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Springtails (Collembola) form the largest of the three lineages of modern hexapods that are no longer considered
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...
s (the other two are the
Protura The Protura, or proturans, and sometimes nicknamed coneheads, are very small (0.6-1.5mm long), soil-dwelling animals, so inconspicuous they were not noticed until the 20th century. The Protura constitute an order of hexapods that were previously ...
and
Diplura The order Diplura ("two-pronged bristletails") is one of three orders of non-insect hexapods within the class Entognatha (alongside Collembola (springtails) and Protura). The name "diplura", or "two tails", refers to the characteristic pair of ...
). Although the three orders are sometimes grouped together in a class called Entognatha because they have internal
mouthparts Mouthparts may refer to: * The parts of a mouth ** Arthropod mouthparts The mouthparts of arthropods have evolved into a number of forms, each adapted to a different style or mode of feeding. Most mouthparts represent modified, paired append ...
, 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 An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nut ...
, 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 and the control of soil microbial communities. The word ''Collembola'' is from the ancient Greek "glue" and "peg"; this name was given due to the existence of the
collophore A collophore is a tube-like structure on the ventral side of the first abdominal segment of the body of springtails (collembolans). It used to be believed that it served to stabilise the animal when it jumped by sticking to the surface on which i ...
, which was previously thought to stick to surfaces to stabilize the creature. Some
DNA sequence DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. T ...
studies suggest that Collembola represent a separate evolutionary line from the other
Hexapoda The subphylum Hexapoda (from Greek for 'six legs') comprises most species of arthropods and includes the insects as well as three much smaller groups of wingless arthropods: Collembola, Protura, and Diplura (all of these were once considere ...
, but others disagree; this seems to be caused by widely divergent patterns of
molecular evolution Molecular evolution is the process of change in the sequence composition of cellular molecules such as DNA, RNA, and proteins across generations. The field of molecular evolution uses principles of evolutionary biology and population genet ...
among the
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s. The adjustments of traditional
taxonomic rank In biological classification, taxonomic rank is the relative level of a group of organisms (a taxon) in an ancestral or hereditary hierarchy. A common system consists of species, genus, family, order, class, phylum, kingdom, domain. While ...
for springtails reflects the occasional incompatibility of traditional groupings with modern
cladistics Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived cha ...
: when they were included with the insects, they were ranked as an
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
; 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 Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differently ...
status.


Morphology

Members of the Collembola are normally less than long, have six or fewer
abdominal The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the to ...
segments, and possess a tubular
appendage An appendage (or outgrowth) is an external body part, or natural prolongation, that protrudes from an organism's body. In arthropods, an appendage refers to any of the homologous body parts that may extend from a body segment, including ante ...
(the
collophore A collophore is a tube-like structure on the ventral side of the first abdominal segment of the body of springtails (collembolans). It used to be believed that it served to stabilise the animal when it jumped by sticking to the surface on which i ...
or ventral tube) with reversible, sticky vesicles, projecting ventrally from the first abdominal segment. It is believed to be associated with fluid uptake and balance, excretion, and orientation of the organism itself. Most species have an abdominal, tail-like appendage known as a
furcula The (Latin for "little fork") or wishbone is a forked bone found in most birds and some species of non-avian dinosaurs, and is formed by the fusion of the two pink clavicles. In birds, its primary function is in the strengthening of the thoracic ...
. 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 A retinaculum (plural ''retinacula'') is a band of thickened deep fascia around tendons that holds them in place. It is not part of any muscle. Its function is mostly to stabilize a tendon. The term retinaculum is New Latin, derived from the Latin ...
(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. Springtails also possess the ability to reduce their body size by as much as 30% through subsequent
ecdyses Ecdysis is the moulting of the cuticle in many invertebrates of the clade Ecdysozoa. Since the cuticle of these animals typically forms a largely inelastic exoskeleton, it is shed during growth and a new, larger covering is formed. The remna ...
(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 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 Porosity or void fraction is a measure of the void (i.e. "empty") spaces in a material, and is a fraction of the volume of voids over the total volume, between 0 and 1, or as a percentage between 0% and 100%. Strictly speaking, some tests measure ...
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
, except for the two families
Sminthuridae Sminthuridae is a family of springtails of the order Symphypleona. Sminthurids are commonly referred to as globular springtails. Description Like other Symphypleona, Sminthuridae are globular in shape and have a furcula that allows them to ju ...
and Actaletidae, which exhibit a single pair of spiracles between the head and the
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the c ...
, leading to a rudimentary, although fully functional, tracheal system. The anatomical variance present between different species partially depends on soil morphology and composition. Surface-dwellers are generally larger, have darker pigments, have longer antennae and functioning furcula. Sub-surface-dwellers are usually unpigmented, have elongated bodies, and reduced furcula. They can be categorized into four main forms according to soil composition and depth: atmobiotic, epedaphic, hemiedaphic, and euedaphic. Atmobiotic species inhabit macrophytes and litter surfaces. They are generally 8-10 millimeters in length, pigmented, have long limbs, and a full set of
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
(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 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. Poduromorphs inhabit the epedaphic, hemiedaphic, and euedaphic layers and are characterized by their elongated bodies and conspicuous segmentation – three thoracic segments, six abdominal segments, and a
prothorax The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum ( dorsal), the prosternum ( ventral), and the propleuron ( lateral) o ...
. 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.


