Springtail

{{Short description|Class of arthropods}} {{Automatic taxobox | fossil_range = {{Fossil range|Early Devonian | present}} | image = Orchesella cincta.jpg | image_caption = ''Orchesella cincta'' | display_parents = 2 | taxon = Collembola | authority = Lubbock, 1871 | subdivision_ranks = Orders | subdivision = * Entomobryomorpha * Poduromorpha * Symphypleona * Neelipleona <small>(disputed)</small> | synonyms = * Oligentoma * Oligoentoma | synonyms_ref = <ref>{{cite book |last=Gillott |first=Cedric |year=2005 |edition=3rd |title=Entomology |publisher=Springer |location=Berlin |isbn=978-0-306-44967-3 |chapter=Apterygote hexapods |editor-last=Gillott |editor-first=Cedric |pages=113–125 |doi=10.1007/1-4020-3183-1_5 |chapter-url=https://archive.org/details/gillott-2005 |access-date=4 November 2025 }}</ref> }}

'''Springtails''' (class '''Collembola''') form the largest of the three lineages of modern hexapods that are no longer considered insects, the others being Protura and Diplura. Although the three lineages are sometimes grouped together in a class called Entognatha because they have internal mouthparts, they do not appear to be any more closely related to one another than they are to insects, which have external mouthparts.<ref>{{cite journal |last1=Luan |first1=Yun-xia |last2=Mallatt |first2=Jon M. |last3=Xie |first3=Rong-dong |last4=Yang |first4=Yi-ming |last5=Yin |first5=Wen-ying |title=The phylogenetic positions of three basal-hexapod groups (Protura, Diplura, and Collembola) based on ribosomal RNA gene sequences |journal=Molecular Biology and Evolution |date=July 2005 |volume=22 |issue=7 |pages=1579–1592 |doi=10.1093/molbev/msi148 |url=https://www.researchgate.net/publication/7893037 |access-date=4 November 2025 }}</ref> There are more than 9000 species.<ref>{{cite web |last1=Bellinger |first1=Peter F. |last2=Christiansen |first2=Kenneth A. |last3=Janssens |first3=Frans |date=2024 |title=Checklist of the Collembola of the world |url=http://www.collembola.org |access-date=4 November 2025 }}</ref>

Springtails are omnivorous, free-living organisms that prefer moist conditions.<ref name="Hopkin1997">{{cite book |last=Hopkin |first=Stephen P. |title=Biology of the springtails (Insecta: Collembola) |publisher=Oxford University Press |year=1997 |location=Oxford, United Kingdom |url=https://fr.1lib.sk/book/933206/06cab9 |access-date=5 November 2025 }}</ref> They do not directly engage in the decomposition of organic matter, but contribute to it indirectly through the fragmentation of organic matter<ref>{{cite journal |last1=Saur |first1=Étienne |last2=Ponge |first2=Jean-François |title=Alimentary studies on the Collembolan ''Paratullbergia callipygos'' using transmission electron microscopy |journal=Pedobiologia |date=September 1988 |volume=31 |issue=5–6 |pages=355–380 |doi=10.1016/S0031-4056(23)02274-6 |url=https://www.academia.edu/52490540 |access-date=5 November 2025 }}</ref> and the control of soil microbial communities.<ref>{{cite journal |last1=Thimm |first1=Torsten |last2=Hoffmann |first2=Andrea |last3=Borkott |first3=Heinz |last4=Munch |first4=Jean Charles |last5=Tebbe |first5=Christoph C. |date=1 July 1998 |title=The gut of the soil microarthropod ''Folsomia candida'' (Collembola) is a frequently changeable but selective habitat and a vector for microorganisms |journal=Applied and Environmental Microbiology |volume=64 |issue=7 |pages=2660–2669 |doi=10.1128/AEM.64.7.2660-2669.1998 |pmid=9647845 |pmc=106441|bibcode=1998ApEnM..64.2660T |url=https://www.researchgate.net/publication/13636614 |access-date=5 November 2025 }}</ref><ref>{{cite journal |last1=Lartey |first1=Robert T. |last2=Curl |first2=Elroy A. |last3=Peterson |first3=Curt M. |date=January 1994 |title=Interactions of mycophagous Collembola and biological control fungi in the suppression of ''Rhizoctonia solani'' |journal=Soil Biology and Biochemistry |volume=26 |issue=1 |pages=81–88 |doi=10.1016/0038-0717(94)90198-8 |url=https://fr.1lib.sk/book/48531517/b4cec7 |access-date=5 November 2025 }}</ref> The word ''Collembola'' is from the Ancient Greek {{lang|grc|κόλλα}} {{Transliteration|grc|kólla}} meaning 'glue' and {{lang|grc|ἔμβολος}} {{Transliteration|grc|émbolos}} meaning 'peg'; this name was given due to the existence of the collophore, which was previously thought to stick to surfaces to stabilize the creature.<ref>{{cite book |last=Lubbock |first=John |year=1873 |title=Monograph of the Collembola and Thysanura |url=https://archive.org/stream/monographofcolle00lubb#page/36/mode/2up |page=36 |publisher=Ray Society |location=London, United Kingdom |author-link=John Lubbock, 1st Baron Avebury |access-date=5 November 2025 }}</ref>

Early DNA sequence studies<ref>{{cite journal |last1=Nardi |first1=Francesco |last2=Spinsanti |first2=Giacomo |last3=Boore |first3=Jeffrey L. |last4=Carapelli |first4=Antonio |last5=Dallai |first5=Romano |last6=Frati |first6=Francesco |title=Hexapod origins: monophyletic or paraphyletic? |journal=Science |date=21 March 2003 |volume=299 |issue=5614 |pages=1887–1889 |doi=10.1126/science.1078607 |pmid=12649480 |bibcode=2003Sci...299.1887N |s2cid=38792657 |url=https://www.researchgate.net/publication/10845956 |access-date=5 November 2025 }}</ref><ref>{{cite journal |last1=Delsuc |first1=Frédéric |last2=Phillips |first2=Matthew J. |last3=Penny |first3=David |title=Comment on "Hexapod Origins: Monophyletic or Paraphyletic?" |journal=Science |date=12 September 2003 |volume=301 |issue=5639 |page=1482 |doi=10.1126/science.1086558 |pmid=12970547 |s2cid=43942720 |url=https://fr.1lib.sk/book/43827822/4c8103 |access-date=5 November 2025 }}</ref><ref>{{cite journal |last1=Nardi |first1=Francesco |last2=Spinsanti |first2=Giacomo |last3=Boore |first3=Jeffrey L. |last4=Carapelli |first4=Antonio |last5=Dallai |first5=Romano |last6=Frati |first6=Francesco |title=Response to Comment on 'Hexapod Origins: Monophyletic or Paraphyletic?' |journal=Science |date=12 September 2003 |volume=301 |issue=5639 |page=1482 |doi=10.1126/science.1087632 |s2cid=82407120 |doi-access=free }}</ref> suggested that Collembola represent a separate evolutionary line from the other Hexapoda, but others disagree;<ref name="Gao2008">{{cite journal |last1=Gao |first1=Yan |last2=Bu |first2=Yun |last3=Luan |first3=Yun-Xia |title=Phylogenetic relationships of basal Hexapods reconstructed from nearly complete 18S and 28S rRNA gene sequences |journal=Zoological Science |date=25 November 2008 |volume=25 |issue=11 |pages=1139–1145 |doi=10.2108/zsj.25.1139 |pmid=19267625 |s2cid=10783597 |url=https://www.researchgate.net/publication/24183328 |access-date=5 November 2025 }}</ref> this seems to be caused by widely divergent patterns of molecular evolution among the arthropods.<ref>{{cite journal |last1=Hassanin |first1=Alexandre |title=Phylogeny of Arthropoda inferred from mitochondrial sequences: strategies for limiting the misleading effects of multiple changes in pattern and rates of substitution |journal=Molecular Phylogenetics and Evolution |date=January 2006 |volume=38 |issue=1 |pages=100–116 |doi=10.1016/j.ympev.2005.09.012 |pmid=16290034 |bibcode=2006MolPE..38..100H |url=https://www.academia.edu/458847 |access-date=5 November 2025 }}</ref> The adjustments of traditional taxonomic rank for springtails reflect the occasional incompatibility of traditional groupings with modern cladistics: when they were included with the insects, they were ranked as an order; as part of the Entognatha, they are ranked as a subclass. If they are considered within Hexapoda, they are elevated to full class status.

== Morphology == [[File:Isotoma Habitus.jpg|thumb|left|''Isotoma anglicana'' (Entomobryomorpha) with visible furcula]] [[File:Deutonura monticola - Flickr - Philippe Garcelon.jpg|thumb|left|''Deutonura monticola'' (Poduromorpha)]] Members of the Collembola are normally less than {{convert|6|mm|abbr=on}} long, have six or fewer abdominal segments, and possess a tubular appendage (the collophore or ventral tube) with reversible, sticky vesicles, projecting ventrally from the first abdominal segment.<ref name="Davies1927">{{cite journal |last1=Davies |first1=W. Maldwyn |title=On the tracheal system of Collembola, with special reference to that of ''Sminthurus viridis'', Lubb. |journal=Quarterly Journal of Microscopical Science |date=1 May 1927 |volume=S2-71 |issue=281 |pages=15–30 |doi=10.1242/jcs.s2-71.281.15 |url=https://archive.org/details/davies-1927 |access-date=6 November 2025 }}</ref> It is believed to be associated with fluid uptake and balance, excretion, and orientation of the organism itself.<ref name="Hopkin1997"/> Most species have an abdominal, tail-like appendage known as a furcula (or furca). It is located on the ventral side of the fourth abdominal segment and is folded beneath the body, held under tension by a small structure called the retinaculum (or tenaculum). When released, it snaps against the substrate, flinging the springtail into the air and allowing for rapid evasion and travel. All of this takes place in as little as 18&nbsp;milliseconds.<ref name="Piper">{{cite book |last=Piper |first=Ross |year=2007 |title=Extraordinary animals: an encyclopedia of curious and unusual animals |url=https://www.marefa.org/w/images/e/ea/Piper_Extraordinary_Animals-An_Encyclopedia_.pdf |publisher=Greenwood Press |location=Santa Barbara, California |isbn=978-0-313-33922-6 |author-link=Ross Piper |oclc=124074839 |access-date=6 November 2025 }}</ref><ref name="Hopkin1997"/>

Springtails also possess the ability to reduce their body size by as much as 30% through subsequent ecdyses (moulting) if temperatures rise high enough. The shrinkage is genetically controlled. Since warmer conditions increase metabolic rates and energy requirements in organisms, the reduction in body size is advantageous to their survival.<ref>{{cite journal |title=Random Sample |journal=Science |date=30 August 2013 |volume=341 |issue=6149 |page=945 |doi=10.1126/science.341.6149.945-a |url=https://fr.1lib.sk/book/83790217/b54dca |access-date=6 November 2025 }}</ref>

