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

Springtails (class Collembola) form the largest of the three lineages of modern hexapods that are no longer considered insects, i.e. Protura, Diplura and Collembola. 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.[2] There are more than 9000 species.[3]

Springtails are omnivorous, free-living organisms that prefer moist conditions.[4] They do not directly engage in the decomposition of organic matter, but contribute to it indirectly through the fragmentation of organic matter[5] and the control of soil microbial communities.[6][7] The word Collembola is from the Ancient Greek κόλλα kólla meaning 'glue' and ἔμβολος 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.[8]

Early DNA sequence studies[9][10][11] suggested that Collembola represent a separate evolutionary line from the other Hexapoda, but others disagree;[12] this seems to be caused by widely divergent patterns of molecular evolution among the arthropods.[13] 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 a basal lineage of Hexapoda, they are elevated to full class status.

Morphology

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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 abdominal segments, and possess a tubular appendage (the collophore or ventral tube) with reversible, sticky vesicles, projecting ventrally from the first abdominal segment.[14] It is believed to be associated with fluid uptake and balance, excretion, and orientation of the organism itself.[4] 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 milliseconds.[15][4]

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.[16]

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.[17][14] The anatomical variance (life-form variation) present between different species partially depends on their vertical distribution across the various strata of terrestrial ecosystems.[18] 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.[19][20][21]

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.[21]

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.[4] Malpighian tubules are absent.[22]

Genomics

[edit]

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.[23] Even though some earlier genome sequences were produced, they do not meet modern standards (contig N50 > 1 MB). One of the first genomes was presented in 2025, that of Orchesella flavescens, which is 270 MB and was assembled into 6 chromosome-level scaffolds.[24]

Systematics and evolution

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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.[25][26][27] 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.[28]

The term Neopleona is essentially synonymous with Symphypleona + Neelipleona.[29] 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.[12] This phylogenetic relationship was also confirmed using a phylogeny based on mtDNA[26] and whole-genome data.[27]

The latest whole-genome phylogeny supporting four orders of Collembola:[27]

Springtails are attested to since the Early Devonian. The fossil from 400 million years ago, Rhyniella praecursor, is the oldest terrestrial arthropod, and was found in the famous Rhynie chert of Scotland.[30] 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.[31] Additional research concerning the coprolites (fossilized feces) of ancient springtails allowed researchers to track their lineages back some 412 million years.[4]

Fossil Collembola are rare. Instead, most are found in amber.[32] 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,[33] Miocene of Central America,[34] and the mid-Cretaceous of Burma and Canada.[35] 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.[4] However, to the light of ongoing developments in molecular methods and the rise of cryptic species unveiled by DNA barcoding within morphologically-described species,[36] 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.[37]

Ecology

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Eating behavior

[edit]

Specific feeding (foraging) strategies and mechanisms are employed to match specific niches.[38] 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.[39] Carnivorous species maintain populations of small invertebrates such as nematodes, rotifers, and other collembolan species.[4][19] 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.[40] 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.[41] 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.[42] Directional movement towards the food source has been recorded by image analysis in laboratory experiments.[43] 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.[44]

Predators

[edit]

Springtails are consumed by mesostigmatan mites in various families, including Ascidae, Laelapidae, Parasitidae, Rhodacaridae and Veigaiidae.[45]

Cave-dwelling springtails are a food source for spiders and harvestmen in the same environment, such as the endangered harvestman Texella reyesi.[46]

Predators also include various soil centipedes.[47]

Jumping, using the furcula as a spring, is the most common way to avoid predation,[48] 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.[49]

Distribution

[edit]

Springtails are cryptozoa frequently found in leaf litter and other decaying material,[4] where they are primarily detritivores and microbivores, and one of the main biological agents responsible for the control and the dissemination of soil microorganisms.[50] 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.[4]

"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,[51] essentially everywhere on Earth where soil and related habitats (moss cushions, fallen wood, grass tufts, ant and termite nests) occur.[52] 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,[53] are readily observed on warm winter days when it is active and its dark color contrasts sharply with a background of snow.[54]

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.[55][56] In temperate regions, a few species (e.g. Anurophorus laricis, Entomobrya albocincta, Xenylla xavieri, Hypogastrura arborea) are almost exclusively arboreal.[52] In tropical regions a single square meter of canopy habitat can support many species of Collembola.[15]

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.[52] This is a complex factor embracing both nutritional and physiological requirements, together with behavioural trends,[57] dispersal limitation[58] and probable species interactions.[59] Some species have been shown to exhibit negative[60] or positive[57] 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.[61]

