Editing Bdelloidea

{{Short description|Class of parthenogenetic freshwater rotifers}}
{{For|the mite superfamily|Bdelloidea (mite)}}
{{Automatic taxobox
| fossil_range = {{Fossil range|Miocene|present}}
| name = Bdelloid rotifers
| image = Bdelloid.JPG
| image_caption = SEM showing morphological variation of bdelloid rotifers and their jaws
| taxon = Bdelloidea
| authority = [[Charles Thomas Hudson|Hudson]], 1884
| subdivision_ranks = 
| subdivision = 
}}

'''Bdelloidea''' {{IPAc-en|ˈ|d|ɛ||l|ɔɪ|d|i|ə}} (from [[Greek language|Greek]] βδέλλα, ''bdella'' 'leech') is a [[Class (biology)|class]] of [[rotifer]]s found in [[freshwater]] habitats all over the world.  There are over 450 described [[species]] of bdelloid rotifers (or 'bdelloids'),<ref>{{Cite book|author=Donner, Josef |year=1965 |chapter=Ordnung Bdelloidea (Rotatoria, Rädertiere) |title=Bestimmungsbücher zur Bodenfauna Europas, volume 6|location=Berlin |publisher=Akademie Verlag |oclc=6733231}} and {{Cite journal|author=Segers, Hendrik |year=2007 |title=Annotated checklist of the rotifers (Phylum Rotifera), with notes on nomenclature, taxonomy and distribution |journal=Zootaxa |volume=1590 |pages=3–104}} [http://www.mapress.com/j/zt/article/view/3617 Abstract].</ref> distinguished from each other mainly on the basis of [[Morphology (biology)|morphology]].<ref name="Segers2007">{{cite book |last= Segers |first= Hendrick |date= 2007 |title= Annotated checklist of the rotifers (Phylum Rotifera), with notes on nomenclature, taxonomy and distribution |location= Auckland|publisher= Magnolia Press |isbn= 978-1-86977-129-4 | url=http://www.mapress.com/zootaxa/2007f/zt01564p104.pdf }}</ref> The main characteristics that distinguish bdelloids from [[#Evolutionary Relationships|related groups of rotifers]] are exclusively [[parthenogenesis|parthenogenetic reproduction]] and the ability to survive in dry, harsh environments by entering a state of desiccation-induced [[dormancy]] ([[anhydrobiosis]]) at any life stage.<ref name="Ricci2000">{{Cite journal | author=Ricci, Claudia | year=2000 | title=Key to the identification of the genera of bdelloid rotifers | journal=Hydrobiologia | volume=418 | pages=73–80 | doi=10.1023/A:1003840216827 | s2cid=44054669 }}</ref> They are often referred to as "ancient asexuals" due to their unique asexual history that spans back to over 25 million years ago through fossil evidence.<ref>{{Cite journal|title = Bdelloid rotifers in Dominican amber: Evidence for parthenogenetic continuity|journal = Experientia|year=1992 |pages = 408–410 |volume = 48|issue = 4|doi = 10.1007/BF01923444|first1 = G. O.|last1=Poinar Jr.|first2 = Claudia |last2 = Ricci|s2cid = 13098228}}</ref> Bdelloid rotifers are microscopic organisms, typically between 150 and 700&nbsp;μm in length.<ref name=Ricci2000 />  Most are slightly too small to be seen with the naked eye, but appear as tiny white dots through even a weak [[hand lens]], especially in bright light. In June 2021, biologists reported the restoration of bdelloid rotifers after being frozen for 24,000 years in the [[Siberia]]n [[permafrost]].<ref name="NYT-20210607">{{cite news |last=Renault |first=Marion |title=This Tiny Creature Survived 24,000 Years Frozen in Siberian Permafrost - The microscopic animals were frozen when woolly mammoths still roamed the planet, but were restored as though no time had passed. |url=https://www.nytimes.com/2021/06/07/science/frozen-rotifers-siberia.html |date=7 June 2021 |work=[[the New York Times]] |accessdate=7 June 2021 }}</ref>

