Springtail

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Springtails
Temporal range: Early Devonian – present
Orchesella cincta
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Clade: Pancrustacea
Subphylum: Hexapoda
Subclass: Collembola
Lubbock, 1871
Orders
Synonyms[1]
  • Oligentoma
  • Oligoentoma

Springtails (Collembola) form the largest of the three lineages of modern hexapods that are no longer considered insects (the other two are the Protura and Diplura). Although the three orders 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 all insects, which have external mouthparts.

Collembolans are

omnivorous, free-living organisms that prefer moist conditions. They do not directly engage in the decomposition of organic matter, but contribute to it indirectly through the fragmentation of organic matter[2] and the control of soil microbial communities.[3] The word Collembola is from the ancient Greek κόλλα kólla "glue" and ἔμβολος émbolos "peg"; this name was given due to the existence of the collophore, which was previously thought to stick to surfaces to stabilize the creature.[4]

Some

basal lineage of Hexapoda, they are elevated to full class
status.

Morphology

Isotoma anglicana (Entomobryomorpha) with visible furcula
Deutonura monticola (Poduromorpha)

Members of the Collembola are normally less than 6 mm (0.24 in) long, have six or fewer

furcula (or furca). It is located on the fourth abdominal segment of collembolans 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 milliseconds.[12][11]

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.[13]

The

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 length, have dispersed pigmentation, shortened limbs, and a reduced number of ocelli. Euedaphic species inhabit upper mineral layers known as the humus horizon. They are smaller than hemiedaphic species; have soft, elongated bodies; lack pigmentation and ocelli; and have reduced or absent furca.[15][16][17]

Poduromorphs are characterized by their elongated bodies and conspicuous segmentation – three thoracic segments, six abdominal segments, including a well-developed prothorax with tergal chaetae,[17] while the first thoracic segment in Entomobryomorpha is clearly reduced and bears no chaetae.

The digestive tract of collembolan species consists of three main components: the 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.[11] Malpighian tubules are absent.[18]

Systematics and evolution

Allacma fusca (Symphypleona) on rotting wood

Traditionally, the springtails were divided into the

junior synonym of the Collembola.[22]

The term "Neopleona" is essentially synonymous with Symphypleona + Neelipleona.

mtDNA[20] and whole-genome data.[21]

The latest whole-genome phylogeny supporting four orders of Collembola:[21]

Springtails are attested to since the

Rhyniella praecursor, is the oldest terrestrial arthropod, and was found in the famous Rhynie chert of Scotland. Given its morphology resembles extant species quite closely, the radiation of the Hexapoda can be situated in the Silurian, 420 million years ago or more.[25] Additional research concerning the coprolites (fossilized feces) of ancient collembolans allowed researchers to track their lineages back some 412 million years.[11]

Fossil Collembola are rare. Instead, most are found in amber.[26] 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,[27] Miocene of Central America,[28] and the mid-Cretaceous of Burma and Canada.[29] 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 about 3,600 different species.[30]

Ecology

Eating behavior

Specific feeding strategies and mechanisms are employed to match specific niches.[31] Herbivorous and detritivorous species fragment biological material present in soil and leaf litter, supporting decomposition and increasing the availability of nutrients for various species of microbes and fungi.[32] Carnivorous species maintain populations of small invertebrates such as nematodes, rotifers, and other collembolan species.[11][15] 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.[33]

Predators

Springtails are consumed by mesostigmatan mites in various families, including Ascidae, Laelapidae, Parasitidae, Rhodacaridae and Veigaiidae.[34]

Cave-dwelling springtails are a food source for spiders and harvestmen in the same environment, such as the endangered harvestman Texella reyesi.[35]

To protect themselves, some species have evolved chemical defenses.[36]

Distribution

Springtails are

microbivores, and one of the main biological agents responsible for the control and the dissemination of soil microorganisms.[38] 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.[39]

"Snow flea"
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,

grass tufts, ant and termite nests) occur.[41] 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 one springtail, the so-called snow flea (Hypogastrura nivicola), is readily observed on warm winter days when it is active and its dark color contrasts sharply with a background of snow.[42]

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.[43][44] 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.[41] In tropical regions a single square meter of canopy habitat can support many species of Collembola.[12]

The main

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".[48]

Dicyrtomina sp. on leaf

As a group, springtails are highly sensitive to

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.[55] The high humidity environment of many caves also favours springtails and there are numerous cave adapted species,[56][57] including one, Plutomurus ortobalaganensis living 1,980 metres (6,500 ft) down the Krubera Cave.[58]

Anurida maritima on water

The horizontal distribution of springtail species is affected by environmental factors which act at the landscape scale, such as soil

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.[65] A few genera are capable of being submerged, and after molting young springtails lose their water repellent properties and is able to survive submerged under water.[66]

In a variegated landscape, made of a patchwork of closed (

specialist species,[68] a phenomenon the measure of which has been called colonisation credit.[69][70]

Relationship with humans

Tomocerus sp. from Germany

Springtails are well known as

pathogenic fungi in greenhouses and other indoor cultures.[78][79]

Various sources and publications have suggested that some springtails may

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.[39]

In 1952,

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.[85] 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.[86]

Captive springtails are often kept in a terrarium as part of a clean-up crew.[87]

Ecotoxicology laboratory animals

Springtails are currently used in laboratory tests for the early detection of

isotomid Folsomia candida.[88] These tests have been standardized.[89] 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.[90] Care should be taken that different strains of the same species may be conducive to different results. Avoidance tests have been also performed.[91] They have been standardized, too.[92] 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.[93] 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.[94]
The springtail
genomic model organism for soil toxicology.[95][96] 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 toxicology.

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.[97]

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.[98] There are negative,[99][100] positive[101][102] and neutral responses reported.[100][103] Neutral responses to experimental warming have also been reported in studies of non-polar regions.[104] 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.[105] 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.[103]

Reproduction

environmental hazards.[108] 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.[109] Feminizing Wolbachia species are widespread in arthropods[110] and nematodes,[111] where they co-evolved with most of their lineages
.

See also

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External links

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