The Poduromorpha and Entomobryomorpha have an elongated body, while the Symphypleona and Neelipleona have a globular body. Collembola lack a tracheal respiration system, which forces them to respire through a porous cuticle, except for the two families Sminthuridae and Actaletidae, which exhibit a single pair of spiracles between the head and the thorax, leading to a rudimentary, although fully functional, tracheal system.<ref>{{cite book |url=https://books.google.com/books?id=x7vtCAAAQBAJ&dq=sminthuridae+actaletidae+spiracles&pg=PA264 |title=Multicellular animals. Volume II. The phylogenetic system of the Metazoa |first=Peter |last=Ax |date=9 March 2013 |publisher=Springer Science & Business Media |location=Berlin, Germany |isbn=978-3-662-10396-8 |via=Google Books |access-date=6 November 2025 }}</ref><ref name="Davies1927"/> The anatomical variance (life-form variation) present between different species partially depends on their vertical distribution across the various strata of terrestrial ecosystems.<ref>{{cite journal |last1=Salmon |first1=Sandrine |last2=Ponge |first2=Jean-François |last3=Gachet |first3=Sophie |last4=Deharveng |first4=Louis |last5=Lefebvre |first5=Noella |last6=Delabrosse |first6=Florian |title=Linking species, traits and habitat characteristics of Collembola at European scale |journal=Soil Biology and Biochemistry |date=August 2014 |volume=75 |pages=73–85 |doi=10.1016/j.soilbio.2014.04.002 |url=https://www.academia.edu/43676843 |access-date=6 November 2025 }}</ref> Surface-dwellers are generally larger, have darker pigments, longer antennae and a functioning furcula. Sub-surface-dwellers are usually unpigmented, have elongated bodies, and reduced furcula. They can be categorized into four main forms according to vertical distribution: atmobiotic, epedaphic, hemiedaphic, and euedaphic. Atmobiotic species inhabit macrophytes and litter surface. They are generally 8-10 millimeters (about ⅓ in) in length, pigmented, have long limbs, and a full set of ocelli (photoreceptors). Epedaphic species inhabit upper litter layers and fallen logs. They are slightly smaller and have less pronounced pigments, as well as less developed limbs and ocelli than the atmobiotic species. Hemiedaphic species inhabit the lower litter layers of decomposing organic material. They are 1-2 millimeters (about 1/16 in) in length, have dispersed pigmentation, shortened limbs, and a reduced number of ocelli. Euedaphic species inhabit upper mineral layers known as the humus horizon (or humipedon). They are smaller than hemiedaphic species; have soft, elongated bodies; lack pigmentation and ocelli; and have reduced or absent furca.<ref name="Coleman2015">{{cite book |doi=10.1016/B978-0-12-415955-6.00005-0 |chapter=Soil fauna: occurrence, biodiversity, and roles in ecosystem function |title=Soil microbiology, ecology and biochemistry |year=2015 |last1=Coleman |first1=David C. |last2=Wall |first2=Diana H. |pages=111–49 |isbn=978-0-12-415955-6 |editor-last=Paul |editor-first=Eldor A. |chapter-url=https://www.researchgate.net/publication/270902844 |access-date=6 November 2025 }}</ref><ref>{{cite journal |last1=Potapov |first1=Anton A. |last2=Semenina |first2=Eugenia E. |last3=Korotkevich |first3=Anastasiya Yu. |last4=Kuznetsova |first4=Natalia A. |last5=Tiunov |first5=Alexei V. |title=Connecting taxonomy and ecology: trophic niches of springtails as related to taxonomic identity and life forms |journal=Soil Biology and Biochemistry |date=October 2016 |volume=101 |pages=20–31 |doi=10.1016/j.soilbio.2016.07.002 |url=https://fr.1lib.sk/book/91252565/de9547 |access-date=6 November 2025 }}</ref><ref name="Jordana2020">{{cite journal |last1=Jordana |first1=Rafael |last2=Baquero |first2=Enrique |last3=Ledesma |first3=Enrique |last4=Sendra |first4=Alberto |last5=Ortuño |first5=Vicente M. |title=Poduromorpha (Collembola) from a sampling in the mesovoid shallow substratum of the Sierra de Guadarrama National Park (Madrid and Segovia, Spain): taxonomy and biogeography |journal=Zoologischer Anzeiger |date=March 2020 |volume=285 |issue=12 |pages=81–96 |doi=10.1016/j.jcz.2020.02.001 |bibcode=2020ZooAn.285...81J |s2cid=214349457 |url=https://fr.1lib.sk/book/111303681/7cd05e |access-date=6 November 2025 }}</ref>

Poduromorphs are characterized by their elongated bodies and conspicuous segmentation: they have three thoracic segments, six abdominal segments, including a well-developed prothorax with tergal chaetae, while the first thoracic segment in Entomobryomorpha is clearly reduced and bears no chaetae.<ref name="Jordana2020"/>

The digestive tract of springtails consists of three main components: foregut, midgut, and hindgut. The midgut is surrounded by a network of muscles and lined with a monolayer of columnar or cuboidal cells. Its function is to mix and transport food from the lumen into the hindgut through contraction. Many species of syntrophic bacteria, archaea, and fungi are present in the lumen. These different digestive regions have varying pH to support specific enzymatic activities and microbial populations. The anterior portion of the midgut and hindgut is slightly acidic (with a pH of approximately 6.0) while the posterior midgut portion is slightly alkaline (with a pH of approximately 8.0). Between the midgut and hindgut is an alimentary canal called the pyloric region, which is a muscular sphincter.<ref name="Hopkin1997"/> Malpighian tubules are absent.<ref>{{cite journal|last1=Konopová |first1=Barbora |last2=Kolosov |first2=Dennis |last3=O'Donnell |first3=Michael J. |url=https://www.researchgate.net/publication/333230796 |title=Water and ion transport across the eversible vesicles in the collophore of the springtail ''Orchesella cincta'' |journal=Journal of Experimental Biology |date=20 May 2019 |volume=222 |issue=10 |article-number=jeb200691 |doi=10.1242/jeb.200691 |access-date=6 November 2025 }}</ref>

== Genomics ==

Given their small size, springtails have been neglected in terms of genome analysis. They are one of the few arthropod groups that do not have high-quality reference genomes.<ref>{{cite journal |last1=Li |first1=Fei |last2=Wang |first2=Xianhui |last3=Zhou |first3=Xin |date=28 January 2025 |title=The genomics revolution drives a new era in entomology |url=https://www.annualreviews.org/content/journals/10.1146/annurev-ento-013024-013420 |journal=Annual Review of Entomology |language=en |volume=70 |issue=1 |pages=379–400 |doi=10.1146/annurev-ento-013024-013420 |pmid=39874145 |issn=0066-4170 |doi-access=free }}</ref> Even though some earlier genome sequences were produced, they do not meet modern standards (contig N50 > 1&nbsp;MB). One of the first genomes was presented in 2025, that of ''Orchesella flavescens,'' which is 270&nbsp;MB and was assembled into 6 chromosome-level scaffolds.<ref>{{cite journal |last1=McCulloch |first1=James |last2=Crowley |first2=Liam M. |last3=University of Oxford and Wytham Woods Genome Acquisition Lab |last4=Darwin Tree of Life Barcoding collective |last5=Wellcome Sanger Institute Tree of Life Management, Samples and Laboratory team |last6=Wellcome Sanger Institute Scientific Operations: Sequencing Operations |last7=Wellcome Sanger Institute Tree of Life Core Informatics team |last8=Tree of Life Core Informatics collective |last9=Darwin Tree of Life Consortium |date=26 September 2025 |title=The genome sequence of a springtail, ''Orchesella flavescens'' (C.Bourlet, 1839)[version 2; peer review: 2 approved, 1 approved with reservations] |url=https://wellcomeopenresearch.org/articles/10-138 |journal=Wellcome Open Research |language=en |volume=10 |article-number=138 |doi=10.12688/wellcomeopenres.23773.2 |doi-access=free |issn=2398-502X |pmc=12476540 }}</ref>

== Systematics and evolution == [[File:Allacma fusca on rotting wood.ogv|thumb|''Allacma fusca'' (Symphypleona) on rotting wood]] Traditionally, springtails were divided into the orders Arthropleona, Symphypleona, and occasionally also Neelipleona. The Arthropleona were divided into two superfamilies, the Entomobryoidea and the Poduroidea. However, recent phylogenetic studies show Arthropleona is paraphyletic.<ref>{{cite journal |last1=Xiong |first1=Yan |last2=Gao |first2=Yan |last3=Yin |first3=Wen-ying |last4=Luan |first4=Yun-xia |title=Molecular phylogeny of Collembola inferred from ribosomal RNA genes |journal=Molecular Phylogenetics and Evolution |date=December 2008 |volume=49 |issue=3 |pages=728–35 |doi=10.1016/j.ympev.2008.09.007 |pmid=18835455 |bibcode=2008MolPE..49..728X |url=https://fr.1lib.sk/book/35534358/e82f76 |access-date=6 November 2025 }}</ref><ref name="Leo2019">{{cite journal |last1=Leo |first1=Chiara |last2=Carapelli |first2=Antonio |last3=Cicconardi |first3=Francesco |last4=Frati |first4=Francesco |last5=Nardi |first5=Francesco |title=Mitochondrial genome diversity in Collembola: phylogeny, dating and gene order |journal=Diversity |date=17 September 2019 |volume=11 |issue=9 |article-number=169 |doi=10.3390/d11090169 |bibcode=2019Diver..11..169L |doi-access=free |hdl=11365/1079696 |hdl-access=free }}</ref><ref name="Sun2020">{{cite journal |last1=Sun |first1=Xin |last2=Ding |first2=Yinhuan |last3=Orr |first3=Michael C. |last4=Zhang |first4=Feng |title=Streamlining universal single-copy orthologue and ultraconserved element design: a case study in Collembola |journal=Molecular Ecology Resources |date=May 2020 |volume=20 |issue=3 |pages=706–17 |doi=10.1111/1755-0998.13146 |pmid=32065730 |bibcode=2020MolER..20..706S |s2cid=211133755 |url=https://www.researchgate.net/publication/339145746 |access-date=6 November 2025 }}</ref> Thus, the Arthropleona are abolished in modern classifications, and their superfamilies are raised in rank accordingly, being now orders Entomobryomorpha and the Poduromorpha. Technically, the Arthropleona are thus a partial junior synonym of the Collembola.<ref>{{cite web |url=https://www.collembola.org/ |title=Checklist of the Collembola |access-date=6 November 2025 }}</ref>

The term ''Neopleona'' is essentially synonymous with Symphypleona + Neelipleona.<ref>{{cite journal |last1=Sánchez-García |first1=Alba |last2=Engel |first2=Michael S. |title=Long-term stasis in a diverse fauna of Early Cretaceous springtails (Collembola: Symphypleona) |journal=Journal of Systematic Palaeontology |date=3 July 2017 |volume=15 |issue=7 |pages=513–37 |doi=10.1080/14772019.2016.1194575 |bibcode=2017JSPal..15..513S |s2cid=88641582 |url=https://fr.1lib.sk/book/87410321/d12a5e |access-date=6 November 2025 |hdl=10550/112294 |hdl-access=free }}</ref> The Neelipleona was originally seen as a particularly advanced lineage of Symphypleona, based on the shared global body shape, but the global body of the Neelipleona is realized in a completely different way than in Symphypleona. Subsequently, the Neelipleona were considered as being derived from the Entomobryomorpha. Analysis of 18S and 28S rRNA sequence data, though, suggests that they form the most ancient lineage of springtails, which would explain their peculiar apomorphies.<ref name="Gao2008"/> This phylogenetic relationship was also confirmed using a phylogeny based on mtDNA<ref name="Leo2019"/> and whole-genome data.<ref name="Sun2020"/>

