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#REDIRECT [[Mite]]
{{other uses}}
{{Automatic taxobox
| fossil_range = {{Fossil range|Early Devonian|recent}}
| image = Peacock mite, Tuckerella sp.jpg
| image_caption = [[Peacock mite]] (''Tuckerella'' sp.),<br />[[false-color|false-colour]] [[Scanning electron microscope|SEM]], magnified 260×
| taxon = Acari
| authority = [[William Elford Leach|Leach]], 1817
| subdivision_ranks = [[Superorder]]s
| subdivision =
* [[Acariformes]]
* [[Parasitiformes]]
and see [[#Taxonomy and phylogeny|text]]
}}


{{Rcat shell|
The '''Acari''' {{IPAc-en|ˈ|æ|k|ər|aɪ}} (or '''Acarina''' {{IPAc-en|æ|k|ə|ˈ|r|aɪ|n|ə}}) are a [[taxon]] of [[arachnids]] that contains [[mite]]s and [[tick]]s. The diversity of the Acari is extraordinary and their [[fossil record|fossil history]] goes back to at least the early [[Devonian]] period.<ref>{{cite web |first1= David Evans | last1= Walter| first2= Gerald |last2= Krantz| first3= Evert |last3= Lindquist |title=Acari. The Mites |url= http://tolweb.org/Acari/2554/1996.12.13 |date=December 13, 1996 |publisher= [[Tree of Life Web Project]] |accessdate=June 21, 2010}}</ref> Acarologists (people who study the Acari) have proposed a complex set of taxonomic ranks to classify mites. In most modern treatments, the Acari are considered a [[Class (biology)|subclass]] of the Arachnida and are composed of two or three [[superorder]]s or [[order (biology)|orders]]: [[Acariformes]] (or Actinotrichida), [[Parasitiformes]] (or Anactinotrichida), and [[Opilioacariformes]]; the latter is often considered a subgroup within the Parasitiformes. The [[monophyly]] of the Acari is open to debate, and the relationships of the acarines to other arachnids is not at all clear.<ref name=Garw>{{cite journal|title=Three-dimensional reconstruction and the phylogeny of extinct chelicerate orders|first=Russell J.|last=Garwood|first2=Jason A.| last2= Dunlop |year= 2014| journal= PeerJ| volume= 2| pages= e641| doi= 10.7717/peerj.641 |pmid= 25405073 |pmc= 4232842}}</ref> In older treatments, the subgroups of the Acarina were placed at order rank, but as their own subdivisions have become better understood, treating them at the superorder rank is more usual.
{{R to related topic}}

{{R from scientific name|arthropod}}
Most acarines are minute to small (for example, {{cvt|0.08|-|1.00|mm|disp=or|3}}), but the largest (some ticks and [[red velvet mite]]s) may reach lengths of {{cvt|10|-|20|mm|1}}. Over 50,000 species have been described (as of 1999) and an estimated million or more species may exist. The study of mites and ticks is called '''[[acarology]]''' (from [[Ancient Greek|Greek]] {{Lang|el|ἀκαρί}}/{{Lang|el|ἄκαρι}}, ''{{Lang|el|akari}}'', a type of mite; and {{Lang|el|-λογία}}, ''[[-logy|-logia]]''),<ref name="Walter">{{cite book |first1= D. E. | last1= Walter| first2=H. C.| last2= Proctor |year=1999 |title=Mites: Ecology, Evolution and Behaviour |publisher=University of NSW Press, Sydney and [[CAB International|CABI]], Wallingford |isbn=978-0-86840-529-2}}</ref> and the leading [[scientific journal]]s for acarology include ''[[Acarologia]]'', ''[[Experimental and Applied Acarology]]'' and the ''[[International Journal of Acarology]]''.
{{R with history}}

==Morphology==
Mites are [[arachnid]]s, and as such, evolved from a segmented body with the segments organised into two [[tagma (biology)|tagmata]]: a [[prosoma]] (cephalothorax) and an [[opisthosoma]] (abdomen). However, only the faintest traces of primary segmentation remain in mites; the prosoma and opisthosoma are fused, and a region of flexible cuticle (the circumcapitular furrow) separates the chelicerae and pedipalps from the rest of the body. This anterior body region is called the capitulum or [[gnathosoma]], and according to some works, is also found in the [[Ricinulei]]. The remainder of the body is called the idiosoma and is unique to mites.