==Evolutionary relationships==
The [[phylum]] [[Rotifera]] traditionally included three [[Class (biology)|classes]]: Bdelloidea, [[Monogononta]] and [[Seisonidea]].<ref name="Kingetal2005">{{cite journal|last=King|first=Charles E.|author2=Ricci, Claudia |author3=Schonfeld, Justin |author4= Serra, Manuel |title=Evolutionary Dynamics of 'the' Bdelloid and Monogonont Rotifer Life-history Patterns|journal=Hydrobiologia|date=September 2005|volume=546|issue=1|pages=55–70|doi=10.1007/s10750-005-4102-9|citeseerx=10.1.1.455.6499|s2cid=25975998}}</ref>  Prior to 1990, [[phylogenetic]] studies based on morphology seemed to indicate that the sister group to bdelloid rotifers was Monogononta, with seisonid rotifers as an early-diverging [[Outgroup (cladistics)|outgroup]].<ref>{{cite journal|last=Wallace|first=Robert Lee|author2=Colburn, Rebecca Arlene |title=Phylogenetic relationships within phylum Rotifera: orders and genus Notholca|journal=Hydrobiologia|date=December 1989|volume=186–187|issue=1|pages=311–318|doi=10.1007/BF00048926|s2cid=20809514}}</ref>

[[File:Phylogenetic hypotheses for syndermata (modified from Lasek-Nesselquist 2012).png|thumb|upright=1.8| Cladograms showing alternative possible relationships within Syndermata (or Rotifera). [[Transcriptome]] results published in 2014 <ref name="Wey2014">{{cite journal|last=Wey-Fabrizius|first=Alexandra R.|author2=Herlyn, Holger |author3=Rieger, Benjamin |author4=Rosenkranz, David |author5=Witek, Alexander |author6=Welch, David B. Mark |author7=Ebersberger, Ingo |author8=Hankeln, Thomas |author9= Schmitz, Jürgen  |title=Transcriptome Data Reveal Syndermatan Relationships and Suggest the Evolution of Endoparasitism in Acanthocephala via an Epizoic Stage|journal=PLOS ONE|date=10 February 2014|volume=9|issue=2|pages=e88618|doi=10.1371/journal.pone.0088618 |pmid=24520404 |pmc=3919803|bibcode=2014PLoSO...988618W|doi-access=free}}</ref> support a refined version of the scenario in the bottom left, with Bdelloidea as a sister group to Seisonidea + Acanthocephala, and Monogononta as an outgroup. Cladograms modified from Fig. 3, Lasek-Nesselquist 2012.<ref>{{cite journal|last1=Lasek-Nesselquist|first1=Erica|title=A Mitogenomic Re-Evaluation of the Bdelloid Phylogeny and Relationships among the Syndermata|journal=PLOS ONE|date=23 August 2012|volume=7|issue=8|pages=e43554|doi=10.1371/journal.pone.0043554|pmid=22927990|pmc=3426538|bibcode=2012PLoSO...743554L|doi-access=free}}</ref> ]]

Modern [[molecular phylogenetic]] studies demonstrate that this classic understanding of 'Rotifera' is incomplete ([[paraphyletic]]), because it omits a fourth [[clade]] of closely related organisms: the [[Acanthocephala]], or thorny-headed worms.<ref>{{cite journal|last=Welch|first=David B. Mark |year=2001 |title=Early contributions of molecular phylogenetics to understanding the evolution of Rotifera |journal=Hydrobiologia|volume=446|pages=315–322|doi=10.1023/A:1017502923286|s2cid=28895228 }}</ref>  Originally classified as a separate [[phylum]], molecular and morphological evidence accumulated between 1994 and 2014 to indicate that Acanthocephala forms a [[monophyletic]] group with Bdelloidea, Monogononta and Seisonidea.<ref name="Wey2014"/><ref name="Ahlrichs1997">{{cite journal|last=Ahlrichs|first=Wilko H. |year=1997 |title=Epidermal ultrastructure of ''Seison nebaliae'' and ''Seison annulatus'', and a comparison of epidermal structures within the Gnathifera |journal=Zoomorphology |volume=117|issue=1|pages=41–48|doi=10.1007/s004350050028|s2cid=24719056 }}</ref>  To accommodate this finding, some authors extend the term 'Rotifera' to include the highly modified, parasitic 'acanthocephalan rotifers' alongside bdelloid, monogonont and seisonid rotifers.<ref>{{cite book|last=Nielsen|first=Claus|title=Animal evolution : interrelationships of the living phyla|date=2012|publisher=Oxford University Press|location=Oxford|isbn=978-0199606030|edition=3rd}}</ref>  Others refer to the grouping of the four taxa as Syndermata, a term derived from their shared [[syncytial]] epidermis.<ref name="Ahlrichs1997" />

