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Springtail
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Springtails (Collembola) form the largest of the three lineages of

modern hexapods that are no longer considered insects (the other two are Springtails
the Protura and Diplura). Although the three orders are sometimes Temporal range:
grouped together in a class called Entognatha because they have internal
Early Devonian–Recent
mouthparts, they do not appear to be any more closely related to one
another than they all are to insects, which have external mouthparts. PreЄ Є O S D C P T J K Pg N

Some DNA sequence studies[2][3][4] suggest that Collembola represent a

separate evolutionary line from the other Hexapoda, but others
disagree;[5] this seems to be caused by widely divergent patterns of
molecular evolution among the arthropods.[6] The adjustments of
traditional taxonomic rank for springtails reflects 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 a
basal lineage of Hexapoda, they are elevated to full class status.

Orchesella cincta
Contents Scientific classification

1 Description Kingdom: Animalia

2 Systematics and evolution Phylum: Arthropoda
3 Ecology Subphylum: Hexapoda
3.1 Distribution
3.2 Relationship with humans Class: Entognatha (but see text)
3.3 Ecotoxicology laboratory animals Subclass: Collembola
4 Reproduction
Lubbock, 1870
5 See also
6 References Orders
7 External links
Entomobryomorpha
Poduromorpha
Description Symphypleona
Neelipleona (disputed)
Members of Collembola are normally less than 6 mm (0.24 in) long, have
six or fewer abdominal segments and possess a tubular appendage (the Synonyms [1]
collophore or ventral tube) with eversible vesicles, projecting ventrally
from the first abdominal segment. The Poduromorpha and
Entomobryomorpha have an elongated body, while the Symphypleona Oligentoma
have a globular body. Collembola lack a tracheal respiration system, Oligoentoma
which forces them to respire through a porous cuticle, to the notable
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exception of Sminthuridae which exhibit a rudimentary, although fully

functional, tracheal system.[7]

Most species have an abdominal, tail-like appendage, the furcula, that is folded beneath the body to be used for
jumping when the animal is threatened. It is held under tension by a small structure called the retinaculum and when
released, snaps against the substrate, flinging the springtail into the air. All of this takes place in as little as 18
milliseconds.[8]

Systematics and evolution

Traditionally, the springtails were divided into the orders Arthropleona,
Symphypleona and occasionally also Neelipleona. The Arthropleona
were divided into two superfamilies, the Entomobryoidea and the
Poduroidea. But actually, these two and the Symphypleona form three
lineages, each of which is equally distant from the other two. Thus, the
Isotoma with visible furcula Arthropleona are abolished in modern classifications, and their
superfamilies are raised in rank accordingly, being now the
Entomobryomorpha and the Poduromorpha. Technically, the Arthropleona are thus a partial junior synonym of the
Collembola. The term "Neopleona" is essentially synonymous with Symphypleona + Neelipleona.

The Neelipleona were originally seen as a particular advanced lineage of Symphypleona, based on the shared
global body shape. But the global body of Neelipleona is realised in a completely different way than in
Symphypleona. Subsequently, the Neelipleona were considered as being derived from the Entomobryomorpha. But
analysis of 18S and 28S rRNA sequence data suggests that they form the most ancient lineage of springtails, which
would explain their peculiar apomorphies.[5]

Springtails are attested to since the Early Devonian.[9] The fossil from 400 million years ago, Rhyniella praecursor,
is the oldest terrestrial arthropod, and was found in the famous Rhynie chert of Scotland. Given its morphology
resembles extant species quite well, the radiation of the Hexapoda can be situated in the Silurian,
420 million years ago or more.[10]

Fossil collembola are rare. Most fossils are found in amber. 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, Miocene of Central
America, and the mid-Cretaceous of Burma and Canada. They display some unusual 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.

Ecology
Distribution

Springtails are cryptozoa frequently found in leaf litter and other decaying material,[11] where they are primarily
detritivores and microbivores, and one of the main biological agents responsible for the control and the
dissemination of soil microorganisms.[12]

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In sheer numbers, they are reputed to be

one of the most abundant of all
macroscopic animals, with estimates of
100,000 individuals per cubic meter of
topsoil, essentially everywhere on Earth
where soil and related habitats (moss
cushions, fallen wood, grass tufts, ant and
termite nests) occur; 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
"Snow flea" cannot be used for absolute comparisons,
it is notable that nematodes are a phylum A species of Sminthurinae
and crustaceans a subphylum. Most (Symphypleona:
springtails are small and difficult to see by casual observation, but one springtail, Sminthuridoidea:
the so-called snow flea (Hypogastrura nivicola), is readily observed on warm Sminthuridae)
winter days when it is active and its dark color contrasts sharply with a
background of snow.[13]

