control fungus gnats, poultry red mites and various soil pests.[67]

In culture

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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".[68] 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.[69] 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.[68]

See also

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References

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[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Mite

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from Grokipedia
Mites are a diverse group of minute arachnids in the subclass Acari (class Arachnida), encompassing approximately 48,000 described species (as of 2025) that inhabit virtually every terrestrial and aquatic ecosystem on Earth, from soil and freshwater to marine depths and extreme environments like Antarctica. Typically measuring 0.08 to 1 mm in length—though some, like certain ticks and velvet mites, can reach 10–20 mm—they lack antennae and possess a body divided into a gnathosoma (containing mouthparts) and an idiosoma (bearing legs and sensory structures), with most species featuring eight legs in adults following a hexapod larval stage. Their life cycles generally include a hexapod larva and up to three octopod nymphal stages, enabling adaptations to free-living, parasitic, or predatory lifestyles. Traditionally classified into two main superorders—Acariformes (including groups like oribatid mites and astigmatans) and Parasitiformes (including mesostigmatans and ticks)—mites play crucial ecological roles as decomposers, predators of insects and nematodes, and indicators of and . In agriculture, species such as spider mites (Tetranychidae) act as pests damaging crops, while others, like predatory Phytoseiidae, serve as biological control agents. Medically and veterinarily, many mites and ticks transmit pathogens causing diseases like and , underscoring their significance in public health. Recent phylogenetic studies have challenged the of Acari, suggesting that "mites" form an artificial rather than natural taxonomic group, with more closely related to other chelicerates (such as horseshoe crabs) than to . This ongoing debate highlights the evolutionary complexity of these organisms, estimated to include 1 million or more undescribed species, representing a substantial portion of diversity.

Classification and Evolution

Taxonomy

Mites belong to the subclass Acari within the class Arachnida, phylum Arthropoda, kingdom Animalia. The Acari encompass a highly diverse group of arachnids, distinguished from other arachnids by the fusion of their abdominal segments and the incorporation of mouthparts into a forward-projecting . Traditionally, the subclass Acari is divided into two superorders: and . The , also known as Actinotrichida, represent the more speciose superorder and include three main orders: Trombidiformes (encompassing subgroups like , which includes plant-feeding spider mites), Sarcoptiformes (including , or oribatid mites, and Astigmatina, such as house dust mites), and Endeostigmata (primitive, soil-dwelling forms). The , or Anactinotrichida, comprise four orders: (predatory and parasitic mites), Ixodida (ticks), Opilioacarida (primitive, tropical forms), and Holothyrida (rare, glandular mites). These superorders account for over 55,000 described species worldwide, with estimates suggesting up to one million species in total, reflecting their extraordinary diversity across terrestrial, freshwater, and marine habitats. Taxonomic revisions in the , driven by molecular phylogenetic analyses, have refined the structure within by confirming its and establishing Endeostigmata as a distinct basal lineage within the order , separate from the derived and Astigmatina clades. This reclassification resolved long-standing debates on the position of Endeostigmata, previously considered potentially paraphyletic, based on 18S rDNA sequence data that highlighted conflicts with morphological long-branch attraction artifacts. Diagnostic traits for major taxa often center on the gnathosoma, the capitulum bearing the and palps. In , the gnathosoma is typically more robust and distinctly articulated to the idiosoma, with often adapted for piercing and sucking in parasitic forms like ticks. In contrast, exhibit a more integrated or variably fused gnathosoma, with frequently stylet-like for scraping or fluid-feeding, as seen in prostigmatans and oribatids. These structural differences, while not supporting overall Acari in recent studies, remain key for distinguishing the superorders.