Most adult mites have four pairs of legs, like other arachnids, but some have fewer. For example, gall mites like ''[[Phyllocoptes variabilis]]'' (family [[Eriophyidae]]) have a worm-like body with only two pairs of legs; some parasitic mites have only one or three pairs of legs in the adult stage. Larval and prelarval stages have a maximum of three pairs of legs; adult mites with only three pairs of legs may be called 'larviform'. Also, members of the Nematalycidae within the [[Endeostigmata]], which live between sand grains, have often worm-like and elongated bodies with reduced legs.<ref>{{cite web| url= https://macromite.wordpress.com/category/endeostigmata/ |website= macromite.wordpress.com| title= Of Knots & Worms Not: Gordialycus| date= May 23, 2010| access-date= October 31, 2017}}</ref>

The mouth parts of mites may be adapted for biting, stinging, sawing, or sucking. They breathe through [[Invertebrate trachea|tracheae]], stigmata (small openings of the skin), intestines, and the skin itself. Species hunting for other mites have very acute senses, but many mites are eyeless. The central eyes of arachnids are always missing, or they are fused into a single eye. Thus, any eye number from none to five may occur.<ref name=schmidt93>{{cite book |first= Günther |last= Schmidt |year=1993 |title=Giftige und gefährliche Spinnentiere |language= German |publisher=Westarp Wissenschaften |isbn=978-3-89432-405-6 |pages=58ff}}</ref>

==Ontogeny==
[[File:Ontogeny Chaetodactylus krombeini.jpg|thumb|right|Life stages of ''[[Chaetodactylus krombeini]]'' (Astigmata); male not shown]]
[[File:Argas spec columbidae.jpg|thumb|right|A soft-bodied tick of the family [[Argasidae]], beside eggs it has just laid]]
Acarine [[ontogeny]] typically consists of an egg, a prelarval stage (often absent), a larval stage (hexapod except in the mite superfamily [[Eriophyoidea]], which have only two pairs of legs), and a series of nymphal stages. Any or all of these stages except the adult may be suppressed or occur only within the body of a previous stage. Larvae (and prelarvae) have a maximum of three pairs of legs (legs are often reduced to stubs or absent in prelarvae); legs IV are added at the first nymphal stage. Usually, a maximum of three nymphal stages are present and they are referred to in sequence as the protonymph, deutonymph, and tritonymph; however, some soft ticks have supernumerary nymphal stages. The females of some [[Tarsonemidae]] bear sexually mature young. If any nymphal stages are absent, then authors may disagree on which stages are present. Only the [[Oribatida]] pass through all developmental stages.<ref name=schmidt93/>

==Diversity and lifestyles==
Acarines are extremely diverse. They live in practically every habitat, and include aquatic (fresh and sea water) and terrestrial species. They outnumber other [[arthropod]]s in the soil [[organic material|organic matter]] and [[detritus (biology)|detritus]]. Many are [[parasitism|parasitic]], and they affect both [[vertebrate]]s and [[invertebrate]]s. Most parasitic forms are external parasites, while the free-living forms are generally [[predation|predatory]] and may even be used to control undesirable arthropods. Others are [[detritivore]]s that help to break down forest [[plant litter|litter]] and dead organic matter, such as [[skin]] cells. Others still are [[herbivory|plant feeders]] and may damage [[crops]].

The [[feather mite]]s, Astigmata, are found on almost all species of birds, except for penguins, and are highly specialized for life on their hosts. They may feed on [[Uropygial gland|uropygial oil]], skin flakes, fungus, bacteria, and feathers, depending on the taxon to which they belong. Their lifestyles are affected by the microclimate (ambient temperature and relative humidity); for example, seasonal change in temperature causes feather mites to shift their microhabitats on [[Eurasian blue tit|blue tits]]. However, no evidence shows microclimate affecting mite diversity.<ref>{{Cite journal|url = |title = Diversity of Feather Mites (Acari: Astigmata) on Darwin's Finches|last = Villa|first = Scott M|date = October 2013|journal = The Journal of Parasitology|doi = 10.1645/12-112.1|pmid = 23691947|access-date = |last2 = Bohec|first2 = Celine Le |volume=99 |issue = 5|pages=756–762|pmc = 4098782}}</ref>