Dicyrtomina sp. on leaf

As a group, springtails are highly sensitive to desiccation, because of their tegumentary respiration,[62] although some species with thin, permeable cuticles have been shown to resist severe drought by regulating the osmotic pressure of their body fluid.[63] The gregarious behaviour of Collembola, mostly driven by the attractive power of pheromones excreted by adults,[64] 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.[65] Sensitivity to drought varies from species to species[66] and increases during ecdysis.[67] 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.[68] The high humidity of many caves also favours springtails and there are numerous cave adapted species,[69][70] including one, Plutomurus ortobalaganensis living 1,980 metres (6,500 ft) down the Krubera Cave.[71]

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.[52] Requirements for pH can be reconstructed experimentally.[72] Altitudinal changes in species distribution can be at least partly explained by increased acidity at higher elevation.[73] Moisture requirements, among other ecological and behavioural factors, explain why some species cannot live aboveground,[74] or retreat in the soil during dry seasons,[75] but also why some epigeal springtails are always found in the vicinity of ponds and lakes, such as the hygrophilous Isotomurus palustris.[76] 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.[77] 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.[78] 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.[79]

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.[52][80] As a consequence of dispersal limitation, landuse change, when too rapid, may cause the local disappearance of slow-moving, specialist species,[81] a phenomenon the measure of which has been called colonisation credit.[82][83]

Relationship with humans

[edit]
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,[84] and is considered as a pest in Australia.[85][86] Onychiuridae are also known to feed on tubers and to damage them to some extent.[87] 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.[88] They also contribute to controlling plant fungal diseases through their active consumption of mycelia and spores of damping-off and pathogenic fungi.[89][90] It has been suggested that they could be reared to be used for the control of pathogenic fungi in greenhouses and other indoor cultures.[91][92]

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.[93]

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.[94] 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.[94][95][96][97][98] 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.[4]

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.[99] 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.[100]

Captive springtails are often kept in a terrarium as part of a clean-up crew.[101]

Ecotoxicology laboratory animals

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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.[102] These tests have been standardized.[103] 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.[104] Care should be taken that different strains of the same species may be conducive to different results. Avoidance tests have been also performed.[105] They have been standardized, too.[106] 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.[107] 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.[108] The springtail Folsomia candida is also becoming a genomic model organism for soil toxicology.[109][110] 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.[111]

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.[112]

Climate warming impact

[edit]

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.[113] There are negative,[114][115] positive[116][117] and neutral responses reported.[115][118] Neutral responses to experimental warming have also been reported in studies of non-polar regions.[119] 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.[120] 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.[118]

Reproduction

[edit]

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.[121] Mating behavior can be observed in Symphypleona.[122] Among Symphypleona, males of some Sminthuridae use a clasping organ located on their antenna.[4] 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.[123][124] 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.[125] Feminizing Wolbachia species are widespread in arthropods[126] and nematodes,[127] where they co-evolved with most of their lineages.

See also

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References

[edit]
[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Springtail

View on Grokipedia
from Grokipedia
Springtails, of the order Collembola within the subclass , are minute, wingless hexapods distinguished from by internalized mouthparts and the absence of a true . Typically measuring less than 6 mm in length, they possess a forked abdominal appendage termed the , which folds under the body and snaps against the substrate to propel jumps of up to 10 cm or more, enabling escape from predators despite their soft-bodied form. This jumping mechanism, actuated by sudden muscular release from a clasp-like retinaculum, exemplifies their adaptive morphology for survival in microhabitats. Abundant across global terrestrial ecosystems, particularly in damp , leaf litter, and decaying , springtails number in the thousands of and dominate arthropod communities, often exceeding densities of millions per square meter. They function as key microbivores and detritivores, accelerating litter decomposition and facilitating nutrient cycling essential to and growth. While generally beneficial, certain may proliferate in high-moisture human environments, prompting occasional pest management concerns, though they pose no direct harm to structures or crops.