Phylogenetic Relationships

Traditionally, the Acari are considered to occupy a monophyletic position within the class Arachnida, forming a major alongside other orders such as Araneae (spiders) and Scorpiones (scorpions), with earlier phylogenetic analyses based on 18S rRNA sequences and mitochondrial genomes supporting this placement and the overall unity of Arachnida. These molecular datasets indicated that Acari diverged early within the arachnid lineage, sharing derived traits like the absence of a on the pedipalps with other arachnids, while exhibiting unique adaptations in cheliceral morphology. However, recent phylogenomic studies as of 2025 have challenged the of Acari, suggesting that may be more closely related to (horseshoe crabs) than to , rendering Acari an artificial rather than natural group. Internally, the traditional phylogeny of Acari is characterized by a deep bifurcation between the superorders (encompassing diverse groups like sarcoptiform mites) and (including ticks and mesostigmatid mites), a division corroborated by older and mitochondrial protein-coding gene analyses. Within , Opilioacariformes emerges as the basal , serving as a critical outgroup to other parasitiform lineages and highlighting the group's evolutionary progression from primitive, soil-dwelling forms to more specialized parasitic taxa. Molecular clock analyses, calibrated using fossil constraints and relaxed clock models on multigene datasets, estimate the origin of Acari around 410–435 million years ago in the , aligning with the colonization of terrestrial environments by early arthropods. This timeline positions Acari as one of the oldest radiating lineages, with the Acariformes-Parasitiformes split occurring shortly thereafter, potentially in the late to . The monophyly of Acari has faced increasing challenges from molecular phylogenies, with recent 2025 studies using phylogenomic data and re-evaluating morphological traits like the gnathosoma and tritonymph stage as likely convergent rather than synapomorphic. These findings highlight an ongoing debate about the evolutionary unity of mites and ticks, with genomic-scale studies suggesting Acari may not form a natural clade distinct from other chelicerates.

Fossil Record

The fossil record of mites (Acari) is sparse compared to their extraordinary modern diversity, exceeding 60,000 described , primarily due to their minute size, which hinders preservation and discovery. The oldest putative evidence of mite-like arthropods dates to the in , approximately 410 million years ago, where five specimens of acariform mites have been identified among early terrestrial ecosystems. These fossils, preserved in silicified plant material, represent early soil-dwelling forms but lack sufficient detail for precise taxonomic placement, suggesting they are stem-group acarines rather than crown-group mites. The earliest unequivocal mite fossils appear in the Late Triassic amber from the Dolomite Alps of northeastern , dated to about 230 million years ago. These include two species of eriophyoid mites, Triasacarus fedelei and Ampezzoa triassica, preserved in droplets likely produced by extinct , indicating that gall-forming and sap-sucking behaviors were already established in early mite lineages. This discovery extends the confirmed record of Acari by over 100 million years beyond previous amber finds and highlights the role of in capturing small arthropods during the . Major fossil deposits of mites are predominantly from amber inclusions, providing exceptional preservation of soft-bodied features. In the mid-Cretaceous from (approximately 99 million years old), diverse are documented, including opilioacarids, mesostigmatids, and the oldest known ticks, such as Cornupalpiger baculatus, often found in association with feathered dinosaurs and early birds, revealing ancient host-parasite dynamics. Eocene (about 44 million years old) yields abundant Astigmata, including transitional forms like Levantoglyphus sidorchukae in the family Levantoglyphidae, which bridge free-living and parasitic lifestyles, alongside numerous phoretic specimens attached to . These deposits, spanning the to , document a radiation of mite subgroups but remain geographically biased toward amber-producing forests in and . Fossils provide key insights into early mite adaptations, particularly and phoresy. Ectoparasitic associations are evident in and Eocene ambers, where mites such as mesostigmatids are preserved attached to hosts like (Myrmozercon nataliae on a formicine ant) and flies, suggesting blood-feeding or tissue-damaging behaviors similar to modern ectoparasites. Phoretic associations, in which mites hitchhike on larger arthropods for dispersal, are frequently captured, as seen in with mites on dolichopodid flies and springtails on , indicating that this dispersal strategy predates the diversification of modern ecosystems and facilitated mite colonization of new habitats. Despite these glimpses, significant gaps persist in the mite fossil record, largely attributable to their small size—most under 1 mm—which reduces the likelihood of preservation in sedimentary rocks outside of . and early deposits are particularly underrepresented, with estimates suggesting that fewer than 1% of mite lineages are captured as s, compared to their dominance in contemporary soils and . This implies that the true evolutionary history of mites, including transitions to , is likely far more ancient and complex than currently evidenced.