==Economic and medical importance==
[[File:Tick male (aka).jpg|right|thumb|Male tick (size: 2&nbsp;mm)]]

Damage to crops is perhaps the most costly economic effect of mites, especially by the [[spider mite]]s and their relatives (Tetranychoidea), earth mites ([[Penthaleidae]]), thread-footed mites ([[Tarsonemidae]]) and the gall and rust mites ([[Eriophyidae]]). The [[honey bee]] parasite ''[[Varroa destructor]]'' has caused or contributed to large-scale die-offs of commercial pollinating populations.<ref>{{Cite journal|last=Potts|first=Simon G.|last2=Biesmeijer|first2=Jacobus C.|last3=Kremen|first3=Claire|last4=Neumann|first4=Peter|last5=Schweiger|first5=Oliver|last6=Kunin|first6=William E.|date=2010-06-01|title=Global pollinator declines: trends, impacts and drivers|journal=Trends in Ecology & Evolution|volume=25|issue=6|pages=345–353|doi=10.1016/j.tree.2010.01.007|pmid=20188434|citeseerx=10.1.1.693.292}}</ref>

Some parasitic forms affect [[human]]s and other [[mammal]]s, causing damage by their feeding, and can even be [[tick-borne disease|vectors of diseases]], such as [[scrub typhus]], [[rickettsialpox]], [[Lyme disease]], [[Q fever]], [[Colorado tick fever]], [[tularemia]], tick-borne [[relapsing fever]], [[babesiosis]], [[ehrlichiosis]], and [[tick-borne meningoencephalitis]].<ref>{{cite web| url=http://publichealth.lacounty.gov/eh/progs/consumer/vectman/vcdocs/ticks.pdf | title =Managing Common Tick Pests in Los Angeles County | publisher= Vector Management Program, Department of Health Services, Los Angeles County, California | accessdate=May 20, 2009}}</ref> A well-known effect of mites on humans is their role as [[allergen]]s and the stimulation of [[asthma]] in people affected by [[respiratory disease]].

The use of predatory mites (for example, [[Phytoseiidae]]) in [[pest control]] and herbivorous mites that infest weeds are also of importance. An unquantified, but major positive contribution of the Acari is their normal functioning in ecosystems, especially their roles in the decomposer subsystem.<ref name="Walter"/> In this context, the association of many species with carcasses and decaying organic matter qualify them as potential medicolegal indicators in [[forensic entomology]].<ref>{{cite journal | author1=González Medina, A|author2=González Herrera, L|author3=Perotti, MA|author4=Jiménez Ríos, G | title=Occurrence of ''Poecilochirus austroasiaticus'' (Acari: Parasitidae) in forensic autopsies and its application on postmortem interval estimation | journal=Exp. Appl. Acarol. | year=2013 | volume=59 | issue=3 | pages= 297–305 | doi=10.1007/s10493-012-9606-1|pmid=22914911}}</ref>

Chemical agents used to control ticks and mites include [[sulfur|dusting sulfur]] and [[ivermectin]].

==Taxonomy and phylogeny==
[[File:Rust Mite, Aceria anthocoptes.jpg|right|thumb|Rust mite, ''[[Aceria anthocoptes]]'' (size: 50&nbsp;μm)]]
[[File:Acari.tif|thumb|Unidentified acarid from a leaf. Microscopic image, manual z-stacking.]]

The [[phylogeny]] of the Acari is still disputed and several taxonomic schemes have been proposed for their classification. The third edition (2009) of the standard textbook ''A Manual of Acarology'' uses a system of six [[Order (biology)|orders]], grouped into three superorders:<ref>{{cite book |title=A Manual of Acarology |edition=3rd |editor-first1= Gerald W. |editor-last1= Krantz|editor-first2= D. E.|editor-last2= Walter |year=2009 |publisher= [[Texas Tech University Press]] |isbn=978-0-89672-620-8}}</ref>
* Superorder '''[[Opilioacariformes]]''' – mites that superficially resemble [[harvestmen]] ([[Opiliones]]), hence their name
* Superorder '''[[Parasitiformes]]''' – ticks and a variety of mites
** [[Holothyrida]]
** [[Ixodida]] – ticks
** [[Mesostigmata]] – bird mites, phytoseiid mites, ''Raubmilben''
*** [[Trigynaspida]]
*** [[Monogynaspida]]
* Superorder '''[[Acariformes]]''' – the most diverse group of mites
** [[Trombidiformes]] – plant parasitic mites (spider mites, peacock mites, gall mites, red-legged earth mites, etc.), snout mites, chiggers, hair follicle mites, velvet mites, water mites, etc.
*** [[Sphaerolichida]]
*** [[Prostigmata]]
** [[Sarcoptiformes]]
*** [[Oribatida]] – oribatid mites, beetle mites, armored mites (also cryptostigmata)
*** [[Astigmata]] – stored product, fur, feather, dust, and human itch mites, etc.