The position of Bdelloidea within Syndermata (or Rotifera) is not entirely clear.  Alternative possible phylogenetic relationships within the clade are illustrated by the accompanying cladograms.  As of 2014, the "most comprehensive phylogenomic analysis of syndermatan relationships" to date was based on [[transcriptome]] data from all four groups,<ref name="Wey2014"/> and provided "strong support" for the hypothesis illustrated in the bottom left of the figure, in which Seisonidea and Acanthocephala are sister taxa.  The study further indicated that the sister group to this taxon is Bdelloidea, whereas Monogononta is the [[Outgroup (cladistics)|outgroup]] to all three.  This would mean that the closest living relatives of bdelloid rotifers are not monogonont rotifers, as previously believed, but seisonid rotifers and acanthocephalans, despite their highly modified morphology.

==Classification and identification==
A 2016 [[Phylogeny|phylogenetic]] analysis of the gene order in the mitochondria suggests that Seisonidea and Acanthocephala are sister clades and that the Eurotatoria are the sister clade to this group, producing the cladogram below.<ref name=Sielaff2015>{{cite journal |last1=Sielaff |first1=M. |last2=Schmidt |first2=H. |last3=Struck |first3=T. H. |last4=Rosenkranz |first4=D. |last5=Mark Welch |first5=D. B. |last6=Hankeln |first6=T |last7=Herlyn |first7=H. |date=March 2016 |title=Phylogeny of Syndermata (syn. Rotifera): Mitochondrial gene order verifies epizoic ''Seisonidea'' as sister to endoparasitic ''Acanthocephala'' within monophyletic ''Hemirotifera'' |journal=Molecular Phylogenetics and Evolution |volume=96 |pages= 79–92|doi=10.1016/j.ympev.2015.11.017 |pmid=26702959 |bibcode=2016MolPE..96...79S }}</ref>

{{clade|{{clade
|label1=[[Syndermata]]
|1={{clade
   |1=[[Seisonida]]
   |2=[[Acanthocephala]]
   }}
|label2=[[Eurotatoria]]
|2={{clade
   |1='''Bdelloidea'''
   |2=[[Monogononta]]
   }}
}}|style=line-height:100%}}

Bdelloidea is a [[Class (biology)|class]] of the [[phylum]] [[Rotifera]], consisting of three orders: Philodinavida, Philodinida and Adinetida.<ref>{{Cite journal|author=Melone, Giulio|author2=Ricci, Claudia|name-list-style=amp|year=1995 |title=Rotatory apparatus in Bdelloids |journal=Hydrobiologia |volume=313 |issue=1 |pages=91–98 |doi=10.1007/BF00025935|s2cid=35937088}}</ref> These orders are divided into four [[Family (biology)|families]] and about 450 [[species]].<ref name="Donner1965">{{cite book |last= Donner |first= Josef |year= 1965 |title= Ordnung Bdelloidea |publisher= Akademie-Verlag |pages= 297|isbn= 9789031908851 }}</ref> Since these organisms are asexual the usual definition of a [[species]] as a group of organisms capable of creating fertile offspring is inapplicable, therefore the [[species problem|species concept]] in these organisms is based on a mixture of morphological and molecular data instead. DNA studies suggest that the diversity is much greater than the original morphological classifications suggest.<ref>{{Cite journal|author=Kaya, Murat|author2=Herniou, Elisabeth A.|author3=Barraclough, Timothy G.|author4=Fontaneto, Diego|name-list-style=amp|year=2009 |title=Inconsistent estimates of diversity between traditional and DNA taxonomy in bdelloid rotifers |journal=Organisms Diversity & Evolution |volume=9 |issue=1 |pages=3–12 |doi=10.1016/j.ode.2008.10.002|doi-access=free|bibcode=2009ODivE...9....3K }}</ref><ref>{{Cite journal|author=Fontaneto, Diego|author2=Kaya, Murat|author3=Herniou, Elisabeth A.|author4=Barraclough, Timothy G.|name-list-style=amp|year=2009 |title=Extreme levels of hidden diversity in microscopic animals (Rotifera) revealed by DNA taxonomy |journal=Molecular Phylogenetics and Evolution |volume=53 |issue=1 |pages=182–189|pmid=19398026 |doi=10.1016/j.ympev.2009.04.011|bibcode=2009MolPE..53..182F }}</ref>