Systematics and evolution

Traditionally, the springtails were divided into the
orders Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
Arthropleona, Symphypleona, and occasionally also Neelipleona. The Arthropleona were divided into two superfamilies, the
Entomobryoidea :''Entomobryoidea is also the old name of Entomobryomorpha, when these were placed in the "Arthropleona" The Entomobryoidea are a superfamily of springtails (Collembola), tiny hexapods related to insects. In the modern sense, this group is plac ...
and the
Poduroidea Poduridae is a small family of stout-bodied springtails containing only the single genus ''Podura'', and making up the monotypic superfamily Poduroidea. The genus contains four species: *''Podura aquatica'' Linnaeus, 1758 *''Podura infernalis'' ...
. However, recent phylogenetic studies show Arthropleona is
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
. 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 The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linn ...
of the Collembola. The term "Neopleona" is essentially synonymous with Symphypleona + Neelipleona. 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 28S ribosomal RNA is the structural ribosomal RNA (rRNA) for the large subunit (LSU) of eukaryotic cytoplasmic ribosomes, and thus one of the basic components of all eukaryotic cells. It has a size of 25S in plants and 28S in mammals, hence th ...
rRNA Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from ribos ...
sequence In mathematics, a sequence is an enumerated collection of objects in which repetitions are allowed and order matters. Like a set, it contains members (also called ''elements'', or ''terms''). The number of elements (possibly infinite) is called ...
data, though, suggests that they form the most ancient lineage of springtails, which would explain their peculiar apomorphies. This phylogenetic relationship was also confirmed using a phylogeny based on
mtDNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial DNA ...
and whole-genome data. The latest whole-genome phylogeny supporting four orders of collembola: Springtails are attested to since the
Early Devonian The Early Devonian is the first of three epochs comprising the Devonian period, corresponding to the Lower Devonian series. It lasted from and began with the Lochkovian Stage , which was followed by the Pragian from and then by the Emsian, ...
. The fossil from , ''
Rhyniella praecursor ''Rhyniella'' is a genus of fossil springtails (Collembola) from the Rhynie chert, which formed during the Pragian stage of the Early Devonian. One species has been described, ''Rhyniella praecursor''. For some time it was believed to be the only ...
'', is the oldest terrestrial arthropod, and was found in the famous
Rhynie chert The Rhynie chert is a Lower Devonian sedimentary deposit exhibiting extraordinary fossil detail or completeness (a Lagerstätte). It is exposed near the village of Rhynie, Aberdeenshire, Scotland; a second unit, the Windyfield chert, is located ...
of
Scotland Scotland (, ) is a country that is part of the United Kingdom. Covering the northern third of the island of Great Britain, mainland Scotland has a border with England to the southeast and is otherwise surrounded by the Atlantic Ocean to ...
. Given its morphology resembles extant species quite closely, the radiation of the
Hexapoda The subphylum Hexapoda (from Greek for 'six legs') comprises most species of arthropods and includes the insects as well as three much smaller groups of wingless arthropods: Collembola, Protura, and Diplura (all of these were once considere ...
can be situated in the
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleoz ...
, or more. Additional research concerning the coprolites (fossilized feces) of ancient collembolans allowed researchers to track their lineages back some 412 million years. 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 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.