The latest whole-genome phylogeny supporting four orders of Collembola:<ref name="Sun2020"/> {{clade |1={{clade |1=Neelipleona 60 px |2={{clade |1=Poduromorpha 60px |2={{clade |1=Symphypleona 60 px |2=Entomobryomorpha 60 px }} }} }} }}

Springtails are attested to since the Early Devonian. The fossil from {{Ma|400}}, ''Rhyniella praecursor'', is the oldest terrestrial arthropod, and was found in the famous Rhynie chert of Scotland.<ref>{{cite journal |last1=Dunlop |first1=Jason A. |last2=Garwood |first2=Russell J. |title=Terrestrial invertebrates in the Rhynie chert ecosystem |journal=Philosophical Transactions of the Royal Society B |date=18 December 2017 |volume=373 |issue=1739 |article-number=20160493 |doi=10.1098/rstb.2016.0493 |url=https://www.researchgate.net/publication/321910003 |access-date=6 November 2025 |pmc=5745329 }}</ref> Given its morphology resembles extant species quite closely, the radiation of the Hexapoda can be situated in the Silurian, {{Ma|420}} or more.<ref>{{cite journal |last1=Wang |first1=Yan-hui |last2=Engel |first2=Michael S. |last3=Rafael |first3=José A. |last4=Wu |first4=Hao-yang |last5=Rédei |first5=Dávid |last6=Xie |first6=Qiang |last7=Wang |first7=Gang |last8=Liu |first8=Xiao-guang |last9=Bu |first9=Wen-jun |title=Fossil record of stem groups employed in evaluating the chronogram of insects (Arthropoda: Hexapoda) |journal=Scientific Reports |date=13 December 2016 |volume=6 |article-number=38939 |doi=10.1038/srep38939 |doi-access=free |hdl=1808/22667 |hdl-access=free |pmc=5154178 }}</ref> Additional research concerning the coprolites (fossilized feces) of ancient springtails allowed researchers to track their lineages back some 412 million years.<ref name="Hopkin1997"/>

Fossil Collembola are rare. Instead, most are found in amber.<ref>{{cite journal |last=Mari Mutt |first=José A. |year=1983 |title=Collembola in amber from the Dominican Republic |journal=Proceedings of the Entomological Society of Washington |volume=85 |issue=3 |pages=575–87 |url=https://archive.org/details/biostor-75378 |access-date=7 November 2025 }}</ref> Even these are rare and many amber deposits carry few or no Collembola. The best deposits are from the early Eocene of Canada and Europe,<ref>{{cite journal |last1=Nel |first1=André |last2=De Ploëg |first2=Gaėl |last3=Milliet |first3=Jacqueline |last4=Menier |first4=Jean-Jacques |last5=Waller |first5=Alain |date=11 January 2004 |title=The French ambers: a general conspectus and the Lowermost Eocene amber deposit of Le Quesnoy in the Paris Basin |journal=Geologica Acta |volume=2 |issue=1 |pages=3–8 |doi=10.1344/105.000001628 |doi-access=free }}</ref> Miocene of Central America,<ref>{{cite journal |last1=Penney |first1=David |last2=McNeil |first2=Andrew |last3=Green |first3=David I. |last4=Bradley |first4=Robert S. |last5=Jepson |first5=James E. |last6=Withers |first6=Philip J. |last7=Preziosi |first7=Richard F. |date=17 October 2012 |title=Ancient Ephemeroptera-Collembola symbiosis fossilized in amber predicts contemporary phoretic associations |journal=PLOS ONE |volume=7 |issue=10 |article-number=e47651 |doi=10.1371/journal.pone.0047651 |pmid=23082186 |pmc=3474712 |bibcode=2012PLoSO...747651P |doi-access=free }}</ref> and the mid-Cretaceous of Burma and Canada.<ref>{{cite journal |last1=Christiansen |first1=Kenneth |last2=Nascimbene |first2=Paul |title=Collembola (Arthropoda, Hexapoda) from the mid Cretaceous of Myanmar (Burma) |journal=Cretaceous Research |date=June 2006 |volume=27 |issue=3 |pages=318–63 |doi=10.1016/j.cretres.2005.07.003 |bibcode=2006CrRes..27..318C |url=https://www.academia.edu/2626139 |access-date=7 November 2025 }}</ref> They display some unexplained characteristics: first, all but one of the fossils from the Cretaceous belong to extinct genera, whereas none of the specimens from the Eocene or the Miocene are of extinct genera; second, the species from Burma are more similar to the modern fauna of Canada than are the Canadian Cretaceous specimens.

There are now about 8,000 described species of Collembola.<ref name="Hopkin1997"/> However, to the light of ongoing developments in molecular methods and the rise of cryptic species unveiled by DNA barcoding within morphologically-described species,<ref>{{cite journal |last1=Zhang |first1=Bing |last2=Chen |first2=Ting-Wen |last3=Mateos |first3=Eduardo |last4=Scheu |first4=Stefan |last5=Schaefer |first5=Ina |title=Cryptic species in ''Lepidocyrtus lanuginosus'' (Collembola: Entomobryidae) are sorted by habitat type |journal=Pedobiologia |date=May 2018 |volume=68 |pages=12–9 |doi=10.1016/j.pedobi.2018.03.001 |url=https://www.researchgate.net/publication/324071644 |access-date=7 November 2025 }}</ref> it has been speculated that the global species richness of Collembola could be at least an order of magnitude greater than a previous estimate of 50,000 species.<ref>{{cite journal |last1=Cicconardi |first1=Francesco |last2=Fanciulli |first2=Pietro P. |last3=Emerson |first3=Brent C. |title=Collembola, the biological species concept and the underestimation of global species richness |journal=Molecular Ecology |date=November 2013 |volume=22 |issue=21 |pages=5282–96 |doi=10.1111/mec.12472 |url=https://www.academia.edu/17784282 |access-date=7 November 2025 }}</ref>

== Ecology ==

=== Eating behavior ===

Specific feeding (foraging) strategies and mechanisms are employed to match specific niches.<ref>{{cite journal |last1=Sechi |first1=Valentina |last2=D'Annibale |first2=Alessandra |last3=Ambus |first3=Per |last4=Sárossy |first4=Zsuzsa |last5=Krogh |first5=Paul Henning |last6=Eriksen |first6=Jørgen |last7=Holmstrup |first7=Martin |title=Collembola feeding habits and niche specialization in agricultural grasslands of different composition |journal=Soil Biology and Biochemistry |date=July 2014 |volume=74 |pages=31–38 |doi=10.1016/j.soilbio.2014.02.019 |bibcode=2014SBiBi..74...31S |url=https://www.researchgate.net/publication/260680327 |access-date=7 November 2025 }}</ref> Herbivorous and detritivorous species fragment biological material present in soil and leaf litter, supporting decomposition and increasing the availability of nutrients for plants and various species of microbes and fungi.<ref>{{cite journal |last1=Rusek |first1=Josef |title=Biodiversity of Collembola and their functional role in the ecosystem |journal=Biodiversity and Conservation |date=September 1998 |volume=7 |issue=9 |pages=1207–19 |doi=10.1023/a:1008887817883 |bibcode=1998BiCon...7.1207R |s2cid=22883809 |url=https://fr.1lib.sk/book/43218797/a22e8e |access-date=7 November 2025 |url-access=subscription }}</ref> Carnivorous species maintain populations of small invertebrates such as nematodes, rotifers, and other collembolan species.<ref name="Hopkin1997"/><ref name="Coleman2015"/> Springtails commonly consume fungal hyphae and spores, but also have been found to consume plant material and pollen, animal remains, colloidal materials, minerals and bacteria.<ref>{{cite journal |last1=Chen |first1=Benrong |last2=Snider |first2=Richard J. |last3=Snider |first3=Renate M. |date=March 1996 |title=Food consumption by Collembola from northern Michigan deciduous forest |journal=Pedobiologia |volume=40 |issue=2 |pages=149–61 |doi=10.1016/S0031-4056(24)00348-2 |url=https://archive.org/details/chen-et-al.-1996 |access-date=7 November 2025 }}</ref> Some springtail species like ''Anurophorus laricis'' can form mutually beneficial relationships with lichens, using them for shelter while feeding on surrounding free-living algae, which helps reduce competition between algae and lichens for space and nutrients.<ref>{{cite journal |author-link=André Aptroot |last1=Aptroot |first1=André |last2=Berg |first2=Matty P. |title=Collembola help lichens in competition with algae |journal=The Lichenologist |volume=36 |issue=2 |date=13 April 2004 |doi=10.1017/S0024282904014082 |pages=167–69 |bibcode=2004ThLic..36..167A |url=https://research.vu.nl/files/2014058/171904.pdf |access-date=7 November 2025 }}</ref> For finding their preferred food in light-free and complex environments like litter and soil layers Collembola use olfactory cues such as the odors emitted by fungi and other food sources as attractors.<ref>{{cite journal |last1=Bengsson |first1=Göran |last2=Erlandsson |first2=Ann |last3=Rundgren |first3=Sten |title=Fungal odour attracts soil Collembola |journal=Soil Biology and Biochemistry |volume=20 |issue=1 |year=1988 |doi=10.1016/0038-0717(88)90122-8 |pages=25–30 |url=https://fr.1lib.sk/book/51956251/204bcd |access-date=7 November 2025 |url-access=subscription }}</ref> Directional movement towards the food source has been recorded by image analysis in laboratory experiments.<ref>{{cite journal |last1=Auclerc |first1=Appoline |last2=Libourel |first2=Paul-Antoine |last3=Salmon |first3=Sandrine |last4=Bels |first4=Vincent |last5=Ponge |first5=Jean-François |title=Assessment of movement patterns in ''Folsomia candida'' (Hexapoda: Collembola) in the presence of food |journal=Soil Biology and Biochemistry |volume=42 |issue=4 |date=April 2010 |doi=10.1016/j.soilbio.2009.12.012 |pages=657–9 |url=https://www.academia.edu/44300794 |access-date=7 November 2025 }}</ref> However, other experiments showed that attraction and consumption were often, but not always correlated, and that preferred fungal strains were not necessarity those that optimize performances in reproduction (fitness), growth and survival.<ref>{{cite journal |last1=Sadaka-Laulan |first1=Nassima |last2=Ponge |first2=Jean-François |last3=Roquebert |first3=Marie-France |last4=Bury |first4=Edith |last5=Boumezzough |first5=Ali |title=Feeding preferences of the collembolan ''Onychiurus sinensis'' for fungi colonizing holm oak litter (''Quercus rotundifolia'' Lam.) |journal=European Journal of Soil Biology |volume=34 |issue=4 |date=September–December 1998 |doi=10.1016/S1164-5563(00)86660-1 |pages=179–88 |url=https://www.academia.edu/49355417 |access-date=7 November 2025 }}</ref>