Taxonomy and Phylogeny

Classification and Systematics

Collembola, commonly known as springtails, constitute a distinct class within the of the Arthropoda, separate from the class Insecta due to features such as entognathous (internalized) mouthparts, the absence of wings, and unique appendages like the collophore and . The class was formally established by Lubbock in 1871. Within , Collembola occupy a basal phylogenetic position, often regarded as the to Insecta plus , with also included in the broader hexapod lineage. Modern systematics recognizes four orders: Neelipleona, , Poduromorpha, and Entomobryomorpha, encompassing approximately 30 families and over 9,600 described species worldwide, though estimates suggest a total of 50,000 to 65,000 species exist. Neelipleona, the smallest order with one family (Neelidae), is considered plesiomorphic and basal in phylogenies. , characterized by a globular body form (symphypleonous morphology), includes two suborders—Sminthuridida (e.g., family Sminthurididae) and Appendiciphora (e.g., families Katiannidae, Dicyrtomidae)—reflecting a revision that rejects the broader traditional Symphypleona sensu lato. Poduromorpha and Entomobryomorpha together form the Arthropleona , with elongated bodies (arthropleonous morphology); Poduromorpha comprises three superfamilies (Onychiuroidea, Hypogastruroidea, Poduroidea), while Entomobryomorpha includes Tomoceroidea, Isotomoidea, and Entomobryoidea. Phylogenetic analyses based on mitochondrial genomes from 124 across 24 subfamilies and 16 families support the of the four orders, with a placing Neelipleona as sister to the remaining taxa, followed by , and then the clade (Poduromorpha + Entomobryomorpha). This configuration indicates rapid diversification at the base of Collembola, refuting some traditional groupings such as undivided Isotomidae subfamilies and broad , while necessitating revisions in superfamilies like Isotomoidea. Earlier divisions into six informal groups (e.g., Arthropleona, Poduromorpha) have been supplanted by this cladistic framework, informed by molecular data alongside morphology.

Evolutionary Origins and Fossil Record

Collembola occupy a basal position within , with recent phylogenomic analyses using site-heterogeneous models supporting as the earliest-diverging lineage, followed by a clade comprising Collembola sister to , and that combined group sister to Insecta. This configuration revises traditional (now excluding ) and highlights ongoing debates, including support for alternative groupings like Ellipura (Collembola + ) in some site-homogeneous models and datasets. Their evolutionary origins trace to early hexapod diversification, with physiological traits such as haemolymph composed mainly of and exhibiting high osmotic pressures indicating direct descent from marine ancestors rather than via freshwater intermediates. The fossil record of Collembola remains limited owing to their minute size (typically under 2 mm) and lack of , resulting in few compressions and a reliance on exceptional preservational environments like and siliceous sediments. The oldest confirmed specimens occur in the of , dated to approximately 407 million years ago (, Pragian stage), including Rhyniella praecursor, a 1.5-mm-long form with antennal, leg, and furca-like structures resembling extant Poduromorpha. Initially described in 1926 from head capsules and later reassessed as a collembolan rather than an , R. praecursor evidences early terrestrialization among arthropods. Post-Devonian fossils are predominantly from amber, documenting taxonomic diversification; for instance, (Late , ~100 Ma) deposits from yield 93 specimens across multiple morphotypes, while mid-Cretaceous (~99 Ma) preserves over 100 species, reflecting adaptations like phoresy on larger arthropods. Miocene amber (~16 Ma) from the reveals behavioral persistence, with springtails attached to and hosts, suggesting conserved dispersal strategies. Overall, the record spans ~400 million years but underscores evolutionary conservatism, with living forms retaining Devonian-grade traits absent in derived .

Morphology and Physiology

External Features

Springtails exhibit a distinctive consisting of a head capsule, a with three segments, and an typically comprising up to six segments, though segmentation is reduced or fused in some taxa. They are primitively wingless hexapods lacking compound eyes, with simple ocelli present in many . The body ranges in size from 0.1 mm to 10 mm, covered by a thin, flexible that may feature scales, setae, or granular structures for protection and sensory functions. The head bears a pair of antennae with four to six segments, used for chemosensation and mechanoreception, and entognathous mouthparts retracted within a buccal pouch, adapted for or feeding depending on the . The thorax supports three pairs of legs, each comprising a coxa, , and fused tibiotarsus ending in a and empodium; these appendages facilitate locomotion across varied substrates. Abdominal features include the collophore, a ventral tube on the first abdominal segment functioning in adhesion and water regulation, and the , a forked on the fourth or fifth segment that enables jumping via rapid release from a retinaculum on the third segment. Body form varies phylogenetically: elongate in groups like Entomobryomorpha and Poduromorpha, or globular in due to abdominal-thoracic fusion. External coloration arises from pigments or structural elements, often cryptic for habitats.