In addition, a few species routinely climb trees and form a dominant component of canopy faunas, where they may
be collected by beating or insecticide fogging.[14][15] These tend to be the larger (>2 mm) species, mainly in the
genera Entomobrya and Orchesella, though the densities on a per square meter basis are typically 1–2 orders of
magnitude lower than soil populations of the same species. In temperate regions, a few species (e.g. Anurophorus
spp., Entomobrya albocincta, Xenylla xavieri, Hypogastrura arborea) are almost exclusively arboreal. In
tropical regions a single square meter of canopy habitat can support many species of Collembola.[8]

The main ecological factor driving the local distribution of species is the vertical stratification of the environment: 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.[16] This is a complex factor embracing both nutritional and
physiological requirements, together with probable species interactions. Some species have been shown to exhibit
negative[14] or positive[17] gravitropism, which adds a behavioural dimension to this still poorly understood vertical
segregation.

As a group, springtails are highly sensitive to desiccation, because of their

Dicyrtomina sp. on leaf
springtails and there are numerous cave adapted species.[20][21]
tegumentary respiration. The gregarious behaviour of Collembola, mostly
driven by the attractive power of pheromones excreted by adults,[18]
gives more chance to every juvenile or adult individual to find suitable,
better protected places, where desiccation could be avoided and
reproduction rate could be kept at an optimum. Sensitivity to drought
varies from species to species and increases during ecdysis.[19] Given
that springtails are moulting 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. The high humidity environment of many caves also favours

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The horizontal distribution of springtail species is affected by

environmental factors which act at the landscape scale, such as soil
acidity, moisture and light.[16] Requirements for pH can be reconstructed
experimentally.[22] Altitudinal changes in species distribution can be at
least partly explained by increased acidity at higher elevation.[23]
Moisture requirements explain why some species cannot live
aboveground (vertical stratification), but also why some epigeal
springtails are always found in the vicinity of ponds and lakes, such as the
hygrophilous Isotomurus palustris. Adaptive features, such as the
presence of a fan-like wettable mucro, allow some species to move at
the surface of water (Sminthurides aquaticus, Sminthurides Anurida maritima on water
malmgreni). Podura aquatica, a unique representative of the family
Poduridae (and one of the first springtails to have been described by
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.[24]

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.[16][25] As a consequence of
dispersal limitation, landuse change, when too rapid, may cause the local disappearance of slow-moving, specialist
species.[26]

Relationship with humans

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,[27] and is considered as a pest in Australia.[28][29] Also Onychiuridae are
known to feed on tubers and to damage them to some extent.[30] 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.[31] They also contribute to
controlling plant fungal diseases through their active consumption of mycelia and
spores of damping-off and pathogenic fungi.[32][33] It has been suggested that
they could be reared to be used for the control of pathogenic fungi in greenhouses
and other indoor cultures.[34][35]

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 Tomocerus sp. from
documented that the scales or hairs from collembolans can cause irritation when Germany
rubbed onto the skin.[36] 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
Morgellons Syndrome, or delusory 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
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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.[36][37] However, 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.[11]

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.[38] These
tests have been standardized.[39] More recently, avoidance tests have been also performed.[40] They have been
standardized, too.[41] 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 may move far from pollution sources.[42] 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.[43]
The springtail Folsomia candida is also becoming a genomic model organism for soil toxicology.[44][45] With
microarray technology the expression of thousands of genes can be measured in parallel. The gene expression
profiles of F. candida exposed to environmental toxicants allow fast and sensitive detection of pollution, and
additionally clarifies molecular mechanisms causing toxicology.

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.[46]
Mating behaviour can be observed in Symphypleona.[47] Among Symphypleona, males of some Sminthuridae use a
clasping organ located on their antenna. Many collembolan species, mostly those living in deeper soil horizons, are
parthenogenetic, which favours reproduction to the detriment of genetic diversity and thereby to population
tolerance of environmental hazards. 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.[48] Feminizing Wolbachia species are widespread in arthropods[49] and nematodes,[50] where they
co-evolved with most of their lineages.

See also
Texella reddelli, a predator of Collembola

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PAHs) (PDF). Soil Biology and Biochemistry 38 (8): 2199–2204. doi:10.1016/j.soilbio.2006.01.026
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External links
Checklist of the Collembola of the World (http://www.collembola.org/)
Maps of Collembola (Britain and Ireland) (http://ws1.roehampton.ac.uk/collembola/taxonomy/index.html)
Maps from 2006 of UK Collembola, plus Photolibrary (http://www.stevehopkin.co.uk/collembolamaps/)
General information on Collembola (http://www.earthlife.net/insects/collembo.html)
The Biology of the Collembola (http://www.fathom.com/feature/122603/)
Tree of Life (http://tolweb.org/tree?group=Collembola&contgroup=Hexapoda)
North American Collembola (http://www.nearctica.com/nomina/oddbugs/collem.htm)
Springtails, Kansas State University (http://www.ksre.ksu.edu/library/entml2/EP124.pdf)

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Categories: Collembola Arthropods

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