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The Acariformes, also known as the Actinotrichida, are the most diverse of the two superorders of
mite 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, but genetic analysis does not show clear evid ...
s. Over 32,000 described
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
are found in 351
families Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideal ...
, with an estimated total of 440,000 to 929,000 species, including
undescribed species In taxonomy, an undescribed taxon is a taxon (for example, a species) that has been discovered, but not yet formally described and named. The various Nomenclature Codes specify the requirements for a new taxon to be validly described and named. U ...
.


Systematics and taxonomy

The Acariformes can be divided into two main
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
s – Sarcoptiformes and
Trombidiformes The Trombidiformes are a large, diverse order of mites. Taxonomy In 1998, Trombidiformes was divided into the Sphaerolichida and the Prostigmata. The group has few synapomorphies by which it can be defined, unlike the other major group of ac ...
. In addition, a
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
group containing primitive forms, the
Endeostigmata Endeostigmata is a suborder of acariform mites. There are about ten families in Endeostigmata. The grouping is strongly suspected to be paraphyletic, containing unrelated early diverging lineages of mites. Taxonomy * Suborder Endeostigmata Reut ...
, was formerly also considered distinct. The latter is composed of only 10 families of little-studied, minute, soft-bodied mites that ingest solid food, such as
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
,
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
, and soft-bodied
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s such as
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
s,
rotifer The rotifers (, from the Latin , "wheel", and , "bearing"), commonly called wheel animals or wheel animalcules, make up a phylum (Rotifera ) of microscopic and near-microscopic pseudocoelomate animals. They were first described by Rev. John H ...
s, and
tardigrade Tardigrades (), known colloquially as water bears or moss piglets, are a phylum of eight-legged segmented micro-animals. They were first described by the German zoologist Johann August Ephraim Goeze in 1773, who called them Kleiner Wasserbä ...
s. These clades were formerly considered suborders, but this does not allow for a sufficiently precise classification of the mites and is abolished in more modern treatments; the Endeostigmata are variously considered to form a suborder on their own (the old view) or are included mainly in the Sarcoptiformes, thus making both groups
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
. The superfamily
Eriophyoidea Eriophyoidea are a superfamily of herbivorous mites. All post-embryonic instars lack the third and fourth pairs of legs. The respiratory system is also absent. The taxonomy of this group has always been confused. There were families created for ...
, traditionally considered members of the Trombidiformes, have been found to be basal mites in genomic analyses, sister to the clade containing Sarcoptiformes and Trombidiformes. Another group often mentioned is the Actinedida, but in treatments like the present one, this is split up between the Sarcoptiformes (and formerly the separate Endeostigmata) and Trombidiformes (which contains the bulk of the "Actinedida"), because it appears to be a massively paraphyletic "
wastebin taxon Wastebasket taxon (also called a wastebin taxon, dustbin taxon or catch-all taxon) is a term used by some taxonomists to refer to a taxon that has the sole purpose of classifying organisms that do not fit anywhere else. They are typically defined ...
", uniting all Acariformes that are not "typical"
Oribatida Oribatida (formerly Cryptostigmata), also known as oribatid mites, moss mites or beetle mites, are an order (biology), order of mites, in the "chewing Acariformes" clade Sarcoptiformes. They range in size from . There are currently 12,000 species ...
and
Astigmatina 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 (synonym Desmonomata) in the order Sarc ...
. The Trombidiformes present their own problems. The small group Sphaerolichida appears to be the most ancient lineage among them. However, the
Prostigmata The Prostigmata is a suborder of mites belonging to the order Trombidiformes, which contains the "sucking" members of the "true mites" (Acariformes). Many species are notorious pests on plants. Well-known examples of prostigmatan plant par ...
are variously subdivided into the
Anystina The Prostigmata is a suborder of mites belonging to the order Trombidiformes, which contains the "sucking" members of the "true mites" (Acariformes). Many species are notorious pests on plants. Well-known examples of prostigmatan plant parasit ...
and
Eleutherengona The Prostigmata is a suborder of mites belonging to the order Trombidiformes, which contains the "sucking" members of the "true mites" (Acariformes). Many species are notorious pests on plants. Well-known examples of prostigmatan plant parasit ...
, and
Eupodina The Prostigmata is a suborder of mites belonging to the order Trombidiformes, which contains the "sucking" members of the "true mites" (Acariformes). Many species are notorious pests on plants. Well-known examples of prostigmatan plant parasit ...
. The delimitation and interrelationships of these groups are entirely unclear; while most analyses find one of the latter two, but not the other to be a subgroup of the Anystina; neither of these mutually contradicting hypotheses is very robust; possibly this is a simple error because phylogenetic software usually fails in handling non
dichotomous A dichotomy is a partition of a whole (or a set) into two parts (subsets). In other words, this couple of parts must be * jointly exhaustive: everything must belong to one part or the other, and * mutually exclusive: nothing can belong simult ...
phylogenies. Consequently, it may be best for the time being to consider each of the three main prostigmatan lineages to be equally distinct from the other two.