Recent genetic research has caused a change in the naming scheme, however, and recent publications have changed the superorder Parasitiformes to an order.<ref>{{cite journal |first1=S.C. |last1=Barker |first2=A. |last2=Murrell |date=2004 |title=Systematics and evolution of ticks with a list of valid genus and species names |journal=Parasitology |volume=129 |issue=7 |pages=S15–S36 |doi=10.1017/S0031182004005207 |lastauthoramp=yes }}</ref> Other recent research has suggested that Acari is [[polyphyletic]], with ticks and spiders being more closely related than ticks and mites.<ref>{{cite journal|last1=Sanggaard|first1=Kristian W.|last2=Bechsgaard|first2=Jesper S.|last3=Fang|first3=Xiaodong|title=Spider genomes provide insight into composition and evolution of venom and silk|journal=Nature Communications |date=6 May 2014| volume= 5|pages=3765| doi= 10.1038/ncomms4765|language=en|issn=2041-1723| pmid= 24801114| pmc= 4273655|bibcode=2014NatCo...5.3765S}}</ref> The cladogram is based on Dabert et al. 2010, which used molecular data. It showed the Acariformes sister to the [[Solifugae]] (camel spiders), while the Parasitiformes were sister to the [[Pseudoscorpionida]].<ref name= "DabertWitalinski2010">{{cite journal |last1=Dabert |first1=Miroslawa |last2=Witalinski |first2=Wojciech |last3=Kazmierski |first3=Andrzej |last4=Olszanowski |first4=Ziemowit |last5=Dabert |first5=Jacek |title=Molecular phylogeny of acariform mites (Acari, Arachnida): Strong conflict between phylogenetic signal and long-branch attraction artifacts |journal= Molecular Phylogenetics and Evolution|volume=56 |issue=1 |year=2010|pages=222–241 |issn= 1055-7903 |doi=10.1016/j.ympev.2009.12.020|pmid=20060051 }}</ref>

{{barlabel
|size=5
|at1=2|label1="'''Acari'''" (mites and ticks)
|cladogram={{clade
|label1=part of [[Arachnida]]
|1={{cladex
|1={{cladex
|1={{cladex
|label1=[[Pseudoscorpionida]] (false scorpions)
|1=[[File:Pseudoscorpion - Soil Fauna Diversity.jpeg|70px]]
|label2=[[Parasitiformes]] (parasitic mites, ticks)
|2={{cladex|barbegin1=red
|1=[[Ixodida]] (ticks) [[File:Tick_male_(aka).jpg|50px]]
|2=Parasitic mites, inc. ''[[Varroa]]'' [[File:5-Varroa destructor on head bee pupa by Gilles San Martin.jpg|50px]]
}}
}}
}}
|2={{cladex
|1={{cladex
|label1=[[Acariformes]] (other mites)
|1={{cladex
|1=[[Trombidiformes]] (chiggers, gall & velvet mites, etc) [[File:Trombidium holosericeum (aka).jpg|50px]]&nbsp;
|2=[[Sarcoptiformes]] (dust mites, fur mites, etc) [[File:Plateremaeoidea.jpg|50px]]
}}
|label2=[[Solifugae]] (camel spiders) |barend1=red
|2=[[File:MojaveSolifugid.JPG|50px]]
}}
}}
}}
}}
}}
}}

==See also==
*[[Mite]]