Bdelloids can only be identified by eye while they are alive because many of the characteristics significant to classification are related to feeding and crawling; however, genetic identification of bdelloids is possible on dead individuals. Once preserved, the individuals contract into "blobs" which restricts analysis.<ref name="Shiel1995" /> There are currently three morphological identification methodologies, two of which are considered dated: Bartoš (1951)<ref name="Bartos1951">{{Cite journal|author=Bartoš, Emanuel |year=1951 | title=The Czechoslovak Rotatoria of the order Bdelloidea | journal=Mémoiresde la Société Zoologique Tchécoslovaque de Prague | volume=15 | pages=241–500 }}</ref> and Donner (1965).<ref name="Donner1965" /> The third method is a diagnostic key developed in 1995 by Shiel.<ref name="Shiel1995">{{cite book |last= Shiel |first= Russell J. |year= 1995 |title= A guide to identification of rotifers, cladocerans and copepods from Australian inland waters |location= Australia|publisher= Co-operative Research Centre for Freshwater Ecology |isbn=978-0-646-22410-7 }}</ref>

===Morphology===
[[File:Bdelloid body areas.gif|200px | thumb |Figure 1: [[Scanning electron microscope|SEM]] pictures of some species of the genus ''[[Rotaria]]'' with head (red), tail (white) and trunk (blue) areas highlighted]]
There are three main regions of the body of bdelloids: head, trunk and foot. The adjacent image depicts each area to show how body parts can be very different although they are named the same depending on the species involved. Bdelloids typically have a well-developed corona, divided into two parts, on a retractable head.

Some identifiable features of the bdelloids include :
*Well-developed foot glands<ref name="Shiel1995" />
*A mouth opening with a long [[oesophagus]]<ref name="Shiel1995" />
*Strong teeth (labelled by a tooth index)<ref name="Shiel1995" />
*Many [[cilia]]<ref name="Shiel1995" />
*Species-specific upper lip shape<ref name="Shiel1995" />
*Order-specific corona type<ref name="Ricci2000" />
**[[Philodinida]] consist of two ciliated discs
**[[Adinetida]] consist of a ventral ciliated field
**[[Philodinavida]] have a small corona

The bdelloid digestive and reproductive systems can be found within the trunk sections of their bodies, with the stomach being the most visible of the [[Organ (anatomy)|organs]]. In certain [[genera]], (''Habrotrocha'', ''Otostephanos'' and ''Scepanotrocha'') the bdelloid can actually be identified by the appearance of distinct spherical pellets within the stomach, which will be released as faeces. These pellets are a distinguishing characteristic since all the other genera release faeces as loose material.<ref name="Ricci2000" />

Most bdelloids retract the foot while they eat, but there are four genera that lack a foot: ''Adineta'', ''Bradyscela'', ''Henoceros'' and ''Philodinavus''. This affects not only how they feed but also how they crawl; for instance ''Adineta'' and ''Bradyscela'' slide whereas the other genera loop.<ref name="Ricci2000" />

==Behaviour==
[[File:Бделлоидная коловратка.webm|thumb|A bdelloid feeding]]
The behaviour of bdelloids can be split into four categories: feeding, locomotion, reproduction and stress-induced behaviours.

===Feeding===
The specific [[Rotifer#Feeding|feeding behaviour]] of bdelloids is varied but most use rings of cilia in the corona organ to create currents of water which blow food through the mouth to the [[Rotifer#Digestive system|mastax organ]] which has been adapted specifically for grinding food.<ref name="Klusemann1990">{{Cite journal | author=Klusemann, J. | author2=Kleinow, W. | author3=Peters, W.|year=1990 | title=The hard parts (trophi) of the rotifer mastax do contain chitin: evidence from studies on Brachionus plicatilis | journal=Histochem. Cell Biol. | volume=94 | issue = 3  | pages=277–283 | doi = 10.1007/bf00266628 | pmid=2401635 | s2cid=26501940 }}</ref> Food includes suspended bacteria, algae, detritus, and other things.