=== Predators ===

Springtails are consumed by mesostigmatan mites in various families, including Ascidae, Laelapidae, Parasitidae, Rhodacaridae and Veigaiidae.<ref>{{cite journal |last=Koehler |first=Harmut K. |title=Predatory mites (Gamasina, Mesostigmata) |journal=Agriculture, Ecosystems & Environment |volume=74 |issue=1–3 |date=June 1999 |doi=10.1016/S0167-8809(99)00045-6 |pages=395–410 |url=https://fr.1lib.sk/book/46329860/33af3d |access-date=7 November 2025 }}</ref>

Cave-dwelling springtails are a food source for spiders and harvestmen in the same environment, such as the endangered harvestman ''Texella reyesi''.<ref>{{cite web |last=City of Austin and Travis County |date=3 September 2020 |title=Support for maintaining endangered status for the Bone Cave harvestman (''Texella reyesi'') |url=https://www.austintexas.gov/edims/document.cfm?id=347313 |access-date=7 November 2025 }}</ref>

Predators also include various soil centipedes.<ref>{{cite journal |last1=Bortolin |first1=Francesca |last2=Fusco |first2=Giuseppe |last3=Bonato |first3=Lucio |date=1 December 2018 |title=Comparative analysis of diet in syntopic geophilomorph species (Chilopoda, Geophilomorpha) using a DNA-based approach |url=https://fr.1lib.sk/book/104051102/b8a7f6 |journal=Soil Biology and Biochemistry |volume=127 |pages=223–9 |doi=10.1016/j.soilbio.2018.09.021 |bibcode=2018SBiBi.127..223B |issn=0038-0717 |access-date=7 November 2025 }}</ref>

Jumping, using the furcula as a spring, is the most common way to avoid predation,<ref>{{cite journal |last1=Reznikova |first1=Zhanna I. |last2=Panteleeva |first2=S. Ninel |date=September 2001 |title=Interaction of the ant ''Myrmica rubra'' L. as a predator with springtails (Collembola) as a mass prey |url=https://www.researchgate.net/publication/10614381 |journal=Doklady Biological Sciences |volume=380 |pages=475–7 |doi=10.1023/A:1012383607521 |access-date=7 November 2025 }}</ref> but jump escape cannot be performed easily in concealed environments such as litter, and more especially soil layers. To protect themselves against predators, some species without jumping abilities have evolved chemical defenses.<ref>{{cite book |url=https://books.google.com/books?id=HoMqBgAAQBAJ&dq=springtails+chemical+defenses&pg=PA44 |title=Insect molecular biology and ecology |first=Klaus H. |last=Hoffmann |date=19 December 2014 |publisher=CRC Press |location=Boca Raton, Florida |isbn=978-1-4822-3189-2 |page=44 |via=Google Books |access-date=7 November 2025 }}</ref>

=== Distribution ===

Springtails are cryptozoa frequently found in leaf litter and other decaying material,<ref name="Hopkin1997"/> where they are primarily detritivores and microbivores, and one of the main biological agents responsible for the control and the dissemination of soil microorganisms.<ref>{{cite journal |last=Ponge |first=Jean-François |date=June 1991 |title=Food resources and diets of soil animals in a small area of Scots pine litter |journal=Geoderma |volume=49 |issue=1–2 |pages=33–62 |doi=10.1016/0016-7061(91)90090-G |bibcode=1991Geode..49...33P |citeseerx=10.1.1.635.8529 |url=https://www.academia.edu/20508971 |access-date=7 November 2025 }}</ref> In a mature deciduous woodland in temperate climate, leaf litter and vegetation typically support 30 to 40 species of springtails, and in the tropics the number may be over 100.<ref name="Hopkin1997"/>

thumb|left|"Snow flea" [[File:CollembolaBMNH.jpg|thumb|right|upright|A species of Sminthurinae (Symphypleona: Sminthuridae)]]

In sheer numbers, they are reputed to be one of the most abundant of all macroscopic animals, with estimates of 100,000 individuals per square meter of ground,<ref>{{cite journal |last1=Ponge |first1=Jean-François |last2=Arpin |first2=Pierre |last3=Sondag |first3=Francis |last4=Delecour |first4=Ferdinand |title=Soil fauna and site assessment in beech stands of the Belgian Ardennes |journal=Canadian Journal of Forest Research |date=December 1997 |volume=27 |issue=12 |pages=2053–64 |doi=10.1139/x97-169 |bibcode=1997CaJFR..27.2053P |url=https://www.academia.edu/49973519 |access-date=7 November 2025 }}</ref> essentially everywhere on Earth where soil and related habitats (moss cushions, fallen wood, grass tufts, ant and termite nests) occur.<ref name="Ponge 1993">{{cite journal |last1=Ponge |first1=Jean-François |title=Biocenoses of Collembola in atlantic temperate grass-woodland ecosystems |journal=Pedobiologia |date=1993 |volume=37 |issue=4 |pages=223–244 |doi=10.1016/S0031-4056(24)00100-8 |bibcode=1993Pedob..37..223P |url=https://www.academia.edu/50930736 |access-date=7 November 2025 }}</ref> Only nematodes, crustaceans, and mites are likely to have global populations of similar magnitude, and each of those groups except mites is more inclusive. Though taxonomic rank cannot be used for absolute comparisons, it is notable that nematodes are a phylum and crustaceans a subphylum. Most springtails are small and difficult to see by casual observation, but some springtails, like snow fleas,<ref>{{cite journal |last1=Valle |first1=Barbara |last2=Porco |first2=David |last3=Skarżyński |first3=Dariusz |last4=Frati |first4=Francesco |last5=Caccianiga |first5=Marco |last6=Rodriguez-Prieto |first6=Ana |last7=Zeni |first7=Michele |last8=Gobbi |first8=Mauro |date=1 March 2024 |title=Alpine blooming of "snow fleas": the importance of snow for Alpine springtails (Hexapoda: Collembola) ecology and biodiversity |journal=Rendiconti Lincei, Scienze Fisiche e Naturali |language=en |volume=35 |issue=1 |pages=163–80 |doi=10.1007/s12210-023-01211-y |bibcode=2024RLSFN..35..163V |issn=1720-0776 |doi-access=free |hdl=2434/1043029 |hdl-access=free }}</ref> are readily observed on warm winter days when it is active and its dark color contrasts sharply with a background of snow.<ref>{{cite journal |last1=Lyford |first1=Walter H. |title=Overland migration of Collembola (''Hypogastrura nivicola'' Fitch) colonies |journal=The American Midland Naturalist |date=July 1975 |volume=94 |issue=1 |pages=205–9 |doi=10.2307/2424550 |jstor=2424550 |url=https://fr.1lib.sk/book/81982092/d7f694 |access-date=7 November 2025 }}</ref>

In addition, a few species routinely climb trees and form a dominant component of canopy fauna, where they may be collected by beating or insecticide fogging.<ref>{{cite journal |last1=Shaw |first1=Peter |last2=Ozanne |first2=Claire |last3=Speight |first3=Martin |last4=Palmer |first4=Imogen |title=Edge effects and arboreal Collembola in coniferous plantations |journal=Pedobiologia |date=October 2007 |volume=51 |issue=4 |pages=287–93 |doi=10.1016/j.pedobi.2007.04.010 |bibcode=2007Pedob..51..287S |url=https://fr.1lib.sk/book/49385694/ca5d0f |access-date=10 November 2025 }}</ref><ref>{{cite journal |last1=Zettel |first1=Joerg |last2=Zettel |first2=Ursula |last3=Egger |first3=Beatrice |title=Jumping technique and climbing behaviour of the collembolan ''Ceratophysella sigillata'' (Collembola: Hypogastruridae) |journal=European Journal of Entomology |date=15 April 2000 |volume=97 |issue=1 |pages=41–5 |doi=10.14411/eje.2000.010 |doi-access=free }}</ref> In temperate regions, a few species (e.g. ''Anurophorus laricis'', ''Entomobrya albocincta'', ''Xenylla xavieri'', ''Hypogastrura arborea'') are almost exclusively arboreal.<ref name="Ponge 1993"/> In tropical regions a single square meter of canopy habitat can support many species of Collembola.<ref name="Piper"/>

The main ecological factor driving the local distribution of species is the vertical stratification of the ecosystem: in woodland a continuous change in species assemblages can be observed from tree canopies to ground vegetation then to plant litter down to deeper soil horizons.<ref name="Ponge 1993"/> This is a complex factor embracing both nutritional and physiological requirements, together with behavioural trends,<ref name="Didden 1987">{{cite journal |last=Didden |first=Wim A. M. |date=March 1987 |title=Reactions of ''Onychiurus fimatus'' (Collembola) to loose and compact soil: methods and first results |journal=Pedobiologia |volume=30 |issue=2 |pages=93–100 |doi=10.1016/S0031-4056(23)00360-8 |bibcode=1987Pedob..30...93D |s2cid=258140448 |doi-access=free }}</ref> dispersal limitation<ref>{{cite journal |last1=Rodgers |first1=Denis J. |last2=Kitching |first2=Rodger L. |year=1998 |title=Vertical stratification of rainforest collembolan (Collembola: Insecta) assemblages: description of ecological patterns and hypotheses concerning their generation |journal=Ecography |volume=21 |issue=4 |pages=392–400 |doi=10.1111/j.1600-0587.1998.tb00404.x |bibcode=1998Ecogr..21..392R |citeseerx=10.1.1.476.6663 |s2cid=85133291 |url=https://fr.1lib.sk/book/42046928/275cc1 |access-date=10 November 2025 }}</ref> and probable species interactions.<ref>{{cite journal |last1=Widenfalk |first1=Lina A. |last2=Malmström |first2=Anna |last3=Berg |first3=Matty P. |last4=Bengtsson |first4=Jan |title=Small-scale Collembola community composition in a pine forest soil: overdispersion in functional traits indicates the importance of species interactions |journal=Soil Biology and Biochemistry |volume=103 |pages=52–62 |date=December 2016 |doi=10.1016/j.soilbio.2016.08.006 |url=https://www.researchgate.net/publication/306347013 |access-date=10 November 2025 |hdl=1871.1/a3a13719-8bc9-4233-9d2e-fa81b6093ec9 |hdl-access=free }}</ref> Some species have been shown to exhibit negative<ref>{{cite journal |last1=Bowden |first1=John |last2=Haines |first2=Ian H. |last3=Mercer |first3=D. |year=1976 |title=Climbing Collembola |journal=Pedobiologia |volume=16 |issue=4 |pages=298–312 |doi=10.1016/S0031-4056(23)02178-9 |bibcode=1976Pedob..16..298B |s2cid=258572651 |doi-access=free }}</ref> or positive<ref name="Didden 1987"/> gravitropism, which adds a behavioural dimension to this still poorly understood vertical segregation. Experiments with peat samples turned upside down showed two types of responses to disturbance of this vertical gradient, called ''stayers'' and ''movers''.<ref>{{cite journal |last1=Krab |first1=Eveline J. |last2=Oorsprong |first2=Hilde |last3=Berg |first3=Matty P. |last4=Cornelissen |first4=Johannes H. C. |date=December 2010 |title=Turning northern peatlands upside down: disentangling microclimate and substrate quality effects on vertical distribution of Collembola |journal=Functional Ecology |volume=24 |issue=6 |pages=1362–9 |doi=10.1111/j.1365-2435.2010.01754.x |doi-access=free |bibcode=2010FuEco..24.1362K }}</ref>