Morphology and Anatomy

External Features

Mites exhibit a characteristic body structure divided into two primary tagmata: the gnathosoma and the idiosoma. The gnathosoma, often referred to as the capitulum, is the anterior, head-like region housing the mouthparts, including the paired for feeding and manipulation, and the pedipalps for sensory functions and grasping. The idiosoma forms the larger, oval to elongate posterior body, bearing the legs and dorsal and ventral shields, with the original segmentation obscured by fusion, resulting in an apparently unsegmented appearance. This division reflects the evolutionary reduction from more segmented arachnids, emphasizing compactness for diverse microhabitats. The appendages of mites are adapted for locomotion, attachment, and sensory , with adults typically possessing four pairs of legs attached to the idiosoma, while larvae bear only three pairs. Each consists of several segments—coxa, , , genu, , and tarsus—with setation patterns (arrangement and number of setae) serving as key diagnostic traits for identification across mite groups. The pretarsal structures at the tips vary, including empodia (fleshy, pad-like extensions for ) and ambulacra (paired claws or suckers for gripping surfaces), which enable mites to navigate smooth tissues or host skins effectively. The external of mites is a thin, flexible composed of epicuticle and procuticle layers, often marked by fine striations that provide extensibility and prevent cracking during movement or molting. In certain taxa, such as oribatid mites, the features hardened sclerites, including the prodorsal shield covering the anterior dorsum and the propodosomal shield over the leg-bearing region, which offer and . These sclerotized elements contrast with the softer, striated in more agile groups like prostigmatids. Mite sizes vary dramatically, from as small as 0.1 mm in eriophyid mites, which are worm-like and adapted for concealed plant feeding, to over 30 mm in fully engorged ticks, the largest acarine representatives. in external features is common, particularly in parasitic groups like ticks, where females often exhibit longer legs or expanded scuta for blood meal accommodation, while males display ornate patterns or shorter appendages for mate location.

Internal Anatomy

The digestive system of mites is divided into , , and regions, with the foregut including a muscular that facilitates ingestion through a pumping action driven by its thick-walled structure. The connects to the and opens into the midgut via an esophageal , allowing liquefied food to pass for further processing. The , the primary site of extracellular and , features epithelial cells that secrete enzymes and absorb nutrients, often organized into a ventriculus and caeca for efficient breakdown of ingested materials such as or host tissues. in mites is mainly accomplished by paired coxal glands located near the bases of the first and third pairs of legs, which function as osmoregulatory organs by filtering through a thin-walled sacculus to produce urine-like waste. Mites exhibit an open comprising a spacious hemocoel cavity that bathes internal organs in , with most species lacking a dedicated heart and relying on body muscle contractions to circulate the fluid. occurs primarily through paired that lead to a system of tracheae and tracheoles in many taxa, delivering oxygen directly to tissues, while smaller or aquatic mites often supplement this with across the thin . The in mites is typically dioecious, with females possessing paired ovaries that produce ova within a tubular leading to a and , often including spermathecae for storage near the genital opening. Males have paired testes connected by vasa deferentia to ejaculatory ducts, with varying by group; for instance, in gamasid mites, males use modified to transfer spermatophores externally before they are taken up by the female. The of mites forms a synganglion, a fused mass divided into supraesophageal and subesophageal regions encircling the , with the subesophageal portion including pedal ganglia that innervate the legs. Sensory input is processed through these ganglia, including connections to trichobothria—specialized setae that detect substrate vibrations and air movements, linking external sensory organs to the central nervous mass. In some species like those in Acaridae, the ventral retains distinct ganglionic aggregations for coordinated .