Sensory and Locomotory Adaptations

Springtails exhibit sensory adaptations suited to their terrestrial microhabitats, primarily involving chemoreception, mechanoreception, and limited vision. The antennae, typically four-segmented, are equipped with tactile setae and chemoreceptors on multiple segments, facilitating detection of mechanical stimuli and chemical cues such as pheromones or environmental volatiles. Many species possess a postantennal organ (PAO), a specialized chemosensory structure posterior to the antennae that detects chemical molecules and compensates for reduced visual input in some taxa. Visual organs include reduced compound eyes with a maximum of eight ommatidia per side, often supplemented by a dorsal light-sensing organ between the antennae for basic phototaxis. Sensory setae distributed across the , especially on antennae, tarsi, and ventral structures, further enhance tactile and chemosensory capabilities, with antennal length and PAO presence influencing overall environmental sensitivity. Locomotory adaptations center on the , a tail-like unique to Collembola that enables ballistic as an escape mechanism. The , comprising a manubrium, dens, and mucro, folds ventrally against the fourth abdominal segment and latches to the retinaculum; muscular contraction stores , followed by rapid release that propels the springtail upward and backward, achieving rotations and distances proportional to body size (up to several centimeters in larger ). This catapult mechanism relies on cuticular elasticity rather than direct muscle power for propulsion, allowing jumps in varied directions despite post-jump tumbling. Ambulatory movement occurs via six thoracic legs adapted for crawling and on substrates, with tarsal setae aiding ; however, predominates for evasion, as sustained locomotion fatigues these small arthropods quickly. Habitat-specific variations, such as elongated furcae in arboreal , optimize jump trajectories for vertical escape.

Internal Anatomy

The digestive system of springtails consists of a tubular alimentary canal divided into , , and regions, extending from the to the anus along the body length. The and are ectodermal invaginations lined with , while the is endodermal and surrounded by a network of muscles that facilitate and nutrient absorption through microvilli-lined . The is an open type, with filling the (hemocoel) and bathing internal organs directly, lacking closed vessels in most species. A dorsal heart, typically a simple tubular vessel located in the , propels hemolymph anteriorly through ostia valves, though this structure is reduced or absent in miniaturized species such as Mesaphorura sylvatica, where circulation relies primarily on body movements. Respiration occurs primarily through cutaneous across the thin, porous , supplemented by via the eversible vesicles of the collophore (ventral tube) on the first abdominal segment, which increases surface area for oxygen uptake in moist environments. Unlike most hexapods, springtails generally lack tracheae, though rudimentary tracheal systems are present in families like Sminthuridae and Actaletidae. Excretion is handled by paired labial nephridia, tubular glands located in the posterior head region, which produce as the primary nitrogenous waste and discharge it via nephridiopores near the mouthparts; Malpighian tubules are absent. The features a dorsal in the head, comprising fused protocerebrum, deutocerebrum, and tritocerebrum connected to a subesophageal , linked by circumenteric connectives to a ventral nerve cord with segmental in the and . This chain supports sensory integration, including connections to antennal and postantennal organs. Reproductive organs include paired ovaries or testes in the abdomen, with gonoducts converging to a genital opening on the fifth abdominal segment; females possess a spermatheca for sperm storage in indirect transfer via spermatophores. A complex muscular system pervades the body, including longitudinal and transverse fibers for locomotion, a specialized retractor muscle for the furcula (jumping appendage), and intrinsic muscles in appendages and the collophore for eversion and adhesion. The fat body, interspersed with muscles and hemolymph spaces, serves storage and metabolic functions, occupying a significant portion of the body volume in small species.

Genetics and Genomics

Genome Characteristics

Springtail exhibit variation in size, typically ranging from 150 to 400 megabase pairs (Mbp), reflecting diversity across Collembola . numbers generally fall between 5 and 11 pairs, contributing to compact karyotypes compared to many other arthropods. This low count facilitates chromosomal-level assemblies in several sequenced , often revealing 5 to 6 pseudochromosomes. High-quality genome assemblies have been produced for model species, enabling detailed genomic analyses. The parthenogenetic Folsomia candida, widely used in ecotoxicology, has an assembled of approximately 221 Mbp across 113–162 scaffolds, with a GC content of 37.5% and evidence of functional parthenogenesis without recent hybridization. In contrast, the sexually reproducing Orchesella cincta features a draft of 283.8 Mbp, with expansions in gene families linked to soil stress responses such as detoxification and DNA repair. Entomobrya proxima yields a 362.37 Mbp assembly, 97% anchored to six chromosomes with a scaffold N50 of 57.67 Mbp. Tomocerus qinae has a 334.44 Mbp forming five chromosomes. Certain species display specialized systems, as in Allacma fusca, where the 392.8 Mbp scaffolds into six pseudomolecules including X1 and X2 chromosomes. Genomic studies also highlight features like paternal genome elimination in some lineages and cryptic driven by , underscoring adaptive in habitats. Overall, these characteristics reveal Collembola's genetic compactness and resilience, with relatively low GC contents (often below 40%) and scaffold efficiencies improving via long-read technologies like PacBio.