===Locomotion===
There appear to be three main methods of movement: [[free swimming]], inch-worming along a substrate, or [[Sessility (zoology)|sessility]]. Inch-worming (or crawling) involves taking alternate steps with the head and tail, as do certain [[leech]]es, which gives the group their name ([[Greek language|Greek]] ''βδέλλα'' or ''bdella'', meaning leech).

===Reproduction===
Bdelloids are of interest in the study of the [[evolution of sex]] because a male has never been observed,<ref>{{Cite journal|title = Ancient asexual scandals|journal = Trends in Ecology & Evolution|year=1996 |pages = 41–46|volume = 11|issue = 2|doi = 10.1016/0169-5347(96)81040-8|first1 = Olivia P.|last1 = Judson|first2 = Benjamin B.|last2 = Normark|pmid=21237759| bibcode=1996TEcoE..11...41J }}</ref> and females reproduce exclusively by [[parthenogenesis#Rotifers|parthenogenesis]],{{Broken anchor|date=2024-10-14|bot=User:Cewbot/log/20201008/configuration|target_link=parthenogenesis#Rotifers|reason=Anchor "parthenogenesis#Rotifers" links to a specific web page: "Rotifers". The anchor (Rotifers) [[Special:Diff/1251095294|has been deleted]].}} a form of asexual reproduction where embryos grow and develop without the need for fertilization; this is akin to the [[apomixis]] seen in some plants.<ref>{{cite magazine| author = Milius, Susan | title=Bdelloids: No sex for over 40 million years |date=1 November 2002|magazine=Science News |url=http://www.thefreelibrary.com/Bdelloids%3A+No+sex+for+over+40+million+years.-a062685144  |access-date=6 November 2016 }}</ref> Each individual has paired [[gonad]]s. Despite having been asexual for millions of years, they have diversified into more than 450 species and are fairly similar to other sexually reproducing rotifer species.

However, a new study provided evidence for interindividual genetic exchange and recombination in ''Adineta vaga'', a species previously thought to be anciently asexual.<ref>{{cite journal|last1= Vakhrusheva|first1= O.A.|last2= Mnatsakanova|first2= E.A.|last3= Galimov|first3= Y.R.|title=Genomic signatures of recombination in a natural population of the bdelloid rotifer Adineta vaga |date=18 December 2020|journal=Nature|volume= 11|issue= 1|page= 6421|doi= 10.1038/s41467-020-19614-y|pmid= 33339818|pmc= 7749112|bibcode= 2020NatCo..11.6421V}}</ref>

''Adineta vaga'' is capable of carrying out [[DNA repair]] by a nonreductional [[meiosis]].<ref name = Terwagne2022>{{cite journal|pmid=36449626 |date=2022 |last1=Terwagne |first1=M. |last2=Nicolas |first2=E. |last3=Hespeels |first3=B. |last4=Herter |first4=L. |last5=Virgo |first5=J. |last6=Demazy |first6=C. |last7=Heuskin |first7=A. C. |last8=Hallet |first8=B. |last9=Van Doninck |first9=K. |title=DNA repair during nonreductional meiosis in the asexual rotifer Adineta vaga |journal=Science Advances |volume=8 |issue=48 |pages=eadc8829 |doi=10.1126/sciadv.adc8829 |pmc=9710870 |bibcode=2022SciA....8C8829T }}</ref> [[Germline]] DNA repair occurs in a specific period of [[oogenesis]] during which [[homologous chromosome]]s take on a meiotic-like juxtaposed configuration.<ref name = Terwagne2022/>  This germline DNA repair results in accurate reconstitution of the genetic material transmitted to offspring.