Life History

Reproduction

Mites in the subclass Acari are predominantly dioecious, with separate male and female sexes, though via occurs in several lineages. In sexual species, mating typically involves indirect sperm transfer, where males deposit stalked s on the substrate, which females actively retrieve using their genital structures. This is widespread across groups such as oribatids, water mites (Parasitengona), and gamasids, with males often using specialized cheliceral structures like the spermatodactyl to shape and position the spermatophore precisely. In prostigmatid mites, including eriophyoids, spermatophore deposition occurs without direct physical contact between sexes, reducing male-female interactions. Parthenogenesis is common in oribatid mites (), where thelytokous reproduction produces all-female offspring, enabling rapid population growth in stable soil environments and comprising up to 90% of individuals in some communities. In contrast, pest species like spider mites (Tetranychidae) exhibit arrhenotokous parthenogenesis, a form of where unfertilized eggs develop into haploid males and fertilized eggs into diploid females, allowing virgin females to initiate populations with male progeny. This reproductive mode is also present in some dermanyssine mites, facilitating colonization of new hosts. Female fertility in mites varies by and environmental conditions, with clutch sizes typically ranging from 1 to 100 eggs per female over their lifetime. Oviposition sites are often selected for protection; for example, tetranychid mites deposit eggs on undersides within webs produced by the females, which shield eggs from and predators. Factors such as host plant quality and population density influence egg production, with females reducing clutch sizes under high density to adjust offspring sex ratios. Sex determination in mites frequently follows haplodiploid arrhenotoky, particularly in Tetranychidae and some Parasitiformes, where ploidy dictates sex and unfertilized eggs yield males. Environmental cues, including population density and resource availability, can bias sex ratios, with females producing more daughters in low-density conditions to enhance mating opportunities. In oribatids, sex determination mechanisms remain less understood but may involve genetic factors independent of haplodiploidy in parthenogenetic lineages.

Development and Life Cycle

The development of mites (subclass Acari) typically follows an anamorphic pattern, involving a sequence of postembryonic stages that include an egg, followed by a hexapod , and then up to three nymphal instars (protonymph, deutonymph, and tritonymph) before reaching the adult form. Some species exhibit a non-feeding prelarval stage immediately after hatching, which is inactive and lacks functional mouthparts or legs. The larval stage is characterized by three pairs of legs and basic body segmentation, while nymphal stages acquire the fourth pair of legs and undergo progressive morphological refinements, such as the development of genital structures in the tritonymph. In certain astigmatid mites, the deutonymph may be heteromorphic, forming a specialized, dispersal-oriented hypopal stage adapted for phoresy on . The duration of the mite life cycle varies widely depending on , , and environmental conditions, ranging from as short as 5–12 days in rapidly developing groups like s (Tetranychidae) under optimal warm temperatures to 1–3 years or more in soil-dwelling oribatid mites. For instance, in the two-spotted spider mite (), the larval and nymphal stages together last 4–9 days at 25–30°C, enabling multiple generations per season in agricultural settings. In contrast, oribatid in temperate forest soils often require 12–24 months to complete development due to slower metabolic rates and extended immature periods. Metamorphosis in mites is gradual rather than abrupt, with key transformations occurring during ecdysis between instars, including the addition of legs in the first nymphal molt and maturation of sensory and reproductive organs in later stages. Environmental factors strongly influence these processes; development rates are highly temperature-dependent, accelerating with warmth (e.g., optimal at 25–30°C for many phytophagous species) and slowing or halting below 10–15°C. Quiescent or diapausing stages, often in eggs or deutonymphs, can be induced by adverse conditions like cold, drought, or short photoperiods, allowing survival until favorable cues trigger resumption; for example, overwintering diapause in spider mites involves arrested embryonic development responsive to chilling.