Genetic Diversity and Adaptations

Springtails (Collembola) demonstrate substantial genetic diversity across populations, often exceeding thresholds for species delineation, with mitochondrial COI sequence divergences of 5–11.3% reported in taxa such as Cryptopygus antarcticus, Isotomurus cf. frater, and Parisotoma octooculata, suggesting cryptic driven by historical isolation. In subterranean habitats like Australian calcretes, genetic analyses of species including Bourginsonia sp. and Troglopedetes sp. reveal extreme intraspecific variation, with less than 5% of sampled calcretes harboring highly divergent lineages indicative of numerous undescribed species adapted to fragmented aquifers. Population genetic studies of soil-dwelling species, such as Orchesella cincta, indicate that 96.5% of total variation occurs within populations, reflecting high local heterozygosity despite limited dispersal, as measured by markers across metal-polluted and reference sites. This genetic variability facilitates adaptations to harsh terrestrial environments. in Orchesella cincta shows expansion of families involved in metabolism and stress response, correlating with tolerance to contaminants like and , where duplicated and genes enhance detoxification efficiency. Transcriptomic analyses across Collembola species highlight evolutionary shifts in hexapod genes for sclerotization and resistance, with upregulated aquaporins and heat shock proteins in arid-adapted lineages enabling survival in low-water s. In polar species, high genetic diversity supports physiological plasticity, including broad thermal tolerance ranges (-30°C to +15°C in some Cryptopygus populations) and behavioral , as evidenced by variant alleles in genes linked to cold acclimation. Mitochondrial genome studies further underscore adaptive divergence, with four distinct gene arrangements (e.g., GO1 predominant in Entomobryomorpha) reflecting phylogenetic splits and rearrangements that may optimize metabolism under fluctuating oxygen in litter layers. assemblies, such as that of Folsomia candida, reveal chromosomal stability with adaptations in sensory and reproductive , including parthenogenesis-promoting loci that maintain diversity in uniparental lineages despite reduced recombination. Overall, this genetic architecture enables Collembola to occupy diverse microhabitats, from Arctic tundra to metal-enriched soils, with ongoing cryptic inferred from divergence levels of 1.7–14.7% in multi-species surveys.

Reproduction and Life History

Mating Behaviors and Strategies

Springtails (Collembola) primarily utilize indirect sperm transfer for , wherein males deposit stalked spermatophores on substrates such as , litter, or water surfaces, which females subsequently uptake using their genital openings, obviating direct copulation. This prevails across taxa, with males often placing spermatophores individually or in clusters to increase uptake probability. Fertilization success hinges on environmental conditions and behavioral cues guiding females to the deposits, reflecting adaptations to terrestrial or semi-aquatic microhabitats. Courtship rituals exhibit considerable variation by family and habitat. In , particularly Sminthurididae, males engage in elaborate, dance-like maneuvers on water surfaces, employing modified antennae as clasping organs to secure females and position them over spermatophores, as documented in extant Sminthurus aquaticus (males 0.3–0.4 mm) and Pseudosminthurides stoechus (~105 million years old). These behaviors, accompanied by in antennal structure, enhance precision in aquatic environments where spermatophores risk dispersal. Conversely, in Entomobryomorpha like Isotomidae, may involve simpler aggregative tendencies or novel tactile interactions, such as males using antennae to fasten the female's during approach, though less ritualized than in globular forms. Male-male competition shapes strategies, particularly in species like Orchesella cincta (), where rival presence triggers ejaculate economy: males deposit fewer but render them more attractive to females, as evidenced by preferential uptake in lab assays, thereby optimizing under risk. This plasticity underscores causal trade-offs between quantity and quality in mating effort. Aggregations, sometimes exceeding 45 individuals as in fossil Proisotoma communis, may facilitate synchronized reproductive episodes, amplifying encounter rates in litter or . Although sexual amphimixis dominates, parthenogenetic reproduction occurs in certain populations and , bypassing entirely and altering effective strategies toward uniparental clonal propagation, with reports across Collembola indicating variable ratios influenced by environmental or genetic factors. Such facultative shifts highlight reproductive flexibility, though detailed behaviors remain centered on spermatophore-mediated in gonochoristic lineages.