==References==
{{Reflist|28em}}

==Further reading==
* {{cite book |first= Wojciech |last= Niedbala |year=1992 |title=Phthiracaroidea (Acari, Oribatida): Systematic Studies |publisher=Warsaw: [[Polish Scientific Publishers PWN|PWN]], Amsterdam: [[Elsevier]] |isbn=978-8-301-09740-0}}
* [[Experimental and Applied Acarology]], {{ISSN|1572-9702}} (electronic) {{ISSN|0168-8162}} (paper), Springer
* {{cite book |first= E. |last= Baker |year=1952 |title=An Introduction to Acarology |url= https://archive.org/details/introductiontoac00bake |publisher=New York: The MacMillan Company}}
* {{cite book |first= T. |last= Woolley |year=1988 |title=Acarology: Mites and Human Welfare |publisher=New York: [[Wiley Interscience]] |isbn=978-0-471-04168-9}}
* {{cite book |first1= R. B.| last1= Halliday|first2= D. E.| last2= Walter| first3= H. C. | last3= Proctor| first4= R. A.| last4= Norton| first5= M. J.| last5= Colloff |year=2001 |title=Acarology, Proceedings of the 10th International Congress [5–10 July 1998] |publisher=[[CSIRO Publishing]], Melbourne |pages=960 pp |isbn=978-0-643-06658-8 |location=Collingwood, Victoria}}
* {{cite book |first1= D. E.| last1= Walter| first2=H. C. |last2= Proctor |year=2001 |title=Mites in soil, an interactive key to mites and other soil microarthropods |publisher=ABRS Identification Series, CSIRO Publishing, Collingwood, Victoria}}
* {{cite book |title=A Manual of Acarology |edition=3rd |editor-first1=Gerald W. | editor-last1= Krantz| editor-first2= D. E. | editor-last2= Walter |year=2009 |publisher= [[Texas Tech University Press]] |isbn=978-0-89672-620-8}}

==External links==
{{Commons category|Acari}}
{{Wikispecies|Acari}}
{{Wikibooks|Dichotomous Key|Acari}}
*[[Tree of Life Web Project]]
** {{cite web |url=http://tolweb.org/Acari/2554 |title=Acari. The Mites |first1= David Evans | last1= Walter| first2= Gerald |last2= Krantz |first3= Evert | last3= Lindquist |date=December 13, 1996 |website= tolweb.org |publisher= Tree of Life Web Project}}
** {{cite web |url=http://tolweb.org/Acariformes/2563 | first= Heather |last=Proctor |date=August 9, 1998 |title=Acariformes. The "mite-like" mites |website= tolweb.org |publisher= Tree of Life Web Project}}
** {{cite web |url=http://tolweb.org/Parasitiformes |first1= David Evans | last1= Walter|title= Parasitiformes. Holothyrans, ticks and mesostigmatic mites |date=December 13, 1996 |website= tolweb.org |publisher= Tree of Life Web Project}}
* {{cite web|url=http://www.lucidcentral.com/keys/lwrrdc/public/aquatics/aghydra/html/about.htm |title=Key to Families and Subfamilies of Water Mites (Hydracarina) in Australia |first1= Mark | last1= Harvey | first2= Heather |last2= Proctor |website= lucidcentral.com |url-status=dead |archiveurl= https://web.archive.org/web/20060825162404/http://www.lucidcentral.com/keys/lwrrdc/public/Aquatics/aghydra/html/about.htm |archivedate=2006-08-25 }}
* {{cite web|url=http://www.lucidcentral.org/keys/v3/mites/Invasive_Mite_Identification/key/Whole_site/Home_whole_key.html |title=Invasive Mite Identification |first1= David Evans |website= lucidcentral.org | last1= Walter| date=September 15, 2006 |publisher=[[Colorado State University]], USDA/APHIS/PPQ Center for Plant Health Science and Technology |url-status=dead |archiveurl= https://web.archive.org/web/20070313140656/http://www.lucidcentral.org/keys/v3/mites/Invasive_Mite_Identification/key/Whole_site/Home_whole_key.html |archivedate=March 13, 2007 }}
* [http://www.jove.com/index/details.stp?ID=737 Non-invasive 3D-visualization with sub-micron resolution using synchrotron-X-ray-tomography], jove.com

{{Arachnida}}
{{Acari}}

{{Taxonbar|from=Q19137}}
{{Authority control}}

[[Category:Arthropod subclasses]]
[[Category:Acari| ]]
[[Category:Extant Early Devonian first appearances]]

Latest revision as of 03:59, 16 April 2022

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