Ecological Roles

Habitats and Niches

Mites occupy a wide array of environmental habitats, demonstrating remarkable adaptability across terrestrial, freshwater, and marine ecosystems. In terrestrial environments, serves as a primary niche, where oribatid mites () predominate as key decomposers, facilitating nutrient cycling by breaking down in floors and grasslands. These mites thrive in the upper soil layers, contributing to and fertility through their feeding on and fungi. In freshwater systems, hydrachnidian mites (Hydrachnidia) are ubiquitous, inhabiting lotic (running water) and lentic (standing water) habitats such as streams, ponds, and wetlands, where they often associate with aquatic vegetation and substrates. Marine interstitial spaces, particularly sandy sediments and algal mats, host halacarid mites (Halacaridae), which exploit the pore waters of intertidal and subtidal zones for feeding on and . Beyond broad habitats, mites exploit diverse microhabitats that reflect their varied ecological roles. Phytophagous mites, such as spider mites (Tetranychidae), colonize plant surfaces like leaf undersides and stems, where they pierce cells to extract sap, often forming dense colonies on crops and ornamentals. Commensal mites inhabit the external or internal surfaces of animal hosts, including feather mites on birds that feed on secretions without harming the host. Certain mites also endure extreme conditions; for instance, thermacarid water mites (Thermacaridae) occupy hot springs with temperatures exceeding 40°C, while oribatid species persist in polar regions like , enduring freezing temperatures and in and . Mites are numerically dominant in many soil ecosystems, often comprising a significant portion of arthropod abundance; for example, in poplar plantation soils, Prostigmata can account for up to 40% of total soil arthropod abundance, with Oribatida contributing substantially. Their distribution exhibits vertical stratification within litter layers, with higher densities and diversity in the surface litter and fermentation layers compared to deeper humus and mineral soil, influenced by moisture, oxygen, and food availability. Specialized adaptations enable mites to persist in these niches. In arid terrestrial environments, many species possess cuticles with thickened layers and hydrocarbons that minimize transcuticular water loss, allowing survival in dry soils and deserts. Aquatic mites, conversely, employ osmoregulatory mechanisms, such as genital acetabula that facilitate ion and , preventing osmotic stress in fluctuating salinities of freshwater and marine habitats.

Interactions and Symbioses

Mites exhibit a diverse array of interactions with other organisms, ranging from predation and to symbiotic associations that influence ecological dynamics. Predatory mites, particularly those in the order , play a key role in controlling populations of smaller . These mites actively hunt nematodes and using chemosensory systems that detect chemical cues from prey, enabling efficient in and environments. For instance, mesostigmatid mites associated with bark beetles predominantly feed on arthropods or nematodes, with over half of studied species classified as predators. Such predation helps regulate pest populations, as soil predatory mites target plant-parasitic nematodes and other arthropods, contributing to biological control in agricultural and natural systems. Parasitic interactions further highlight mites' impact on host organisms. Many mites function as ectoparasites, attaching to the external surfaces of vertebrates and to feed on skin fluids or tissues. Chiggers, larval mites in the family , exemplify this mode by infesting vertebrates and occasionally transmitting bacterial diseases like during feeding. In insects, some mites adopt endoparasitic strategies, invading the hemocoel to feed on internal fluids; for example, pierces the host's to access and tissue, weakening the host while potentially vectoring pathogens. These mites often serve as vectors in disease transmission, facilitating the spread of bacteria, viruses, and other microbes between hosts through their feeding activities. Symbiotic associations among mites include phoresy and more complex mutualisms that aid dispersal and pathogen enhancement. Phoresy involves mites hitchhiking on larger insects, such as beetles or bees, to reach new habitats without direct harm to the carrier, a strategy common in patchy environments. In detrimental symbioses, Varroa mites form a mutualistic relationship with viruses like deformed wing virus (DWV) in honey bees, where the mite vectors the virus, and viral replication within the mite boosts its reproductive success, creating a feedback loop that amplifies harm to bee colonies. This interaction underscores how mite symbioses can evolve from neutral transport to pathogenic alliances. Within trophic levels, mites occupy multiple positions in food webs, particularly as detritivores facilitating . Oribatid mites, abundant in , consume organic and fungal hyphae, accelerating the breakdown of and nutrient cycling in ecosystems. As primary consumers, these detritivores integrate into chains, enhancing by processing dead matter. Mites also serve as prey for larger arthropods, such as predatory beetles and spiders, linking lower trophic levels to higher predators and maintaining balance in and food webs.