Developmental Stages and Life Cycle

Springtails (Collembola) exhibit ametabolous development, lacking distinct larval, pupal, or other metamorphic stages typical of many ; instead, eggs hatch directly into juveniles that morphologically resemble smaller versions of adults. This direct allows for continuous growth through without radical morphological shifts. Juveniles possess functional mouthparts, appendages, and sensory structures from hatching, enabling immediate feeding and dispersal akin to adults. Eggs are typically spherical or ovoid, measuring 0.1–0.2 in diameter, and are deposited in moist , , or on substrates in clutches of 10–100, depending on species and conditions; incubation periods range from 2–20 days, strongly influenced by and . Upon , neonates are minute (often <0.5 ) and undergo 3–8 instars to reach , with each molt increasing body size by 20–50% and refining structures like the for jumping. The number of juvenile instars varies phylogenetically and environmentally; for instance, in Folsomia candida, development from hatch to first averages 15–42 days at 20–26°C, encompassing 4–5 molts. Maturation to adulthood occurs post-final juvenile molt, with adults retaining the capacity for lifelong molting—up to annually or more under stress—unlike most hexapods, facilitating repair, growth, or reproductive adjustments. Total life cycle duration spans weeks to months across species, with parthenogenetic forms like Folsomia achieving multiple generations annually under favorable conditions (e.g., 20–30°C, high moisture), while sexual species may extend cycles due to mate location. Environmental factors, such as drops below 5°C, induce diapause-like delays in development, enhancing survival in temperate zones. This flexible, non-metamorphic strategy supports their abundance in diverse microhabitats, with generation times adapting to resource availability and predation pressures.

Ecology and Behavior

Habitats and Global Distribution

Springtails (Collembola) primarily occupy moist, organic-rich terrestrial habitats, including , leaf litter, , decaying wood, and under bark, where they contribute to processes. They thrive in environments with high humidity, such as forest floors, grasslands, and agricultural s, with population densities often exceeding 10,000 individuals per square meter in optimal conditions, though lower in drier or disturbed sites like woodlands and croplands compared to scrublands. Some exhibit semi-aquatic adaptations, inhabiting freshwater edges, wet , or marine littoral zones, while others, such as "snow fleas," form visible aggregations on snow surfaces during winter in temperate and polar regions. Collembola display a cosmopolitan global distribution, occurring on all continents from the to , and spanning elevations from to over 5,000 meters on mountain peaks. They represent approximately 32% of global terrestrial abundance, with highest biomasses in ecosystems worldwide, adapting to diverse biomes including tundras, deserts (under stones), and caves. Species diversity peaks in temperate regions like , but abundance remains high in polar areas, where they dominate microarthropod communities despite lower taxonomic richness. Vertical distribution within soils varies by species and microhabitat, with many confined to upper litter layers but some utilizing earthworm burrows or deeper profiles to evade desiccation. Their broad habitat tolerance, driven by physiological adaptations to moisture and temperature, enables persistence in human-modified landscapes, though local abundances fluctuate with edaphic factors like organic matter content and pH.

Trophic Interactions and Diet

Collembola primarily function as detritivores and microbivores in soil ecosystems, feeding on decomposing , fungal hyphae, , , and associated organic particles. Gut content analyses reveal that diets often consist of fungal spores and mycelia, humified , and microbial biofilms, with species-specific preferences influenced by depth and life form. Surface-active epedaphic species, such as those in , preferentially graze on fresh litter and epigeic fungi during early decomposition stages, while euedaphic forms like Onychiuridae process deeper, more stabilized and microbes. Feeding strategies exhibit considerable flexibility, with many species classified as omnivores capable of shifting between detrital, fungal, and bacterial resources based on availability. Stable isotope studies, including and profiling combined with gut dissections, demonstrate multichannel trophic niches, where Collembola integrate from both detrital and microbial pathways, often occupying positions as secondary decomposers. of field-collected specimens has shown incorporation of recently fixed carbon (up to contemporary levels), indicating substantial reliance on exudates, living tissues, or associated fresh organic inputs rather than exclusively aged , challenging traditional views of them as strict saprophages. Although predation is uncommon, certain taxa, such as some Neanuridae and poduromorph species, opportunistically consume nematodes, , or conspecifics, elevating their trophic position in microbial food webs. These interactions contribute to population regulation of and enhance nutrient mineralization. In broader trophic networks, Collembola suppresses fungal and alters microbial community structure, accelerating litter breakdown rates by 10–30% in experiments while serving as basal prey for meso- and macro-predators like predatory mites and spiders, thus channeling energy upward.