[[File:Sminthuridae sp 01.jpg|thumb|left|''Dicyrtomina'' sp. on leaf]]

As a group, springtails are highly sensitive to desiccation, because of their tegumentary respiration,<ref>{{cite journal |last1=Nickerl |first1=Julia |last2=Helbig |first2=Ralf |last3=Schulz |first3=Hans-Jürgen |last4=Werner |first4=Carsten |last5=Neinhuis |first5=Christoph |title=Diversity and potential correlations to the function of Collembola cuticle structures |journal=Zoomorphology |date=June 2013 |volume=132 |issue=2 |pages=183–95 |doi=10.1007/s00435-012-0181-0 |s2cid=14442664 |url=https://www.researchgate.net/publication/257425170 |access-date=10 November 2025 }}</ref> although some species with thin, permeable cuticles have been shown to resist severe drought by regulating the osmotic pressure of their body fluid.<ref>{{cite journal |last1=Holmstrup |first1=Martin |last2=Bayley |first2=Mark |date=November 2013 |title=''Protaphorura tricampata'', a euedaphic and highly permeable springtail that can sustain activity by osmoregulation during extreme drought |journal=Journal of Insect Physiology |volume=59 |issue=11 |pages=1104–10 |doi=10.1016/j.jinsphys.2013.08.015|pmid=24035747 |bibcode=2013JInsP..59.1104H |url=https://fr.1lib.sk/book/57718634/393017 |access-date=10 November 2025 }}</ref> The gregarious behaviour of Collembola, mostly driven by the attractive power of pheromones excreted by adults,<ref>{{cite journal |last=Verhoef |first=Herman A. |year=1984 |title=Releaser and primer pheromones in Collembola |journal=Journal of Insect Physiology |volume=30 |issue=8 |pages=665–70 |doi=10.1016/0022-1910(84)90052-0 |bibcode=1984JInsP..30..665V |url=https://www.academia.edu/55863820 |access-date=10 November 2025 }}</ref> gives more chance to every juvenile or adult individual to find suitable, better protected places, where desiccation could be avoided and reproduction and survival rates (thereby fitness) could be kept at an optimum.<ref>{{cite journal |last1=Benoit |first1=Joshua B. |last2=Elnitsky |first2=Michael A. |last3=Schulte |first3=Glen G. |last4=Lee |first4=Richard E. Jr |last5=Denlinger |first5=David L. |date=6 November 2008 |title=Antarctic Collembolans use chemical signals to promote aggregation and egg laying |journal=Journal of Insect Behavior |volume=22 |issue=2 |pages=121–33 |doi=10.1007/s10905-008-9159-7 |bibcode=2009JIBeh..22..121B |s2cid=914560 |url=https://www.researchgate.net/publication/225379648 |access-date=10 November 2025 }}</ref> Sensitivity to drought varies from species to species<ref>{{cite journal |last1=Prinzing |first1=Andreas |last2=D'Haese |first2=Cyrille A. |last3=Pavoine |first3=Sandrine |last4=Ponge |first4=Jean-François |date=February 2014 |title=Species living in harsh environments have low clade rank and are localized on former Laurasian continents: a case study of ''Willemia'' (Collembola) |journal=Journal of Biogeography |volume=41 |issue=2 |pages=353–65 |doi=10.1111/jbi.12188 |bibcode=2014JBiog..41..353P |s2cid=86619537 |url=https://www.researchgate.net/publication/259967386 |access-date=10 November 2025 }}</ref> and increases during ecdysis.<ref>{{cite journal |last=Verhoef |first=Herman A. |year=1981 |title=Water balance in Collembola and its relation to habitat selection: water content, haemolymph osmotic pressure and transpiration during an instar |journal=Journal of Insect Physiology |volume=27 |issue=11 |pages=755–60 |doi=10.1016/0022-1910(81)90065-2 |bibcode=1981JInsP..27..755V |url=https://fr.1lib.sk/book/48428519/a52afb |access-date=10 November 2025 }}</ref> Given that springtails moult repeatedly during their entire life (an ancestral character in Hexapoda) they spend much time in concealed micro-sites where they can find protection against desiccation and predation during ecdysis, an advantage reinforced by synchronized moulting.<ref>{{cite journal |last=Leinaas |first=Hans Petter |year=1983 |title=Synchronized moulting controlled by communication in group-living Collembola |journal=Science |volume=219 |issue=4581 |pages=193–5 |doi=10.1126/science.219.4581.193 |pmid=17841689|bibcode=1983Sci...219..193P |s2cid=41604935 |url=https://fr.1lib.sk/book/71524986/acd55c |access-date=10 November 2025 }}</ref> The high humidity of many caves also favours springtails and there are numerous cave adapted species,<ref>{{cite journal |last=Wilson |first=Jane M. |year=1982 |title=A review of world Troglopedetini (Insecta, Collembola, Paronellidae), including an identification table and descriptions of new species |journal=Cave Science: Transactions of the British Cave Research Association |volume=9 |issue=3 |pages=210–26 |url=https://www.researchgate.net/publication/321870154 |access-date=10 November 2025 |archive-date=2016-08-04 |archive-url=https://web.archive.org/web/20160804122528/http://www.wcmt.org.uk/sites/default/files/report-documents/Jane%20Wilson%20Howarth.pdf |url-status=live }}</ref><ref>{{cite journal |last1=Palacios-Vargas |first1=José G. |last2=Wilson |first2=Jane |year=1990 |title=''Troglobius coprophagus'', a new genus and species of cave collembolan from Madagascar with notes on its ecology |journal=International Journal of Speleology |volume=19 |issue=1–4 |pages=67–73 |doi=10.5038/1827-806x.19.1.6 |bibcode=1990IJSpe..19...67P |url=https://www.researchgate.net/publication/283327501 |access-date=10 November 2025 }}</ref> including one, ''Plutomurus ortobalaganensis'' living {{convert|1980|m|ft}} down the Krubera Cave.<ref>{{cite journal |last1=Jordana |first1=Rafael |last2=Baquero |first2=Enrique |last3=Reboleira |first3=Sofía |last4=Sendra |first4=Alberto |date=1 January 2012 |title=Reviews of the genera ''Schaefferia'' Absolon, 1900, ''Deuteraphorura'' Absolon, 1901, ''Plutomurus'' Yosii, 1956 and the ''Anurida'' Laboulbène, 1865 species group without eyes, with the description of four new species of cave springtails (Collembola) from Krubera-Voronya cave, Arabika Massif, Abkhazia |journal=Terrestrial Arthropod Reviews |volume=5 |issue=1 |pages=35–85 |doi=10.1163/187498312X622430 |hdl=10171/27607 |hdl-access=free |url=https://scispace.com/pdf/reviews-of-the-genera-schaefferia-absolon-1900-26puktblik.pdf |access-date=10 November 2025 }}</ref>

[[File:Anurida maritima.jpg|thumb|right|''Anurida maritima'' on water]]

The horizontal distribution of springtail species is affected by environmental factors which act at the landscape scale, such as soil acidity, moisture and light.<ref name="Ponge 1993"/> Requirements for pH can be reconstructed experimentally.<ref>{{cite journal |last1=Salmon |first1=Sandrine |last2=Ponge |first2=Jean-François |last3=Van Straalen |first3=Nico |year=2002 |title=Ionic identity of pore water influences pH preference in Collembola |journal=Soil Biology and Biochemistry |volume=34 |issue=11 |pages=1663–7 |doi=10.1016/S0038-0717(02)00150-5 |bibcode=2002SBiBi..34.1663S |url=https://www.academia.edu/20508988 |access-date=10 November 2025 }}</ref> Altitudinal changes in species distribution can be at least partly explained by increased acidity at higher elevation.<ref>{{cite journal |last1=Loranger |first1=Gladys |last2=Bandyopadhyaya |first2=Ipsa |last3=Razaka |first3=Barbara |last4=Ponge |first4=Jean-François |date=March 2001 |title=Does soil acidity explain altitudinal sequences in collembolan communities? |journal=Soil Biology and Biochemistry |volume=33 |issue=3 |pages=381–93 |doi=10.1016/S0038-0717(00)00153-X |bibcode=2001SBiBi..33..381L |s2cid=84523833 |url=https://www.academia.edu/9948943 |access-date=10 November 2025 }}</ref> Moisture requirements, among other ecological and behavioural factors, explain why some species cannot live aboveground,<ref>{{cite journal |last1=Faber |first1=Jack H. |last2=Joosse |first2=Els N.G. |date=November 1993 |title=Vertical distribution of Collembola in a ''Pinus nigra'' organic soil |journal=Pedobiologia |volume=37 |issue=6 |pages=336–50 |doi=10.1016/S0031-4056(24)00108-2 |bibcode=1993Pedob..37..336F |doi-access=free }}</ref> or retreat in the soil during dry seasons,<ref>{{cite journal |last=Detsis |first=Vassilis |year=2000 |title=Vertical distribution of Collembola in deciduous forests under Mediterranean climatic conditions |journal=Belgian Journal of Zoology |volume=130 |issue=Supplement 1 |pages=57–61 |url=https://www.researchgate.net/publication/242773445 |access-date=10 November 2025 }}</ref> but also why some epigeal springtails are always found in the vicinity of ponds and lakes, such as the hygrophilous ''Isotomurus palustris''.<ref>{{cite journal |last1=Lesel |first1=Tanja |last2=Marx |first2=Michael Thomas |last3=Eisenbeis |first3=Gerhard |date=20 May 2011 |title=Effects of ecological flooding on the temporal and spatial dynamics of carabid beetles (Coleoptera, Carabidae) and springtails (Collembola) in a polder habitat |journal=Zookeys |volume=100 |pages=421–46 |doi=10.3897/zookeys.100.1538 |doi-access=free |pmc=3131029 }}</ref> Adaptive features, such as the presence of a fan-like wettable mucro, allow some species to move at the surface of water in freshwater and marine environments.<ref>{{cite book|url=https://books.google.com/books?id=dC2gEAAAQBAJ&dq=snow+hydrophilic+P.+aquatica&pg=PA227 |title=Identification and ecology of freshwater arthropods in the Mediterranean Basin |page=227 |first1=Alain |last1=Maasri |first2=James H. |last2=Thorp |date=31 October 2023 |publisher=Elsevier |location=Amsterdam, The Netherlands |isbn=978-0-12-821856-3 |via=Google Books }}</ref> ''Podura aquatica'', a unique representative of the family Poduridae (and one of the first springtails to have been described by Carl Linnaeus), spends its entire life at the surface of water, its wettable eggs dropping in water until the non-wettable first instar hatches then surfaces.<ref>{{cite journal |last=Pichard |first=Sylvain |year=1973 |title=Contribution à l'étude de la biologie de ''Podura aquatica'' (Linné) Collembole |language=fr |journal=Bulletin Biologique de la France et de la Belgique |volume=107 |issue=4 |pages=291–9 |url=https://archive.org/details/pichard-1973 |access-date=11 November 2025 }}</ref> A few genera are capable of being submerged, and after molting young springtails lose their water repellent properties and are able to survive submerged under water.<ref>{{cite book |url=https://books.google.com/books?id=98mcDwAAQBAJ&dq=P.+aquatica+springtail+eggs+water&pg=PA198 |page=198 |title=Aquatic insects: behavior and ecology |first1=Kleber |last1=Del-Claro |first2=Rhainer |last2=Guillermo |date=10 June 2019 |publisher=Springer |isbn=978-3-030-16327-3 |via=Google Books |access-date=11 November 2025 }}</ref>