Dispersal Mechanisms

Mites employ a variety of active and passive strategies to disperse, constrained primarily by their small size, which limits unaided locomotion over long distances. Active dispersal typically involves walking or limited , but these methods are inefficient for most species due to their diminutive bodies, often measuring less than 1 mm in length. For instance, oribatid mites can walk short distances across or surfaces, but such movement rarely exceeds a few centimeters without external aid. In contrast, certain eriophyid mites, such as Aceria tosichella and Abacarus hystrix, exhibit specialized wind-blown dispersal, where active stages position themselves perpendicular to air currents to be carried aloft, achieving higher success rates in windy conditions compared to phoresy. Passive dispersal dominates mite movement, with phoresy being the most prevalent mechanism, wherein mites temporarily attach to larger hosts for transport to new habitats. Uropodine mites (: Uropodina), for example, use modified deutonymph stages equipped with anal pedicels to hitch rides on wood-inhabiting beetles, facilitating colonization of ephemeral resources like decaying logs. Similarly, astigmatid mites such as those in Glycyphagoidea attach to birds or mammals via ventral suckers or clasping structures, often dispersing through nests or during host migration. These associations enable mites to overcome , though attachment success varies with host availability and mite morphology. Long-distance dispersal often integrates active and passive elements, particularly in specialized taxa. Tetranychid spider mites, including , engage in ballooning by producing threads from their spinnerets, aggregating at apices to form wind-borne balls that can transport groups of adults and immatures over kilometers. This collective behavior enhances escape from crowded or depleted patches, with survival depending on prompt takeoff to avoid . Some soil mites, such as oribatids, utilize on floating debris, where individuals survive submersion for up to 365 days and disperse downstream at rates potentially reaching thousands of kilometers in river systems, aided by anti-wetting secretions. Dispersal barriers, including oceanic distances and habitat discontinuities, lead to genetic isolation in island populations, as evidenced by phylogenetic analyses of ameronothroid mites across the . These studies reveal pre-Pleistocene divergences among genera like Podacarus and Halozetes, indicating limited across the Antarctic Polar Front and survival in refugia during glaciation. Human-mediated spread via exacerbates connectivity, introducing invasive mites such as eriophyids and tetranychids through infested material or commodities, often bypassing barriers and accelerating range expansions.

Interactions with Humans

Health and Medical Impacts

Mites play a significant role in various parasitic diseases affecting humans and animals. , a highly contagious , is caused by the mite var. hominis, which burrows into the upper layer of the to lay eggs, leading to intense itching, rashes, and secondary infections. , another parasitic condition, occurs in mammals including dogs, cats, and humans due to overproliferation of mites residing in hair follicles and sebaceous glands, resulting in symptoms like , , and lesions, particularly in immunocompromised individuals. In , is a notable issue in dogs, where it manifests as generalized or localized mange-like . Certain mites act as vectors for bacterial diseases, exacerbating concerns. Lyme disease, the most common tick-borne illness in the , is transmitted by ticks infected with the spirochete bacterium , which the mites acquire from feeding on infected wildlife hosts before biting humans or animals, causing symptoms ranging from fever and rash to severe joint and neurological issues if untreated. Beyond direct , mites contribute to allergic disorders; house dust mites of the Dermatophagoides, such as D. pteronyssinus and D. farinae, produce allergens in their fecal pellets that trigger immunoglobulin E-mediated responses, leading to asthma exacerbations, allergic rhinitis, and atopic dermatitis in sensitized individuals worldwide. In occupational and veterinary contexts, mite-induced allergies and infestations pose additional challenges. Storage mites, including species like Tyrophagus putrescentiae and Acarus siro, cause "baker's itch" or allergic contact dermatitis among bakers and grain handlers through exposure to contaminated flour and stored products, manifesting as pruritic rashes and respiratory symptoms. Among pets, ear mites (Otodectes cynotis) infest the external ear canals of cats and dogs, leading to severe irritation, dark waxy discharge, head shaking, and potential secondary bacterial infections if untreated. In livestock, sarcoptic mange caused by Sarcoptes scabiei varieties results in intense pruritus, alopecia, and thickened skin, impacting animal welfare and productivity in species like cattle, sheep, and pigs. Recent environmental changes have amplified mite-related health risks. In the 2020s, warming climates have driven increases in populations, particularly Ixodes species, by extending active seasons and expanding geographic ranges, thereby heightening the transmission potential of vector-borne diseases like in regions such as and .