Ecology


Eating behavior

Specific feeding strategies and mechanisms are employed to match specific niches. 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. Carnivorous species maintain populations of small invertebrates such as nematodes, rotifers, and other collembolan species. 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.


Predators

Springtails are consumed by mesostigmatan mites in various families, including
Ascidae Ascidae is a family of mites in the order Mesostigmata. Description Ascidae are mites characterised by: seta ''st4'' usually on unsclerotised cuticle, peritrematic shield broadly connected to exopodal shield beside coxa IV, fixed cheliceral d ...
,
Laelapidae The Laelapidae are a family of mites in the order Mesostigmata. The family is also referred to in the literature as Laelaptidae, which may be the correct spelling. Description Laelapidae have a shield covering all or most of the dorsal surface ...
,
Parasitidae Parasitidae is a family of predatory mites in the order Mesostigmata that has worldwide distribution. They are the only family in the superfamily Parasitoidea. Relatively large for mites, their color is often yellowish to dark brown. The family a ...
,
Rhodacaridae Rhodacaridae is a family of mites in the order Mesostigmata Mesostigmata is an order of mites belonging to the Parasitiformes. They are by far the largest group of Parasitiformes, with over 8,000 species in 130 families. Mesostigmata includes ...
and Veigaiidae. Cave-dwelling springtails are a food source for
spiders Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species di ...
and
harvestmen The Opiliones (formerly Phalangida) are an order of arachnids colloquially known as harvestmen, harvesters, harvest spiders, or daddy longlegs. , over 6,650 species of harvestmen have been discovered worldwide, although the total number of extan ...
in the same environment, such as the endangered harvestman ''
Texella reyesi ''Texella reyesi'' is a rare species of arachnid known by the common name Bone Cave harvestman. It is endemic to Texas in the United States, where it lives in limestone caves in Travis and Williamson Counties. It is threatened by the loss of it ...
''.


