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 all are to 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, rather, can indirectly through the fragmentation of organic matter and the control of soil microbial communities. The word "collembola" is from the Greek colle meaning glue and embolon meaning piston or peg.
Members of Collembola are normally less than 6 mm (0.24 in) long, have six or fewer abdominalsegments and possess a tubular appendage (the 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 have a globular body. Collembola lack a tracheal respiration system, which forces them to respire through a porouscuticle, to the notable exception of Sminthuridae which exhibit a rudimentary, although fully functional, tracheal system.
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.
Springtails also possess the ability to reduce their body size by as much as 30% through subsequent ecdyses (molting) 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.
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 28SrRNAsequence data suggests that they form the most ancient lineage of springtails, which would explain their peculiar apomorphies.
Fossil collembola are rare. Instead, most 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.
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, 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 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.
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. 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.
As a group, springtails are highly sensitive to desiccation, because of their tegumentary respiration. although some species with thin, permeable cuticles have been shown to resist severe drought by regulating the osmotic pressure of their body fluid. The gregarious behaviour of Collembola, mostly driven by the attractive power of pheromones excreted by adults, 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. Sensitivity to drought varies from species to species and increases during ecdysis. 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, an advantage reinforced by synchronized moulting. The high humidity environment of many caves also favours springtails and there are numerous cave adapted species, including one, Plutomurus ortobalaganensis living 1,980 metres (6,500 ft) down the Krubera Cave.
The horizontal distribution of springtail species is affected by environmental factors which act at the landscape scale, such as soil acidity, moisture and light. Requirements for pH can be reconstructed experimentally. Altitudinal changes in species distribution can be at least partly explained by increased acidity at higher elevation. Moisture requirements, among other ecological and behavioural factors, explain why some species cannot live aboveground, or retreat in the soil during dry seasons, 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 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.
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. As a consequence of dispersal limitation, landuse change, when too rapid, may cause the local disappearance of slow-moving, specialist species, a phenomenon the measure of which was recently called colonisation credit.
Relationship with humans
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, and is considered as a pest in Australia. Also Onychiuridae are known to feed on tubers and to damage them to some extent. 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. They also contribute to controlling plant fungal diseases through their active consumption of mycelia and spores of damping-off and pathogenic fungi. It has been suggested that they could be reared to be used for the control of pathogenic fungi in greenhouses and other indoor cultures.
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 collembolans can cause irritation when rubbed onto the skin. 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 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. 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.
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 parthenogeneticisotomidFolsomia candida. These tests have been standardized. More recently, avoidance tests have been also performed. They have been standardized, too. 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. 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. The springtail Folsomia candida is also becoming a genomic model organism for soil toxicology. 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.
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^Julia Nickerl, Ralf Helbig, Hans-Jürgen Schulz, Carsten Werner, & Christoph Neinhuis (2013). "Diversity and potential correlations to the function of Collembola cuticle structures". Zoomorphology132 (2): 183–195. doi:10.1007/s00435-012-0181-0.
^Martin Holmstrup & Mark Bayley (2013). "Protaphorura tricampata, a euedaphic and highly permeable springtail that can sustain activity by osmoregulation during extreme drought". Journal of Insect Physiology59 (11): 1104–1110. doi:10.1016/j.jinsphys.2013.08.015.
^Herman A. Verhoef (1981). "Water balance in Collembola and its relation to habitat selection: water content, haemolymph osmotic pressure and transpiration during an instar". Journal of Insect Physiology27 (11): 755–760. doi:10.1016/0022-1910(81)90065-2.
^May Berenbaum (2005). "Face Time" (PDF). The American Entomologist51 (2): 68–69.
^Kenneth Christiansen & Ernest C. Bernard (2008). "Critique of the article "Collembola (Springtails) (Arthropoda: Hexapoda: Entognatha) found in scrapings from individuals diagnosed with delusory parasitosis"". Entomological News119 (5): 537–540. doi:10.3157/0013-872x-119.5.537.
^Michelle T. Fountain & Steve P. Hopkin (2001). "Continuous monitoring of Folsomia candida (Insecta: Collembola) in a metal exposure test". Ecotoxicology and Environmental Safety48 (3): 275–286. doi:10.1006/eesa.2000.2007.
^Benjamin Nota, Martijn J.T.N. Timmermans, Oscar Franken, Kora Montagne-Wajer, Janine Mariën, Muriel E. De Boer, Tjalf E. De Boer, Bauke Ylstra, Nico M. Van Straalen & Dick Roelofs (2008). "Gene expression analysis of Collembola in cadmium containing soil". Environmental Science and Technology42 (21): 8152–8157. doi:10.1021/es801472r. PMID19031917.
^Marek Wojciech Kozlowski & Shi Aoxiang (2006). "Ritual behaviors associated with spermatophore transfer in Deuterosminthurus bicinctus (Collembola : Bourletiellidae)". Journal of Ethology24 (2): 103–110. doi:10.1007/s10164-005-0162-6.
^Alice B. Czarnetzki & Christoph C. Tebbe (2004). "Detection and phylogenetic analysis of Wolbachia in Collembola". Environmental Microbiology6 (1): 35–44. doi:10.1046/j.1462-2920.2003.00537.x.