Fossil record

The oldest fossils of acariform mites are from the
Rhynie Chert The Rhynie chert is a Lower Devonian sedimentary deposit exhibiting extraordinary fossil detail or completeness (a Lagerstätte). It is exposed near the village of Rhynie, Aberdeenshire, Scotland; a second unit, the Windyfield chert, is locate ...
, Scotland, which dates to the early
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
, around 410 million years ago The
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
'' Immensmaris chewbaccei'' had idiosoma of more than in length and was the largest fossil acariform mite and also the largest erythraeoid mite ever recorded.


Diversity

The Sarcoptiformes ingest solid food, being mainly microherbivores,
fungivore Fungivory or mycophagy is the process of organisms consuming fungi. Many different organisms have been recorded to gain their energy from consuming fungi, including birds, mammals, insects, plants, amoebas, gastropods, nematodes, bacteria and oth ...
s and detritivores. Some
Astigmatina 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 (synonym Desmonomata) in the order Sarc ...
– the
Psoroptidia Psoroptidia is a parvorder of the Acari (mite) group Astigma (or Astigmatina). It comprises around 40 families, and apparently originated as parasites of birds, before a secondary radiation saw some taxa become parasites of mammals. Because of t ...
– have become associated with
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s and nest-building
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s. These include the well known
house dust mite House dust mites (HDM, or simply dust mites) are various species of acariform mites belonging to the family Pyroglyphidae that are found in association with dust in dwellings. They are known for causing allergies. Biology Species The curren ...
s, scab mites and mange mites, stored product mites, feather mites and some fur mites. The relationships between their main groups are not well-resolved and subject to revision. In particular it appears as if the
Oribatida Oribatida (formerly Cryptostigmata), also known as oribatid mites, moss mites or beetle mites, are an order (biology), order of mites, in the "chewing Acariformes" clade Sarcoptiformes. They range in size from . There are currently 12,000 species ...
need to be split up in two, as the
Astigmatina 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 (synonym Desmonomata) in the order Sarc ...
are closer to some of them (e.g. certain Desmonomata) than the latter are to other "Oribatida". The Trombidiformes are most noted for the economic damage caused by many
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclud ...
parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
species. All of the most important plant
pests PESTS was an anonymous American activist group formed in 1986 to critique racism, tokenism, and exclusion in the art world. PESTS produced newsletters, posters, and other print material highlighting examples of discrimination in gallery represent ...
among the
Acari 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, but genetic analysis does not show clear evid ...
are trombidiformans, such as spider mites (Tetranychidae) and Eriophyidae. Many species are also predators,
fungivore Fungivory or mycophagy is the process of organisms consuming fungi. Many different organisms have been recorded to gain their energy from consuming fungi, including birds, mammals, insects, plants, amoebas, gastropods, nematodes, bacteria and oth ...
s, and animal parasites. Some of the most conspicuous species of free-living mites are the relatively large and bright red velvet mites, that belong to the family
Trombidiidae Trombidiidae, also known as red velvet mites, true velvet mites, or rain bugs, are small arachnids (eight-legged arthropods) found in plant litter and are known for their bright red color. While adults are typically in length, some, such as t ...
. Oribatid mites and to a much lesser extent others are a source of alkaloids in poison frogs (namely small species like the strawberry poison-dart frog ''Oophaga pumilio''). Such frogs raised without these oribatids in their diets do not develop the strong poisons associated with them in the wild.