Distribution

Springtails are cryptozoa frequently found in
leaf litter Plant litter (also leaf litter, tree litter, soil litter, litterfall or duff) is dead plant material (such as leaves, bark, needles, twigs, and cladodes) that have fallen to the ground. This detritus or dead organic material and its constituent ...
and other decaying material, where they are primarily
detritivores Detritivores (also known as detrivores, detritophages, detritus feeders, or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrates, ...
and microbivores, and one of the main biological agents responsible for the control and the dissemination of soil
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in old ...
s. 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. 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 Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta ('' sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and ...
cushions, fallen
wood Wood is a porous and fibrous structural tissue found in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulose fibers that are strong in tension and embedded in a matrix of lignin ...
,
grass Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns a ...
tufts,
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...
and
termite Termites are small insects that live in colonies and have distinct castes ( eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blat ...
nests) occur. Only
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant- parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a bro ...
s,
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapoda, decapods, ostracoda, seed shrimp, branchiopoda, branchiopods, argulidae, fish lice, krill, remipedes, isopoda, isopods, barnacles, copepods, ...
s, and
mite Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear e ...
s 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 In biology, a phylum (; plural: phyla) is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature ...
and crustaceans a
subphylum In zoological nomenclature, a subphylum is a taxonomic rank below the rank of phylum. The taxonomic rank of " subdivision" in fungi and plant taxonomy is equivalent to "subphylum" in zoological taxonomy. Some plant taxonomists have also used th ...
. Most springtails are small and difficult to see by casual observation, but one springtail, the so-called
snow flea Snow flea is a common name for several arthropods, not including true fleas: * ''Boreidae'', a family of scorpionflies known as snow fleas in the British Isles, especially: ** ''Boreus hyemalis'' a species of scorpionfly that crawls on snow * ''H ...
(''
Hypogastrura nivicola ''Hypogastrura nivicola'' is a species of dark blue springtail. Its (US) English name is snow flea, but there are also insects called by that name. They are often seen jumping about on the surface of snow on a warm winter day in North America. Re ...
''), 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 ''Orchesella'' is a genus of springtails belonging to the family Entomobryidae. This genus includes springtails with subdivided basal antennal segments. Species These 96 species belong to the genus ''Orchesella'': * '' Orchesella adriatica'' S ...
'', 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. The main
ecological factor An environmental factor, ecological factor or eco factor is any factor, abiotic or biotic, that influences living organisms. Abiotic factors include ambient temperature, amount of sunlight, and pH of the water soil in which an organism lives. ...
driving the local distribution of species is the vertical stratification of the environment: in
woodland A woodland () is, in the broad sense, land covered with trees, or in a narrow sense, synonymous with wood (or in the U.S., the '' plurale tantum'' woods), a low-density forest forming open habitats with plenty of sunlight and limited shade (see ...
a continuous change in species assemblages can be observed from tree canopies to ground
vegetation Vegetation is an assemblage of plant species and the ground cover they provide. It is a general term, without specific reference to particular taxa, life forms, structure, spatial extent, or any other specific botanical or geographic charac ...
then to
plant litter Plant litter (also leaf litter, tree litter, soil litter, litterfall or duff) is dead plant material (such as leaves, bark, needles, twigs, and cladodes) that have fallen to the ground. This detritus or dead organic material and its constitue ...
down to deeper
soil horizon A soil horizon is a layer parallel to the soil surface whose physical, chemical and biological characteristics differ from the layers above and beneath. Horizons are defined in many cases by obvious physical features, mainly colour and texture. ...
s. This is a complex factor embracing both
nutritional Nutrition is the biochemical and physiological process by which an organism uses food to support its life. It provides organisms with nutrients, which can be metabolized to create energy and chemical structures. Failure to obtain sufficient nu ...
and
physiological Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemica ...
requirements, together with behavioural trends, dispersal limitation and probable species
interactions Interaction is action that occurs between two or more objects, with broad use in philosophy and the sciences. It may refer to: Science * Interaction hypothesis, a theory of second language acquisition * Interaction (statistics) * Interactions ...
. Some species have been shown to exhibit negative or positive
gravitropism Gravitropism (also known as geotropism) is a coordinated process of differential growth by a plant in response to gravity pulling on it. It also occurs in fungi. Gravity can be either "artificial gravity" or natural gravity. It is a general featu ...
, which adds a
behavioural Behavior (American English) or behaviour (British English) is the range of actions and mannerisms made by individuals, organisms, systems or artificial entities in some environment. These systems can include other systems or organisms as wel ...
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". As a group, springtails are highly sensitive to
desiccation Desiccation () is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. ...
, because of their tegumentary
respiration Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellul ...
, 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 A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
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 Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. Reproduction is a fundamental feature of all known life; each individual or ...
and
survival Survival, or the act of surviving, is the propensity of something to continue existing, particularly when this is done despite conditions that might kill or destroy it. The concept can be applied to humans and other living things (or, hypotheti ...
rates (thereby fitness) could be kept at an optimum. Sensitivity to drought varies from species to species and increases during
ecdysis Ecdysis is the moulting of the cuticle in many invertebrates of the clade Ecdysozoa. Since the cuticle of these animals typically forms a largely inelastic exoskeleton, it is shed during growth and a new, larger covering is formed. The remnan ...
. Given that springtails
moult In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...
repeatedly during their entire life (an
ancestral An ancestor, also known as a forefather, fore-elder or a forebear, is a parent or ( recursively) the parent of an antecedent (i.e., a grandparent, great-grandparent, great-great-grandparent and so forth). ''Ancestor'' is "any person from wh ...
character in
Hexapoda The subphylum Hexapoda (from Greek for 'six legs') comprises most species of arthropods and includes the insects as well as three much smaller groups of wingless arthropods: Collembola, Protura, and Diplura (all of these were once considere ...
) they spend much time in concealed micro-sites where they can find protection against
desiccation Desiccation () is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. ...
and
predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill ...
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 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 In computer science, ACID ( atomicity, consistency, isolation, durability) is a set of properties of database transactions intended to guarantee data validity despite errors, power failures, and other mishaps. In the context of databases, a ...
,
moisture Moisture is the presence of a liquid, especially water, often in trace amounts. Small amounts of water may be found, for example, in the air (humidity), in foods, and in some commercial products. Moisture also refers to the amount of water vapo ...
and
light Light or visible light is electromagnetic radiation that can be perceived by the human eye. Visible light is usually defined as having wavelengths in the range of 400–700 nanometres (nm), corresponding to frequencies of 750–420 t ...
. 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 ''Isotomurus palustris'', the marsh springtail, is a species of elongate-bodied springtail in the family Isotomidae. It is found in Europe, Africa, North America, and Central America. References External links * Collembola Articles ...
''.
Adaptive Adaptation, in biology, is the process or trait by which organisms or population better match their environment Adaptation may also refer to: Arts * Adaptation (arts), a transfer of a work of art from one medium to another ** Film adaptation, a ...
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 ''Podura aquatica'', the water springtail, is a species of springtail, one of only four described species in the family Poduridae. It is an abundant species with a Holarctic distribution. As its common and scientific names suggest, this is an ex ...
'', a unique representative of the family
Poduridae Poduridae is a small family of stout-bodied springtails containing only the single genus ''Podura'', and making up the monotypic superfamily Poduroidea. The genus contains four species: *''Podura aquatica ''Podura aquatica'', the water spring ...
(and one of the first springtails to have been described by
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, ...
), 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 A woodland () is, in the broad sense, land covered with trees, or in a narrow sense, synonymous with wood (or in the U.S., the '' plurale tantum'' woods), a low-density forest forming open habitats with plenty of sunlight and limited shade (see ...
) and open (
meadow A meadow ( ) is an open habitat, or field, vegetated by grasses, herbs, and other non- woody plants. Trees or shrubs may sparsely populate meadows, as long as these areas maintain an open character. Meadows may be naturally occurring or arti ...
s,
cereal crop A cereal is any grass cultivated for the edible components of its grain (botanically, a type of fruit called a caryopsis), composed of the endosperm, germ, and bran. Cereal grain crops are grown in greater quantities and provide more food en ...
s) environments, most
soil Soil, also commonly referred to as earth or dirt, is a mixture of organic matter, minerals, gases, liquids, and organisms that together support life. Some scientific definitions distinguish ''dirt'' from ''soil'' by restricting the former ...
-dwelling species are not specialized and can be found everywhere, but most epigeal and
litter Litter consists of waste products that have been discarded incorrectly, without consent, at an unsuitable location. Litter can also be used as a verb; to litter means to drop and leave objects, often man-made, such as aluminum cans, paper cups ...
-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 generalist species is able to thrive in a wide variety of environmental conditions and can make use of a variety of different resources (for example, a heterotroph with a varied diet). A specialist species can thrive only in a narrow range of env ...
, a phenomenon the measure of which has been called colonisation credit.