Predators, Parasites, and Defenses

Springtails serve as prey for numerous predators, including spiders (such as linyphiids and lycosids), carabid beetles, mesostigmatid mites, and , which actively hunt them in and habitats. predators encompass amphibians like salamanders and frogs, small birds, and larval in aquatic environments, with predation rates varying by habitat density and season. In agroecosystems, Collembola constitute a significant portion of diets for generalist predators, supporting their populations during periods of low alternative prey availability. Parasitic interactions with springtails include ectoparasitic mites, such as those infesting cave-dwelling species like Trogolaphysa (Paronellidae), where mites attach externally and feed on host fluids. Endoparasites encompass protozoans, nematodes, trematodes, and that infect Collembola as intermediate or definitive hosts. Endosymbiotic like Wolbachia occur in various Collembola species, inducing effects ranging from reproductive manipulation to potential pathogenicity, though interactions are context-dependent and not uniformly deleterious. Defensive adaptations in springtails primarily involve the , a tail-like appendage that enables rapid ballistic jumping up to 10-20 cm or more, allowing escape from approaching predators by propelling the animal away at speeds exceeding 2 m/s in some . Certain taxa, such as Tetrodontophora bielanensis, deploy chemical defenses via pseudocells that release a sticky containing pyridopyrazines, which disorients predators and triggers reflexive grooming behaviors. Other exhibit reflex bleeding of laced with deterrents like or highly substituted benzenes, repelling attackers through toxicity or aversion. Unique epicuticular lipids, including higher and esters distinct from those in , contribute to passive protection against predation and environmental stressors by altering surface chemistry. These mechanisms collectively enhance survival in predator-rich microhabitats, though efficacy varies by and threat type.

Human Interactions

Beneficial Roles in Ecosystems and Agriculture

Springtails, or Collembola, contribute to ecosystem functioning primarily through their role as detritivores, accelerating the of organic litter and facilitating nutrient cycling in . By consuming fungal hyphae, , and plant detritus, they enhance microbial processes, promoting the mineralization of organic carbon and into plant-available forms. This activity stimulates soil microbial communities, particularly fungi, thereby improving overall and supporting in terrestrial . In and ecosystems, collembolan on increases carbon loss and transforms organic material into higher-quality resources for further breakdown, influencing plant growth indirectly through enhanced nutrient availability. Their burrowing and feeding behaviors also contribute to aggregation and , albeit on a micro scale, aiding infiltration and root penetration. Studies indicate that collembolans can account for significant portions of breakdown rates, underscoring their integral position in belowground food webs and energy flow. Within agricultural systems, springtails bolster by mirroring benefits, with elevated abundances—up to 20 times higher in no-till versus conventional —indicating effective and reduced disturbance. They function as bioindicators of , sensitive to land-use changes and pollutants, enabling assessments of ; for instance, species diversity and reproduction rates in tests with Folsomia candida reveal chemical impacts on soil biota. In rubber plantations and similar , their presence correlates with improved decomposition under varied , supporting nutrient retention without synthetic inputs.

Perceptions as Pests and Management

Springtails (Collembola) are infrequently regarded as agricultural or structural pests, primarily due to their role as decomposers of rather than direct herbivores or vectors of . In residential settings, they often invade homes during periods of high , such as after heavy rainfall, congregating in basements, bathrooms, and around where damp conditions prevail, leading to perceptions of from their high numbers—sometimes numbering in the thousands—though they cause no structural damage, , or risks beyond aesthetic . In controlled environments like greenhouses or cultivation facilities, certain may feed on germinating , fungal mycelia, or decaying plant material, occasionally resulting in minor seedling damage under excessive , but such impacts are rare and overshadowed by their beneficial decomposition activities. Management of springtail populations emphasizes (IPM) principles, prioritizing environmental modifications to eliminate favorable conditions over chemical interventions, as insecticides show limited efficacy indoors and are unnecessary for outdoor minor issues. Key strategies include reducing through improved ventilation, dehumidifiers, leak repairs, and allowing or to dry out, which disrupts their requirements. Exclusion measures, such as caulking cracks in foundations, screening vents, and minimizing organic debris like litter near buildings, prevent entry and breeding sites. In agricultural contexts, cultural practices like avoiding overwatering and incorporating drainage further mitigate risks without relying on broad-spectrum pesticides, which can harm beneficial biota. For persistent indoor clusters, non-toxic options such as vacuuming followed by soapy water application provide short-term control comparable to chemicals but without associated risks or costs. Perimeter barrier treatments with insecticides may be applied outdoors to deter migration toward structures, though their use is secondary to habitat alteration. Overall, patience and control often suffice, as populations decline naturally with drying conditions.