Parthenogenesis

Acariformes species appear to have evolved from a sexual ancestor and the primary manner of reproduction during the course of evolution has been
sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...
. However, within the super order Acariformes,
parthenogenetic Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and developmen ...
species have arisen numerous times during the course of evolution. In contrast to the commonly held view that parthenogenetic lineages are short lived, four species-rich parthenogenetic clusters of the order
Oribatida Oribatida (formerly Cryptostigmata), also known as oribatid mites, moss mites or beetle mites, are an order (biology), order of mites, in the "chewing Acariformes" clade Sarcoptiformes. They range in size from . There are currently 12,000 species ...
are very ancient and likely arose 400-300 million years ago. In some parthenogenetic species that undergo
automixis ''Automixis'' is the fusion of (typically haploid) nuclei or gametes derived from the same individual. The term covers several reproductive mechanisms, some of which are parthenogenetic. Diploidy might be restored by the doubling of the chromoso ...
(a kind of self-fertilization that retains
meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately resu ...
) sexual reproduction has re-emerged.


Examples

Eriophyidae, plant
parasites Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson ha ...
, e.g. '' Acalitus essigi'' (redberry mite) ; Sarcoptiformes *
Cheese mite Cheese mites (for instance '' Tyrophagus casei'' or other species) are mites that are used to produce such cheeses as ', Cantal and Mimolette. The action of the living mites on the surface of these cheeses contributes to the flavor and gives them ...
s * Epidermoptidae * Gastronyssidae *''
Sarcoptes scabiei ''Sarcoptes scabiei'' or the itch mite is a parasitic mite that burrows into skin and causes scabies. The mite is found in all parts of the world. Humans are not the only mammals that can become infected. Other mammals, such as wild and domesti ...
'' ;
Trombidiformes The Trombidiformes are a large, diverse order of mites. Taxonomy In 1998, Trombidiformes was divided into the Sphaerolichida and the Prostigmata. The group has few synapomorphies by which it can be defined, unlike the other major group of ac ...
*
Demodecidae ''Demodecidae'' is a family of parasitic mites, living on various species of mammals. Each species of mite is usually only found on a single mammal species, whereas a mammal species can have several different species of demodecid mites living o ...
, e.g. Demodex mites *
Erythraeidae Erythraeidae is a family of mites belonging to the Trombidiformes. Larval forms of these mites are parasitic on various other arthropods, for example harvestmen, but the adults are free-living predators. These oval mites are rather large, usually ...
* Labidostommatidae *''
Polydiscia deuterosminthurus ''Polydiscia deuterosminthurus'' is a species of mite recently discovered in the autonomous community of Navarre in Spain. Only the six-legged larval stage is so far known. This lives as a parasite on the springtail '' Deuterosminthurus bisetos ...
'' *
Smarididae Smarididae is a family of mites belonging to the order Trombidiformes. These large predatory mites have long oval bodies, distinctively pointed in front. They are usually red and densely hairy with slender legs, sometimes very long. They have e ...
* Spider mites, e.g. ''
Tetranychus urticae ''Tetranychus urticae'' (common names include red spider mite and two-spotted spider mite) is a species of plant-feeding mite generally considered to be a pest. It is the most widely known member of the family Tetranychidae or spider mites. It ...
'' *
Tarsonemidae Tarsonemidae is a family of mites, also called thread-footed mites or white mites. Only a limited number of tarsonemid genera ('' Steneotarsonemus'', '' Polyphagotarsonemus'', '' Phytonemus'', '' Floridotarsonemus'' and '' Tarsonemus'') are kno ...
, a number of which are plant
pests PESTS was an anonymous American activist group formed in 1986 to critique racism, tokenism, and exclusion in the art world. PESTS produced newsletters, posters, and other print material highlighting examples of discrimination in gallery represent ...
, e.g. ''
Acarapis woodi ''Acarapis woodi'' is an internal parasite affecting honey bees, the symptoms of infestation was originally observed on the Isle of Wight in 1904, but was not species description, described until 1921. ''Acarapis woodi'' mites live and reproduce ...
'' *
Tydeidae Tydeidae is a family of acariform mites. As of 2016, it contained over 300 species in three subfamilies, though more species have been discovered since then. These mites live in a wide range of habitats and there are predatory, fungivorous an ...


References


Further reading

* {{Taxonbar, from=Q1341457 Arthropod superorders