=== Evolution of obligate parthenogenetic reproduction ===
In 2003, the mode of asexual reproduction in the bdelloid rotifers was wholly unknown.<ref>{{Cite journal|first1 = Jean-Christophe|last1 = Simon|first2 = François|last2 = Delmotte|first3 =Claude |last3 = Rispe|first4 = Teresa|last4 = Crease|title = Phylogenetic relationships between parthenogens and their sexual relatives: the possible routes to parthenogenesis in animals|journal = Biological Journal of the Linnean Society|year=2003 |pages=151–163 |volume = 79|issue = 1|doi = 10.1046/j.1095-8312.2003.00175.x|doi-access = free}}</ref> One theory of how obligate [[parthenogenesis]] arose in bdelloid rotifers was that parthenogenic lineages lost the ability to respond to sex-inducing signal, which is why these lineages retained their asexuality.<ref>{{Cite journal|title = Obligate asex in a rotifer and the role of sexual signals|journal = Journal of Evolutionary Biology|year = 2008|pages = 287–293|volume = 21|issue = 1|doi = 10.1111/j.1420-9101.2007.01437.x|first =Claus-Peter |last = Stelzer|pmid=17995949|doi-access = free}}</ref> The obligate parthenogenetic strains of bdelloid rotifers produce a sex-inducing signal but have lost the ability to respond to that signal. It was later discovered that the inability to respond to sex-inducing signals in [[Parthenogenesis|obligate parthenogens]] was caused by simple [[Mendelian inheritance]] of the gene ''op.'' <ref>{{Cite journal|title = Loss of Sexual Reproduction and Dwarfing in a Small Metazoan|journal = PLOS ONE|date = 2010-09-20|pmc = 2942836|pmid = 20862222|pages = e12854|volume = 5|issue = 9|doi = 10.1371/journal.pone.0012854|first1 = Claus-Peter|last1 = Stelzer|first2 = Johanna|last2 = Schmidt|first3 = Anneliese|last3 = Wiedlroither|first4 = Simone|last4 = Riss|bibcode = 2010PLoSO...512854S|doi-access = free}}</ref>

===Stress-induced behaviour===
[[File:Rotifer entering inert state.ogg|thumb|Video of a rotifer transforming into a xerosome]]
Bdelloids are able to survive environmental stresses by entering a state of dormancy known as [[anhydrobiosis]] which enables the organism to rapidly dehydrate and thus resist desiccation. While preparing for this dormant state many metabolic processes are adjusted to equate for the change in state; e.g. the production of protective chemicals.<ref name="Crowe1971">{{Cite journal|author=Crowe, John H. |year=1971 |title=Anhydrobiosis: an unsolved problem |journal= American Naturalist | volume=105 |issue=946 | pages=563–573 | doi=10.1086/282745 | jstor=2459752|s2cid=84512992 }}</ref> The bdelloid can remain in this state, which is known as a 'xerosome' until the return of a sufficient amount of water, at which point they will rehydrate and become active within hours. Hatching of the young will only occur when conditions are at their most favourable.<ref name="Guidetti2002" /> These forms of dormancy are also known as [[cryptobiosis]] or quiescence. Bdelloids have been known to survive in this state for up to 9 years while waiting for favourable conditions to return.<ref name="Guidetti2002">{{Cite journal |author=Guidetti, Roberto |author2=Jönsson, K. Ingemar |name-list-style=amp|year=2002 |title= Long-term anhydrobiotic survival in semi-terrestrial micrometazoans |journal= Journal of Zoology |volume=257 |issue=2 | pages=181–187 | url=http://lup.lub.lu.se/luur/download?func=downloadFile&recordOId=147480&fileOId=625132 | doi=10.1017/s095283690200078x|citeseerx=10.1.1.630.9839 }}</ref>  In addition to surviving desiccation through anhydrobiosis, desiccation stress on two bdelloid species actually helped to maintain fitness and even improved their species [[fecundity]].<ref>{{Cite journal|author=Ricci, Claudia|author2=Fontaneto, Diego|name-list-style=amp|year=2009 |title=The importance of being a bdelloid: ecological and evolutionary consequences of dormancy |journal=Italian Journal of Zoology |volume=76 |issue=3 |pages=240–249 |doi=10.1080/11250000902773484|doi-access=free }}</ref> The rotifers that were consistently kept hydrated fared worse than those desiccated and rehydrated.<ref>{{Cite journal|first1 = Claudia|last1 = Ricci|first2 = Manuela|last2 = Caprioli|first3 = Diego|last3 = Fontaneto|year=2007 |title =Stress and fitness in parthenogens: is dormancy a key feature for bdelloid rotifers? |journal = BMC Evolutionary Biology |pmc = 1963474|pmid = 17767737|pages = S9|volume = 7|issue = Suppl 2|doi = 10.1186/1471-2148-7-S2-S9 | doi-access=free | bibcode=2007BMCEE...7S...9R }}</ref>