Applications in Scientific Research

Springtails (Collembola) are widely employed as model organisms in owing to their high sensitivity to pesticides and environmental contaminants, particularly insecticides, with LC50 values indicating greater compared to fungicides. Species such as Folsomia candida and Folsomia fimetaria undergo standardized life-cycle toxicity tests, including survival, , and development assays, to evaluate and other chemical impacts under controlled conditions. These organisms have been utilized in such assessments since the , providing insights into sublethal effects like reduced and altered in contaminated s. In ecological research, springtails function as bioindicators for assessing and anthropogenic disturbances, with population responses to intensity, applications, and organic amendments revealing influences on and activity . For instance, higher frequencies correlate with increased springtail , while incorporation affects feeding dynamics on fungi and . Physiological studies further explore combined stressors, such as polycyclic aromatic hydrocarbons like alongside , elucidating molecular and metabolic adaptations in species like Folsomia candida. Biomechanical investigations leverage the springtail's appendage for analyzing latch-mediated spring actuation, enabling explosive jumps up to 10 cm despite body lengths under 2 mm. High-speed imaging and kinematic models demonstrate precise directional takeoff, mid-air righting via collophore adhesion, and stable landings, principles validated in bioinspired microrobots that achieve upright orientation in 75% of jumps. These mechanisms, involving hydrostatic and nanotopographical cuticles for reduced bioadhesion, inform micro-robotics and applications mimicking anti-wetting surfaces. Emerging research examines specialized traits, such as endogenous in Lobella sauteri for potential underground signaling studies, and cuticular for analyses. Overall, springtails' parthenogenetic , short generation times, and ease of culturing facilitate their role in both and field experiments across these disciplines.

Environmental Responses

Effects of Climate Variability

Springtails exhibit varied responses to climate variability, including shifts in , , and community composition, primarily driven by alterations in and patterns. Experimental warming treatments have been observed to increase Collembola and abundance across multiple seasons in temperate ecosystems, except during early spring, suggesting enhanced activity and reproduction under moderate temperature elevations. However, long-term exposure to elevated temperatures in forest soils correlates with statistically significant influences from cumulative positive air temperatures on , potentially altering developmental rates and locomotory activity in habitat-specific ways, such as greater sensitivity in forest-dwelling species compared to open habitats. Precipitation deficits and drought impose more pronounced constraints than temperature fluctuations alone, often reducing total abundance, taxonomic diversity, and biomass, particularly in acidic soils where aluminum ions may exacerbate stress. In shrubland and grassland systems, combined warming and drought treatments significantly decrease Collembola density and biomass, with epigeic species demonstrating higher drought resistance and dispersal capacity to seek favorable microhabitats compared to eudaphic forms. Extreme drought events can modify functional traits and community structure at regional scales, with land use moderating resilience—intensively managed areas showing greater vulnerability than natural habitats. In polar regions, climate variability introduces complex stressors; mid-winter under warmer conditions exposes springtails to lethal cold snaps, heightening mortality despite overall warming trends, while and basal thermal tolerances enable recovery and adaptation in some populations. Alien invasive Collembola often display superior critical thermal maxima compared to indigenous species, potentially conferring advantages in warming scenarios and influencing invasion success under altered climates. Overall, Collembola communities demonstrate functional resilience to moderate variability but face risks from intensified extremes, with water availability emerging as a dominant regulator over in many contexts.

Responses to Pollution and Anthropogenic Stressors

Springtails exhibit high sensitivity to contaminants, often experiencing reduced , growth inhibition, and shifts in community structure, which positions them as effective bioindicators of . Studies demonstrate that heavy metal , such as , , and lead, significantly impairs Collembola populations; for instance, contamination leads to decreased body sizes after 14 days of exposure, with greater from As(III) than As(V). Similarly, elevated heavy metal levels in urban soils correlate with lower and abundance, though some tolerant species may persist or dominate in polluted sites. Pesticide exposure further exacerbates these effects, with neonicotinoids like reducing surface- and soil-dwelling springtail abundances by 65–90% in field margins at higher concentrations. Insecticides such as teflubenzuron disrupt lipid metabolism in species like Folsomia candida, altering life history traits including and survival. Realistic mixtures of pesticides observed in agricultural sites also induce , highlighting the compounded risks from multiple chemical stressors. Pre-exposure to pollutants like or fungicides increases vulnerability to , amplifying mortality under combined anthropogenic pressures. Air pollution and urban stressors influence springtail communities independently of soil heavy metals, with species richness declining in areas of higher atmospheric . Remediation efforts, such as application, show potential to mitigate heavy metal impacts by enhancing conditions and supporting population recovery. Overall, these responses underscore Collembola's utility in monitoring anthropogenic disturbances, as their abundance and diversity reliably reflect gradients across urban and agricultural landscapes.

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