In a variegated landscape, made of a patchwork of closed (woodland) and open (meadows, cereal crops) environments, most soil-dwelling species are not specialized and can be found everywhere, but most epigeal and litter-dwelling species are attracted to a particular environment, either forested or not.<ref name="Ponge 1993"/><ref>{{cite journal |last1=Ponge |first1=Jean-François |last2=Gillet |first2=Servane |last3=Dubs |first3=Florence |last4=Fédoroff |first4=Éric|last5=Haese |first5=Lucienne |last6=Sousa |first6=José Paulo |last7=Lavelle |first7=Patrick |date=June 2003 |title=Collembolan communities as bioindicators of land use intensification |journal=Soil Biology and Biochemistry |volume=35 |issue=6 |pages=813–26 |doi=10.1016/S0038-0717(03)00108-1 |bibcode=2003SBiBi..35..813P |hdl=10316/3875 |s2cid=86289373 |hdl-access=free |url=https://www.academia.edu/106027667 |access-date=11 November 2025 }}</ref> As a consequence of dispersal limitation, landuse change, when too rapid, may cause the local disappearance of slow-moving, specialist species,<ref>{{cite journal |last1=Ponge |first1=Jean-François |last2=Dubs |first2=Florence |last3=Gillet |first3=Servane |last4=Sousa |first4=José Paulo |last5=Lavelle |first5=Patrick |date=May 2006 |title=Decreased biodiversity in soil springtail communities: the importance of dispersal and landuse history in heterogeneous landscapes |journal=Soil Biology and Biochemistry |volume=38 |issue=5 |pages=1158–61 |doi=10.1016/j.soilbio.2005.09.004 |bibcode=2006SBiBi..38.1158P |url=https://www.academia.edu/44897122 |access-date=11 November 2025 }}</ref> a phenomenon the measure of which has been called ''colonisation credit''.<ref>{{cite journal |last1=Cristofoli |first1=Sara |last2=Mahy |first2=Grégory |date=May 2010 |title=Colonisation credit in recent wet heathland butterfly communities |journal=Insect Conservation and Diversity |volume=3 |issue=2 |pages=83–91 |doi=10.1111/j.1752-4598.2009.00075.x |s2cid=86558143 |url=https://www.academia.edu/58010063 |access-date=11 November 2025 }}</ref><ref>{{cite journal |last1=Heiniger |first1=Charlène |last2=Barot |first2=Sébastien |last3=Ponge |first3=Jean-François |last4=Salmon |first4=Sandrine |last5=Botton-Divet |first5=Léo |last6=Carmignac |first6=David |last7=Dubs |first7=Florence |date=March 2014 |title=Effect of habitat spatiotemporal structure on collembolan diversity |journal=Pedobiologia |volume=57 |issue=2 |pages=103–17 |doi=10.1016/j.pedobi.2014.01.006 |bibcode=2014Pedob..57..103H |url=https://www.academia.edu/21287032 |access-date=11 November 2025 }}</ref>

===Role in turfgrass ecosystems=== In managed turfgrass and urban lawn ecosystems, Collembola function as "thatch busters", playing a significant role in the decomposition of thatch—the layer of living and dead organic matter between the green vegetation and the soil surface. A common model species in North American turfgrass is ''Isotomiella minor''.{{cn|date=February 2026}}

=== Relationship with humans === [[File:Tomocerus.sp.1.jpg|thumb|right|upright|''Tomocerus'' sp. from Germany]] Springtails are well known as pests of some agricultural crops. ''Sminthurus viridis'', the ''lucerne flea'', has been shown to cause severe damage to agricultural crops,<ref>{{cite journal |last1=Shaw |first1=Michael W. |last2=Haughs |first2=G. M. |date=December 1983 |title=Damage to potato foliage by ''Sminthurus viridis'' (L.) |journal=Plant Pathology |volume=32 |issue=4 |pages=465–6 |doi=10.1111/j.1365-3059.1983.tb02864.x |bibcode=1983PPath..32..465S |url=https://fr.1lib.sk/book/51610413/cc21d0 |access-date=11 November 2025 }}</ref> and is considered as a pest in Australia.<ref>{{cite journal |last1=Bishop |first1=Alan L. |last2=Harris |first2=Anne M. |last3=McKenzie |first3=Harry J. |date=February 2001 |title=Distribution and ecology of the lucerne flea, ''Sminthurus viridis'' (L.) (Collembola: Sminthuridae), in irrigated lucerne in the Hunter dairying region of New South Wales |journal=Australian Journal of Entomology |volume=40 |issue=1 |pages=49–55 |doi=10.1046/j.1440-6055.2001.00202.x |url=https://www.academia.edu/144299100 |access-date=11 November 2025 }}</ref><ref>{{cite web |publisher=State of Victoria |date=29 August 2025 |url=https://agriculture.vic.gov.au/biosecurity/pest-insects-and-mites/priority-pest-insects-and-mites/lucerne-flea |title=Lucerne flea |access-date=11 November 2025 }}</ref> Onychiuridae are also known to feed on tubers and to damage them to some extent.<ref>{{cite journal |last1=Baker |first1=A. N. |last2=Dunning |first2=Andrew R. |date=September 1975 |title=Association of populations of onychiurid Collembola with damage to sugar-beet seedlings |journal=Plant Pathology |volume=24 |issue=3 |pages=150–4 |doi=10.1111/j.1365-3059.1975.tb01882.x |bibcode=1975PPath..24..150B |url=https://fr.1lib.sk/book/41309045/d3aa19 |access-date=11 November 2025 }}</ref> However, by their capacity to carry spores of mycorrhizal fungi and mycorrhiza helper bacteria on their tegument, soil springtails play a positive role in the establishment of plant-fungal symbioses and thus are beneficial to agriculture.<ref>{{cite journal |last1=Klironomos |first1=John N. |last2=Moutoglis |first2=Peter |date=July 1999 |title=Colonization of nonmycorrhizal plants by mycorrhizal neighbours as influenced by the collembolan, ''Folsomia candida'' |journal=Biology and Fertility of Soils |volume=29 |issue=3 |pages=277–81 |doi=10.1007/s003740050553 |bibcode=1999BioFS..29..277K |s2cid=11014525 |url=https://fr.1lib.sk/book/39393157/d7dd18 |access-date=11 November 2025 }}</ref> They also contribute to controlling plant fungal diseases through their active consumption of mycelia and spores of damping-off and pathogenic fungi.<ref>{{cite journal |last1=Sabatini |first1=Maria Agnese |last2=Innocenti |first2=Gloria |date=January 2001 |title=Effects of Collembola on plant-pathogenic fungus interactions in simple experimental systems |journal=Biology and Fertility of Soils |volume=33 |issue=1 |pages=62–6 |doi=10.1007/s003740000290 |bibcode=2001BioFS..33...62S |s2cid=9273050 |hdl=11380/303426 |hdl-access=free |url=https://fr.1lib.sk/book/39392728/3014ff |access-date=11 November 2025 }}</ref><ref>{{cite journal |last1=Shiraishi |first1=Hiroyoshi |last2=Enami |first2=Yoshinari |last3=Okano |first3=Seigo |year=2003 |title=''Folsomia hidakana'' (Collembola) prevents damping-off disease in cabbage and Chinese cabbage by ''Rhizoctonia solani'' |journal=Pedobiologia |volume=47 |issue=1 |pages=33–8 |doi=10.1078/0031-4056-00167 |bibcode=2003Pedob..47...33S |url=https://fr.1lib.sk/book/50527762/21009f |access-date=11 November 2025 }}</ref> It has been suggested that they could be reared to be used for the control of pathogenic fungi in greenhouses and other indoor cultures.<ref>{{cite journal |last1=Ponge |first1=Jean-François |last2=Charpentié |first2=Marie-José |title=Étude des relations microflore-microfaune: expériences sur Pseudosinella alba (Packard), Collembole mycophage |journal=Revue d'Écologie et de Biologie du Sol |year=1981 |volume=18 |issue=3 |pages=291–303 |language=fr |url=https://www.academia.edu/61470655 |access-date=11 November 2025 }}</ref><ref>{{cite journal |last1=Lartey |first1=Robert T. |last2=Curl |first2=Elroy A. |last3=Peterson |first3=Curt M. |last4=Harper |first4=James D. |date=1 April 1989 |title=Mycophagous grazing and food preference of ''Proisotoma minuta'' (Collembola: Isotomidae) and ''Onychiurus encarpatus'' (Collembola: Onychiuridae) |journal=Environmental Entomology |volume=18 |issue=2 |pages=334–7|doi=10.1093/ee/18.2.334 |url=https://fr.1lib.sk/book/83086413/816e7e |access-date=11 November 2025 }}</ref>

Springtails sometimes find their way inside human living spaces. However, most pesticides are not effective against them and moisture-removal strategies are the best line of defense to combat infestations.<ref>{{cite web |title=Springtails |url=https://extension.umn.edu/nuisance-insects/springtails |access-date=11 November 2025 |website=extension.umn.edu |language=en }}</ref>