Bdelloidea have evolved a unique mechanism to help overcome one of the major perils of asexual reproduction.  According to the [[Red Queen hypothesis]] of [[Coevolution|co-evolution]], obligate asexuals will be driven extinct by rapidly changing parasites and pathogens, because they cannot change their genotypes quickly enough to keep up in this never-ending race.  In populations of bdelloid rotifers, however, many parasites are destroyed during periods of extended desiccation.<ref>{{Cite journal|title = Anciently Asexual Bdelloid Rotifers Escape Lethal Fungal Parasites by Drying Up and Blowing Away|journal = Science|date = 29 January 2010 |pmid = 20110504|pages = 574–576|volume = 327|issue = 5965|doi = 10.1126/science.1179252|first1 = Christopher G.|last1 = Wilson|first2 = Paul W.|last2 = Sherman|bibcode = 2010Sci...327..574W|s2cid = 43898914}}</ref>  Moreover, desiccated bdelloid rotifers are easily blown away from parasite-infested habitats by wind, and establish new, healthy populations elsewhere, which allows them to escape the Red Queen by moving in time and space instead of using sex to change their genotype.<ref>{{Cite journal|title = Spatial and temporal escape from fungal parasitism in natural communities of anciently asexual bdelloid rotifers|journal = Proceedings of the Royal Society B|date = 22 August 2013|pmid = 23825214|pages = 20131255|volume = 280|issue = 1765|doi = 10.1098/rspb.2013.1255|first1 = Christopher G.|last1 = Wilson|first2 = Paul W.|last2 = Sherman|pmc=3712457}}</ref>

When these creatures recover from desiccation, it has been shown that they undergo a potentially unique genetic process where [[horizontal gene transfer]] occurs,{{citation needed|date=August 2018}} resulting in a significant proportion of the bdelloid genome, up to 10%, having been obtained through horizontal gene transfer from bacteria, fungi and plants.<ref name="Boschetti2011">{{Cite journal | author=Boschetti, Chiara | author2=Pouchkina-Stantcheva, Natalia | author3=Hoffmann, Pia | author4=Tunnacliffe, Alan | name-list-style=amp| title= Foreign genes and novel hydrophilic protein genes participate in the desiccation response of the bdelloid rotifer ''Adineta ricciae'' |journal= The Journal of Experimental Biology | volume=214 | year=2011 | pages=59–68 | doi=10.1242/jeb.050328 | issue=1 | pmid=21147969| doi-access=free }}</ref> How and why horizontal gene transfer occur in bdelloids is under much debate at present; particularly with regards to possible connections between the foreign genes and the desiccation process as well as possible connections to bdelloids' ancient asexuality.

When they desiccate completely, their DNA breaks up into many pieces. And when they come back to life after being rehydrated, it creates an opportunity for alien DNA fragments to enter their genome. This process was improved 60 million years ago when they captured a bacterial gene this way, which gave them a new gene regulatory system. The new system was used to keep [[Transposable element|transposons]] in check.<ref>[https://www.laboratoryequipment.com/583869-60-M-Year-Old-Bacteria-Sheds-Light-on-New-DNA-Modification-System-in-Animals/ 60-M-Year-Old Bacteria Shed Light on 'New' DNA Modification System in Animals]</ref>

Bdelloid rotifers are extraordinarily resistant to damage from [[ionizing radiation]] due to the same DNA-preserving adaptations used to survive dormancy.<ref name="Gladyshev20081">{{Cite journal | author=Gladyshev, Eugene | author2=Meselson, Matthew | name-list-style=amp| title= Extreme resistance of bdelloid rotifers to ionizing radiation | journal= Proceedings of the National Academy of Sciences | volume=105 | date=1 April 2008 | pages=5139–5144 | doi=10.1073/pnas.0800966105 | issue=13 | pmid=18362355 | pmc=2278216| bibcode=2008PNAS..105.5139G | doi-access=free }}</ref>  These adaptations include an extremely efficient mechanism for repairing DNA double-strand breaks.<ref name="pmid25105197">{{Cite journal |vauthors=Hespeels B, Knapen M, Hanot-Mambres D, Heuskin AC, Pineux F, LUCAS S, Koszul R, Van Doninck K |title=Gateway to genetic exchange? DNA double-strand breaks in the bdelloid rotifer Adineta vaga submitted to desiccation |journal=J. Evol. Biol. |volume=27 |issue=7 |pages=1334–45 |date=July 2014  |pmid=25105197 |doi= 10.1111/jeb.12326|url=https://hal-pasteur.archives-ouvertes.fr/pasteur-01420001/file/hespeels%20et%20al%202014.pdf |doi-access=free }}</ref>  This repair mechanism was studied in two Bdelloidea species, ''Adineta vaga'',<ref name="pmid25105197" /> and ''Philodina roseola''.<ref name="pmid18362354">{{cite journal |author=Welch, David B. Mark |author2=Welch, Jessica L. Mark |author3=Meselson, Matthew |name-list-style=amp|title=Evidence for degenerate tetraploidy in bdelloid rotifers |journal=Proceedings of the National Academy of Sciences |volume=105 |issue=13 |pages=5145–9 |date=1 April 2008 |pmid=18362354 |pmc=2278229 |doi=10.1073/pnas.0800972105 |bibcode=2008PNAS..105.5145M |doi-access=free }}</ref> and appears to involve mitotic recombination between homologous DNA regions within each species.