Economic and Agricultural Significance

Mites represent significant economic challenges in , particularly as pests affecting crop yields and quality. s of the genus Tetranychus, such as the two-spotted spider mite (T. urticae), are notorious for infesting a wide range of crops including , tomatoes, cucumbers, and apples, where their feeding punctures cells, causing characteristic stippling, bronzing, and reduced . This damage can lead to substantial yield losses; for instance, infestations on strawberries have been shown to reduce marketable yields by up to 30% in field-grown plants, with early-season attacks exacerbating the impact on perennial crops. Similarly, gall mites from the family Eriophyidae induce plant deformities by injecting growth-regulating saliva during feeding, resulting in , curling, russeting, and blistering on hosts like ornamentals, fruits, and vegetables, which diminish aesthetic and commercial value. These effects collectively contribute to significant annual global agricultural losses from mite pests, underscoring their role in reducing productivity across diverse farming systems. In stored product systems, mites like Acarus siro (the flour or grain mite) infest commodities such as , , dried fruits, and animal feeds, accelerating spoilage through feeding, fecal contamination, and promotion of fungal growth, which degrades nutritional quality and leads to product rejection. Infestations by A. siro and related species cause , accumulation of mite residues, and hazards in processed , resulting in substantial economic damages when considering broader stored-product pest impacts that include mites. These losses extend to the industry, where even low-level infestations can trigger recalls or market devaluation, amplifying costs for storage and quality . Forestry operations face threats from conifer-infesting mites, such as the spruce spider mite (Oligonychus ununguis), which targets and other evergreens by rasping needle surfaces, leading to defoliation, needle drop, and tree decline that reduces timber volume and aesthetic value in plantations. exacerbates these impacts, with warmer winters and altered precipitation patterns enabling larger mite populations and more frequent outbreaks in coniferous forests, as milder conditions enhance overwintering survival and extend active feeding periods. In regions like , such dynamics have contributed to heightened vulnerability of stands, with infestations causing significant economic setbacks in timber production. Effective relies on monitoring pest densities against established action thresholds to guide interventions and minimize unnecessary treatments. For spider mites, thresholds often range from 5-10 mites per leaf in crops like tomatoes and strawberries, beyond which yield impacts become economically significant, prompting application or other controls. However, widespread resistance to acaricides complicates control; T. urticae has developed tolerance to multiple chemical classes, including pyrethroids and organophosphates, often within 1-4 years of repeated exposure, necessitating rotation of modes of action to sustain efficacy. This resistance pattern, driven by enhanced enzymes and reduced penetration, has led to increased costs and persistent outbreaks in and settings.