Various sources and publications have suggested that some springtails may parasitize humans, but this is entirely inconsistent with their biology, and no such phenomenon has ever been scientifically confirmed, though it has been documented that the scales or hairs from springtails can cause irritation when rubbed onto the skin.<ref name="Janssens2007">{{cite web |last1=Janssens |first1=Frans |last2=Christiansen |first2=Kenneth A. |work=Checklist of the Collembola |url=https://www.collembola.org/publicat/sidney.htm |title=Synanthropic Collembola, springtails in association with Man |date=11 February 2018 |access-date=11 November 2025 }}</ref> They may sometimes be abundant indoors in damp places such as bathrooms and basements, and incidentally found on one's person. More often, claims of persistent human skin infection by springtails may indicate a neurological problem, such as delusional parasitosis, a psychological rather than entomological problem. Researchers themselves may be subject to psychological phenomena. For example, a publication in 2004 claiming that springtails had been found in skin samples was later determined to be a case of pareidolia; that is, no springtail specimens were actually recovered, but the researchers had digitally enhanced photos of sample debris to create images resembling small arthropod heads, which then were claimed to be springtail remnants.<ref name="Janssens2007"/><ref>{{cite journal |last=Berenbaum |first=May |date=1 April 2005 |title=Face time |journal=American Entomologist |volume=51 |issue=2 |pages=68–9 |doi=10.1093/ae/51.2.68 |url=https://fr.1lib.sk/book/81520867/78fd20 |access-date=11 November 2025 }}</ref><ref>{{cite journal |last1=Christiansen |first1=Kenneth |last2=Bernard |first2=Ernest C. |date=1 November 2008 |title=Critique of the article "Collembola (Springtails) (Arthropoda: Hexapoda: Entognatha) found in scrapings from individuals diagnosed with delusory parasitosis" |journal=Entomological News |volume=119 |issue=5 |pages=537–40 |doi=10.3157/0013-872x-119.5.537 |s2cid=83870937 |url=https://fr.1lib.sk/book/62847230/d8c61d |access-date=11 November 2025 }}</ref><ref>{{cite journal |last1=Lim |first1=Christopher Seng-Hong |last2=Lim |first2=Scott L. |last3=Chew |first3=Fook Tim |last4=Deharveng |first4=Louis |date=1 January 2009 |title=Collembola are unlikely to cause human dermatitis |journal=Journal of Insect Science |volume=9 |issue=3 |pages=1–5 |doi=10.1673/031.009.0301 |pmc=3011850 |pmid=19611235 |url=https://www.researchgate.net/publication/26678416 |access-date=11 November 2025 }}</ref><ref>{{cite journal |last=Shelomi |first=Matan |title=Evidence of photo manipulation in a delusional parasitosis paper |journal=Journal of Parasitology |date=1 June 2013 |volume=99 |issue=3 |pages=583–5 |doi=10.1645/12-12.1 |pmid=23198757 |s2cid=6473251 |url=https://www.researchgate.net/publication/233825102 |access-date=11 November 2025 }}</ref> However, Steve Hopkin reports one instance of an entomologist aspirating an ''Isotoma'' species and in the process accidentally inhaling some of their eggs, which hatched in his nasal cavity and made him quite ill until they were flushed out.<ref name="Hopkin1997"/>

In 1952, China accused the United States military of spreading bacteria-laden insects and other objects during the Korean War by dropping them from P-51 fighters above rebel villages over North Korea. In all, the U.S. was accused of dropping ants, beetles, crickets, fleas, flies, grasshoppers, lice, springtails, and stoneflies as part of a biological warfare effort. The alleged associated diseases included anthrax, cholera, dysentery, fowl septicemia, paratyphoid, plague, scrub typhus, small pox, and typhoid. China created an international scientific commission for investigating possible bacterial warfare, eventually ruling that the United States probably did engage in limited biological warfare in Korea. The US government denied all the allegations, and instead proposed that the United Nations send a formal inquiry committee to China and Korea, but China and Korea refused to cooperate. U.S. and Canadian entomologists further claimed that the accusations were ridiculous and argued that anomalous appearances of insects could be explained through natural phenomena.<ref>{{cite web |title=The role of insects as biological weapons |url=https://www.montana.edu/historybug/insects-as-bioweapons.html |access-date=11 November 2025 |website=www.montana.edu |last=Peterson |first=Robert K D. }}</ref> Springtail species cited in allegations of biological warfare in the Korean War were ''Isotoma (Desoria) negishina'' (a local species) and the "white rat springtail" ''Folsomia candida''.<ref>{{cite book |last=Lockwood |first=Jeffrey A. |title=Six-legged soldiers: using insects as weapons of war |edition=1st |publisher=Oxford University Press |location=Oxford, United Kingdom |year=2009 |isbn=978-0-19-533305-3 |url=https://fr.1lib.sk/book/728260/66835b |access-date=11 November 2025 }}</ref>

Captive springtails are often kept in a terrarium as part of a clean-up crew.<ref>{{cite web |last=Tansley |first=Luke |url=https://www.reptilecentre.com/blog/2015/12/top-5-bioactive-clean-up-crew/ |title=The top 5 bioactive clean up crew |work=Northampton Reptile Centre |date=6 September 2024 |access-date=11 November 2025 }}</ref>

=== Ecotoxicology laboratory animals ===

Springtails are currently used in laboratory tests for the early detection of soil pollution. Acute and chronic toxicity tests have been performed by researchers, mostly using the parthenogenetic isotomid ''Folsomia candida''.<ref>{{cite journal |last1=Fountain |first1=Michelle T. |last2=Hopkin |first2=Steve P. |date=March 2001 |title=Continuous monitoring of ''Folsomia candida'' (Insecta: Collembola) in a metal exposure test |journal=Ecotoxicology and Environmental Safety |volume=48 |issue=3 |pages=275–86 |doi=10.1006/eesa.2000.2007 |pmid=11222037 |bibcode=2001EcoES..48..275F |url=https://www.academia.edu/84671891 |access-date=11 November 2025 }}</ref> These tests have been standardized.<ref>{{cite web |author=ISO 11267:2023 |year=2023 |title=Soil quality. Inhibition of reproduction of Collembola (''Folsomia candida'') by soil contaminants |publisher=International Organization for Standardization |location=Geneva, Switzerland |url=https://www.iso.org/standard/79816.html }}</ref> Details on a ringtest, on the biology and ecotoxicology of ''Folsomia candida'' and comparison with the sexual nearby species ''Folsomia fimetaria'' (sometimes preferred to ''Folsomia candida'') are given in a document written by Paul Henning Krogh.<ref>{{cite web |last=Krogh |first=Paul Henning |url=https://www2.mst.dk/udgiv/publications/2009/978-87-7052-881-8/pdf/978-87-7052-882-5.pdf |title=Toxicity testing with the collembolans ''Folsomia fimetaria'' and ''Folsomia candida'' and the results of a ringtest |year=2009 |access-date=11 November 2025 }}</ref> Care should be taken that different strains of the same species may be conducive to different results. Avoidance tests have been also performed.<ref>{{cite journal |last1=Lors |first1=Christine |last2=Martínez Aldaya |first2=Maite |last3=Salmon |first3=Sandrine |last4=Ponge |first4=Jean-François |date=August 2006 |title=Use of an avoidance test for the assessment of microbial degradation of PAHs |journal=Soil Biology and Biochemistry |volume=38 |issue=8 |pages=2199–204 |doi=10.1016/j.soilbio.2006.01.026 |bibcode=2006SBiBi..38.2199L |url=https://www.academia.edu/44838246 |access-date=11 November 2025 }}</ref> They have been standardized, too.<ref>{{cite web |author=ISO 17512-2:2011 |year=2011 |title=Soil quality. Avoidance test for determining the quality of soils and effects of chemicals on behaviour. Part 2. Test with collembolans (''Folsomia candida'') |publisher=International Organization for Standardization |location=Geneva, Switzerland |url=https://www.iso.org/standard/50779.html }}</ref> Avoidance tests are complementary to toxicity tests, but they also offer several advantages: they are more rapid (thus cheaper), more sensitive and they are environmentally more reliable, because in the real world Collembola move actively far from pollution spots.<ref>{{cite journal |last1=Chauvat |first1=Matthieu |last2=Ponge |first2=Jean-François |date=September 2002 |title=Colonization of heavy metal-polluted soils by Collembola: preliminary experiments in compartmented boxes |journal=Applied Soil Ecology |volume=21 |issue=2 |pages=91–106 |doi=10.1016/S0929-1393(02)00087-2 |bibcode=2002AppSE..21...91C |url=https://www.academia.edu/46881277 |access-date=11 November 2025 }}</ref> It may be hypothesized that the soil could become locally depauperated in animals (and thus improper to normal use) while below thresholds of toxicity. Contrary to earthworms, and like many insects and molluscs, Collembola are very sensitive to herbicides and thus are threatened in no-tillage agriculture, which makes a more intense use of herbicides than conventional agriculture.<ref>{{cite journal |last1=Ponge |first1=Jean-François |last2=Bandyopadhyaya |first2=Ipsa |last3=Marchetti |first3=Valérie |date=June 2002 |title=Interaction between humus form and herbicide toxicity to Collembola (Hexapoda) |journal=Applied Soil Ecology |volume=20 |issue=3 |pages=239–53 |doi=10.1016/S0929-1393(02)00026-4 |bibcode=2002AppSE..20..239P |url=https://www.academia.edu/9948933 |access-date=11 November 2025 }}</ref> The springtail ''Folsomia candida'' is also becoming a genomic model organism for soil toxicology.<ref>{{cite journal |last1=Nota |first1=Benjamin |last2=Timmermans |first2=Martijn J.T.N. |last3=Franken |first3=Oscar |last4=Montagne-Wajer |first4=Kora |last5=Mariën |first5=Janine |last6=De Boer |first6=Muriel E. |last7=De Boer |first7=Tjalf E. |last8=Ylstra |first8=Bauke |last9=Van Straalen |first9=Nico M. |last10=Roelofs |first10=Dick |date=23 September 2008 |title=Gene expression analysis of Collembola in cadmium containing soil |journal=Environmental Science and Technology |volume=42 |issue=21 |pages=8152–7 |pmid=19031917 |doi=10.1021/es801472r |bibcode=2008EnST...42.8152N |url=https://www.academia.edu/32226969 |access-date=11 November 2025 }}</ref><ref>{{cite journal |last1=Nota |first1=Benjamin |last2=Bosse |first2=Mirte |last3=Ylstra |first3=Bauke |last4=Van Straalen |first4=Nico M. |last5=Roelofs |first5=Dick |year=2009 |title=Transcriptomics reveals extensive inducible biotransformation in the soil-dwelling invertebrate ''Folsomia candida'' exposed to phenanthrene |journal=BMC Genomics |volume=10 |article-number=236 |pmid=19457238 |doi=10.1186/1471-2164-10-236 |pmc=2688526 |doi-access=free }}</ref> With microarray technology the expression of thousands of genes can be measured in parallel. The gene expression profiles of ''Folsomia candida'' exposed to environmental toxicants allow fast and sensitive detection of pollution, and additionally clarifies molecular mechanisms causing toxicity.<ref>{{cite journal |last1=Pitombeira de Figueirêdo |first1=Livia |last2=Damm |first2=Michiel A. |last3=Mainardi |first3=Giulia |last4=Mariën |first4=Janine |last5=Espíndola |first5=Evaldo Luiz Gaeta |last6=Van Gestel |first6=Cornelis A. M. |last7=Roelofs |first7=Dick |date=January 2019 |title=The use of gene expression to unravel the single and mixture toxicity of abamectin and difenoconazole on survival and reproduction of the springtail ''Folsomia candida'' |journal=Environmental Pollution |volume=244 |pages=342–50 |doi=10.1016/j.envpol.2018.10.077 |url=https://fr.1lib.sk/book/104335935/193f2e |access-date=11 November 2025 |hdl=1871.1/eb4b3414-4882-4ba0-bef2-a6f89271ef8a |hdl-access=free }}</ref>

Collembola have been found to be useful as bio-indicators of soil quality. Laboratory studies have been conducted that validated that the jumping ability of springtails can be used to evaluate the soil quality of Cu- and Ni-polluted sites.<ref>{{cite journal |last1=Kim |first1=Shin Woong |last2=An |first2=Youn-Joo |date=March 2014 |title=Jumping behavior of the springtail ''Folsomia candida'' as a novel soil quality indicator in metal-contaminated soils |journal=Ecological Indicators |volume=38 |pages=67–71 |doi=10.1016/j.ecolind.2013.10.033 |bibcode=2014EcInd..38...67K |url=https://fr.1lib.sk/book/56069691/2f47fa |access-date=11 November 2025 }}</ref>