=== Horizontal gene transfer ===
Large-scale horizontal transfer of bacterial, plant and fungal genes into bdelloid rotifers<ref>{{Cite journal|last1=Gladyshev|first1=Eugene A.|last2=Meselson|first2=Matthew|last3=Arkhipova|first3=Irina R.|date=2008-05-30|title=Massive horizontal gene transfer in bdelloid rotifers|journal=Science|volume=320|issue=5880|pages=1210–1213|doi=10.1126/science.1156407|issn=1095-9203|pmid=18511688|url=http://nrs.harvard.edu/urn-3:HUL.InstRepos:3120157|bibcode=2008Sci...320.1210G|s2cid=11862013|url-access=subscription}}</ref> has been documented, and may represent an important factor in bdelloid evolution.

==Gallery==
<gallery>
File:Bdelloidea1 w.jpg|Lateral view of a bdelloid.
File:Bdelloidea2 w.jpg|Frontal view of a bdelloid's corona.
File:Bdelloid rotifer (2009 04 11).jpg|Lateral view of a bdelloid.
File:Bdelloid Rotifer.jpg|Lateral view of a bdelloid.
File:Bdelloid Rotifier - Danelle Vivier.jpg|Lateral view of a bdelloid in algae-rich water
File:Philodina sp. (Classː Bdelloidea) - Rotifera.jpg|Specimen of the genus ''[[Philodina]]''
</gallery>

== References ==
{{Reflist|2}}

==External links==
{{Commons category|Bdelloidea}}
* [http://www.ucmp.berkeley.edu/phyla/rotifera/rotifera.html Introduction to rotifers]
* [http://www.rotifera.hausdernatur.at Rotifer World Catalog, by Jersabek C.D. & Leitner M.F.]
* [http://judson.blogs.nytimes.com/2008/06/03/the-weird-sisters/ The Weird Sisters] {{Webarchive|url=https://web.archive.org/web/20151001155042/http://judson.blogs.nytimes.com/2008/06/03/the-weird-sisters/ |date=2015-10-01 }}
* [http://www.sciencenews.org/articles/20000520/fob6.asp Bdelloids: No sex for over 40 million years] {{Webarchive|url=https://web.archive.org/web/20080423232627/http://www.sciencenews.org/articles/20000520/fob6.asp |date=2008-04-23 }}
* [https://web.archive.org/web/20061030024212/http://www.harvardmagazine.com/on-line/1100113.html An Evolutionary Scandal, from Harvard Magazine]
* [http://biology.plosjournals.org/perlserv/?request=get-document&doi=10.1371/journal.pbio.0050099 Who Needs Sex (or Males) Anyway?]
* [https://timesofglob.com/world-news/tiny-creature-comes-back-to-life-after-24000-years-in-siberian-deep-freeze/ Tiny Creature Comes Back To Life After 24,000 Years In Siberian Deep Freeze] {{Webarchive|url=https://web.archive.org/web/20210628060357/https://timesofglob.com/world-news/tiny-creature-comes-back-to-life-after-24000-years-in-siberian-deep-freeze/ |date=2021-06-28 }}

{{Taxonbar|from=Q430301}}

[[Category:Bdelloidea| ]]
[[Category:Eurotatoria]]
[[Category:Radiodurants]]
[[Category:Protostome classes]]