Beneficial Applications

Mites play a significant role in biological control programs, particularly through the use of predatory species to manage pest populations in agricultural settings. The predatory mite Phytoseiulus persimilis is widely employed against the two-spotted spider mite () in crops such as strawberries and ornamentals. This species targets all life stages of the pest, with adults exhibiting the highest consumption rates among available biocontrol predatory mites. Releases are recommended at rates of 1-3 mites per for preventative control, 5 per for low to moderate infestations, and 10 or more per for high infestations, often requiring multiple applications in outdoor environments due to dispersal. studies show that maintaining a pest-to-predator ratio of 5-10:1 can significantly reduce populations within 2-3 weeks, with evaluation based on observing shriveled, dead prey mites on foliage. In , the parasitic mite serves as a key indicator for hive through non-chemical monitoring techniques. Mite drop counts, often using sticky boards placed beneath brood frames or methods like powdered sugar shakes or alcohol es on samples of 300 bees, allow beekeepers to estimate infestation levels by calculating the percentage of mites per 100 bees. This method exploits natural mite fall from grooming or brood emergence, enabling early detection to prevent colony decline. Guidelines from the Honey Bee Health Coalition recommend thresholds of less than 1% during dormancy (acceptable), 1-2% (caution), and over 2% (treatment needed), with sampling advised four times annually, post-treatment, and during peak brood periods to inform . As of 2025, V. destructor has caused catastrophic losses in U.S. colonies, with commercial beekeepers reporting an average of 62% colony mortality from June 2024 to March 2025, linked to miticide resistance and associated viruses like , highlighting the urgent need for improved monitoring and control strategies. Beyond , mites contribute to by participating in succession patterns on decomposing remains, aiding in estimation. Various acarid arrive on cadavers via phoresy on or direct dispersal, with community composition shifting across stages—early colonizers like gamasid mites feeding on fluids, followed by oribatids during dry phases. Studies highlight their potential as markers, as specific mite assemblages can indicate conditions or time since , with like Macrocheles common in initial stages. Forensic acarology emphasizes mites' microhabitat specificity, allowing reconstruction of events even without the carrier present. Oribatid mites also function as bioindicators of in agricultural systems, where their diversity and abundance reflect quality and management practices. These mites, comprising a dominant group in microarthropod communities, drive decomposition and nutrient cycling, with higher correlating to sustainable practices like organic fertilization in crop rotations. In Mediterranean vineyards transitioning to , oribatid increases under reduced and cover crops, signaling improved and fertility. Monitoring their communities helps assess impacts from disturbances, as low diversity often indicates degradation, while resilient assemblages support long-term . Emerging research explores mites' indirect contributions to bioremediation through their facilitation of organic matter decomposition in contaminated soils. Oribatid mites enhance litter breakdown by feeding on fungi, bacteria, and detritus, promoting microbial activity that aids in the natural attenuation of pollutants like hydrocarbons. In polluted sites, their presence influences soil processes, though high contaminant levels can suppress populations, underscoring their sensitivity as indicators during remediation efforts.

Cultural Representations

Mites have appeared in philosophical and literary works as symbols of the infinitesimal and human insignificance. In Blaise Pascal's Pensées (1670), he describes a mite's minute body and intricate parts—limbs, veins, and humors—to illustrate the vast scale between the infinitely small and large, emphasizing humanity's precarious position in the universe. This depiction underscores mites as metaphors for the overlooked details of creation, influencing later reflections on scale in natural philosophy. In visual art, mites gained prominence through early . Robert Hooke's (1665) featured detailed engravings of mites, including a "wandering mite" and crab-like forms observed under compound microscopes, revealing their complex anatomy and sparking wonder at the microscopic world. These illustrations, among the first public depictions of mites, portrayed them not as pests but as marvels of design, bridging art and science in the . In modern literature, particularly , mites often symbolize invasive threats or existential perils. Charles Pellegrino's novel (1998) depicts swarms of genetically engineered mites devastating ecosystems, representing uncontrolled biological catastrophe. Similarly, in Mircea Cărtărescu's (2015), a enters a parallel mite world, exploring themes of alienation and microscopic societies. Folklore traditions occasionally associate mites with omens or natural cycles. In oral , proverbs like "más viejo que la sarna" (older than , referring to mites) evoke enduring afflictions as symbols of antiquity and persistence. Some Indigenous Mayan practices in link ticks (a type of mite) to seasonal rituals on St. Francis of Assisi's day, viewing them as harbingers of environmental balance or imbalance. Contemporary media highlights mites' ubiquity through educational documentaries and viral content. The PBS series Deep Look (2016) episode on dust mites examines their role in ecosystems, blending with narrative to demystify these invisible companions. Online memes and social media exhibits, such as artistic renderings of eyelash mites, have popularized their presence on human faces, often evoking humorous horror at their pervasiveness—nearly all adults host them without harm. These representations foster public fascination with mite diversity, estimated at over 1 million .

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