==Climate warming impact==

In polar regions that are expected to experience among the most rapid impact from climate warming, springtails have shown contrasting responses to warming in experimental warming studies.<ref>{{cite journal |last1=Nielsen |first1=Uffe N. |last2=Wall |first2=Diana H. |date=March 2013 |title=The future of soil invertebrate communities in polar regions: different climate change responses in the Arctic and Antarctic? |journal=Ecology Letters |volume=16 |issue=3 |pages=409–19 |pmid=23278945 |doi=10.1111/ele.12058 |bibcode=2013EcolL..16..409N |url=https://fr.1lib.sk/book/54493744/eaae44 |access-date=11 November 2025 }}</ref> There are negative,<ref>{{cite journal |last1=Coulson |first1=Stephen James |last2=Hodkinson |first2=Ian D. |last3=Woolley |first3=Christopher |last4=Webb |first4=Nigel R. |last5=Block |first5=William |last6=Worland |first6=M. Rodger |last7=Bale |first7=Jeff S. |last8=Strathdee |first8=Andrew T. |date=February 1996 |title=Effects of experimental temperature elevation on high-arctic soil microarthropod populations |journal=Polar Biology |volume=16 |issue=2 |pages=147–53 |doi=10.1007/BF02390435 |bibcode=1996PoBio..16..147C |s2cid=13826340 |url=https://www.researchgate.net/publication/227050079 |access-date=11 November 2025 }}</ref><ref name="Sjursen2005">{{cite journal |last1=Sjursen |first1=Heidi |last2=Michelsen |first2=Anders |last3=Jonasson |first3=Sven |date=November 2005 |title=Effects of long-term soil warming and fertilisation on microarthropod abundances in three sub-arctic ecosystems |journal=Applied Soil Ecology |volume=30 |issue=3 |pages=148–61 |doi=10.1016/j.apsoil.2005.02.013 |bibcode=2005AppSE..30..148S |url=https://www.academia.edu/51613927 |access-date=11 November 2025 }}</ref> positive<ref>{{cite journal |last1=Dollery |first1=Rebecca |last2=Hodkinson |first2=Ian D. |last3=Jónsdóttir |first3=Ingibjörg S. |date=February 2006 |title=Impact of warming and timing of snow melt on soil microarthropod assemblages associated with Dryas-dominated plant communities on Svalbard |journal=Ecography |volume=29 |issue=1 |pages=111–9 |doi=10.1111/j.2006.0906-7590.04366.x |bibcode=2006Ecogr..29..111D |doi-access=free }}</ref><ref>{{cite journal |last1=Hågvar |first1=Sigmund |last2=Klanderud |first2=Kari |date=December 2009 |title=Effect of simulated environmental change on alpine soil arthropods |journal=Global Change Biology |volume=15 |issue=12 |pages=2972–80 |doi=10.1111/j.1365-2486.2009.01926.x |bibcode=2009GCBio..15.2972H |citeseerx=10.1.1.658.1606 |s2cid=36287733 |url=https://www.researchgate.net/publication/230184963 |access-date=11 November 2025 }}</ref> and neutral responses reported.<ref name="Sjursen2005"/><ref name="Alatalo2015">{{cite journal |last1=Alatalo |first1=Juha M. |last2=Jägerbrand |first2=Annika K. |last3=Čuchta |first3=Peter |date=16 December 2015 |title=Collembola in three alpine subarctic sites resistant to twenty years of experimental warming |journal=Scientific Reports |volume=5 |article-number=18161 |doi=10.1038/srep18161 |pmc=4680968 |pmid=26670681 |bibcode=2015NatSR...518161A |doi-access=free }}</ref> Neutral responses to experimental warming have also been reported in studies of non-polar regions.<ref>{{cite journal |last1=Kardol |first1=Paul |last2=Reynolds |first2=W. Nicholas |last3=Norby |first3=Richard J. |last4=Classen |first4=Aimée T. |title=Climate change effects on soil microarthropod abundance and community structure |journal=Applied Soil Ecology |date=January 2011 |volume=47 |issue=1 |pages=37–44 |doi=10.1016/j.apsoil.2010.11.001 |bibcode=2011AppSE..47...37K |url=https://www.researchgate.net/publication/225292391 |access-date=11 November 2025 }}</ref> The importance of soil moisture has been demonstrated in experiments using infrared heating in an alpine meadow, which had a negative effect on mesofauna biomass and diversity in drier parts and a positive effect in moist sub-areas.<ref>{{cite journal |last1=Harte |first1=John |last2=Rawa |first2=Agnieszka |last3=Price |first3=Vanessa |date=March 1996 |title=Effects of manipulated soil microclimate on mesofaunal biomass and diversity |journal=Soil Biology and Biochemistry |volume=28 |issue=3 |pages=313–22 |doi=10.1016/0038-0717(95)00139-5 |bibcode=1996SBiBi..28..313H |url=https://www.researchgate.net/publication/222151922 |access-date=11 November 2025 }}</ref> Furthermore, a study with 20 years of experimental warming in three contrasting plant communities found that small scale heterogeneity may buffer springtails to potential climate warming.<ref name="Alatalo2015"/>

== Reproduction ==

Sexual reproduction occurs through the clustered or scattered deposition of spermatophores by male adults. Stimulation of spermatophore deposition by female pheromones has been demonstrated in ''Sinella curviseta''.<ref>{{cite journal |last=Waldorf |first=Elizabeth S. |date=1 December 1974 |title=Sex pheromone in the springtail, ''Sinella curviseta'' |journal=Environmental Entomology |volume=3 |issue=6 |pages=916–8 |doi=10.1093/ee/3.6.916 |url=https://fr.1lib.sk/book/87093677/7c97d6 |access-date=11 November 2025 }}</ref> Mating behavior can be observed in Symphypleona.<ref>{{cite journal |last1=Kozlowski |first1=Marek Wojciech |last2=Aoxiang |first2=Shi |date=22 September 2005 |title=Ritual behaviors associated with spermatophore transfer in ''Deuterosminthurus bicinctus'' (Collembola: Bourletiellidae) |journal=Journal of Ethology |volume=24 |issue=2 |pages=103–9 |doi=10.1007/s10164-005-0162-6 |s2cid=12056710 |url=https://fr.1lib.sk/book/39655957/cbc3ea |access-date=11 November 2025 }}</ref> Among Symphypleona, males of some Sminthuridae use a clasping organ located on their antenna.<ref name="Hopkin1997"/> Many springtails, mostly those living in deeper soil horizons, are parthenogenetic, which favors reproduction to the detriment of genetic diversity and thereby to population tolerance of environmental hazards.<ref>{{cite journal |last1=Simon |first1=Jean-Christophe |last2=Delmote |first2=François |last3=Rispe |first3=Claude |last4=Crease |first4=Teresa |date=May 2003 |title=Phylogenetic relationships between parthenogens and their sexual relatives: the possible routes to parthenogenesis in animals |journal=Biological Journal of the Linnean Society |volume=79 |issue=1 |pages=151–3 |doi=10.1046/j.1095-8312.2003.00175.x |url=https://www.researchgate.net/publication/228707432 |access-date=11 November 2025 }}</ref><ref>{{cite journal |last=Ponge |first1=Jean-François |date=March 2020 |title=Move or change, an eco-evolutionary dilemma: the case of Collembola |journal=Pedobiologia |volume=79 |article-number=150625 |doi=10.1016/j.pedobi.2020.150625 |url=https://www.academia.edu/42005378 |access-date=11 November 2025 }}</ref> Parthenogenesis (also called thelytoky) is under the control of symbiotic bacteria of the genus ''Wolbachia'', which live, reproduce and are carried in female reproductive organs and eggs of Collembola.<ref>{{cite journal |last1=Czarnetzki |first1=Alice B. |last2=Tebbe |first2=Christoph C. |date=January 2004 |title=Detection and phylogenetic analysis of ''Wolbachia'' in Collembola |journal=Environmental Microbiology |volume=6 |issue=1 |pages=35–44 |doi=10.1046/j.1462-2920.2003.00537.x |pmid=14686939 |bibcode=2004EnvMi...6...35C |url=https://www.academia.edu/12808291 |access-date=11 November 2025 }}</ref> Feminizing ''Wolbachia'' species are widespread in arthropods<ref>{{cite journal |last1=Werren |first1=John H. |last2=Zhang |first2=Wan |last3=Guo |first3=Li Rong |date=22 July 1995 |title=Evolution and phylogeny of ''Wolbachia'': reproductive parasites of arthropods |journal=Proceedings of the Royal Society B |volume=261 |issue=1360 |pages=55–63 |pmid=7644549 |jstor=50047 |doi=10.1098/rspb.1995.0117 |bibcode=1995RSPSB.261...55W |s2cid=8585323 |url=https://fr.1lib.sk/book/53171160/6615d9 |access-date=11 November 2025 }}</ref> and nematodes,<ref>{{cite journal |last1=Fenn |first1=Katelyn |last2=Blaxter |first2=Mark |date=April 2004 |title=Are filarial nematode ''Wolbachia'' obligate mutualist symbionts? |journal=Trends in Ecology and Evolution |volume=19 |issue=4 |pages=163–6 |doi=10.1016/j.tree.2004.01.002 |pmid=16701248 |url=https://fr.1lib.sk/book/49511648/896697 |access-date=11 November 2025 }}</ref> where they co-evolved with most of their lineages.

== See also == {{Portal|Arthropods}} * Hexapoda

== References == {{Reflist|30em}}

== External links == <!-- {{Wikispecies|Collembola}} [has an odd "semi-modern" classification from ITIS which should not be used here. Useful for order and family articles though] --> {{Commons category|Collembola}} * {{cite EB1911|wstitle=Springtail |volume=25 |short=x}} * [https://www.collembola.org/ Checklist of the Collembola of the World] * [http://urweb.roehampton.ac.uk/collembola/taxonomy/ Summary information about the distribution and ecology of Collembola (springtails) in the UK and Ireland] {{Webarchive|url=https://web.archive.org/web/20190422055046/http://urweb.roehampton.ac.uk/collembola/taxonomy/ |date=22 April 2019 }} * [https://earthlife.net/collembola-cave-dwelling-springtails/ General information on Collembola] * [https://www.nhm.ac.uk/resources-rx/files/35feat_springtails_most_abundent-3056.pdf A small lecture from Steve Hopkin] * [https://www.janvanduinen.nl/collembolaengels.php General information on Collembola, with many macrophotographs from Dutch springtails] * [https://collembola.co.za/ Collembola species currently recorded from South Africa]

{{Arthropods}} {{Taxonbar|from=Q190701}} {{Authority control}}

Category:Springtails <!-- Category:Subclasses moved to "Collembola" redirect --> Category:Articles containing video clips Category:Extant Early Devonian first appearances Category:Taxa named by John Lubbock, 1st Baron Avebury