Relationship with humans

Springtails are well known as
pests PESTS was an anonymous American activist group formed in 1986 to critique racism, tokenism, and exclusion in the art world. PESTS produced newsletters, posters, and other print material highlighting examples of discrimination in gallery represent ...
of some agricultural crops. ''
Sminthurus viridis ''Sminthurus viridis'' is a member of the Collembola, the springtails, an order in the subphylum Hexapoda. The species is known by common names such as clover springtail, lucerne flea, or lucerne earth flea. Common names such as lucerne flea are ...
'', the lucerne flea, has been shown to cause severe damage to agricultural crops, and is considered as a pest in Australia. Onychiuridae are also known to feed on tubers and to damage them to some extent. However, by their capacity to carry spores of
mycorrhizal fungi   A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the pla ...
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 Pathogenic fungi are fungi that cause disease in humans or other organisms. Approximately 300 fungi are known to be pathogenic to humans. Markedly more fungi are known to be pathogenic to plant life than those of the animal kingdom. The study of fu ...
through their active consumption of
mycelia Mycelium (plural mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Fungal colonies composed of mycelium are found in and on soil and many other substrates. A typical single spore germinates in ...
and
spores In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...
of damping-off and
pathogenic In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a ger ...
fungi. It has been suggested that they could be reared to be used for the control of
pathogenic fungi Pathogenic fungi are fungi that cause disease in humans or other organisms. Approximately 300 fungi are known to be pathogenic to humans. Markedly more fungi are known to be pathogenic to plant life than those of the animal kingdom. The study of ...
in greenhouses and other indoor cultures. Various sources and publications have suggested that some springtails may
parasitize Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson ha ...
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
delusional parasitosis Delusional parasitosis (DP) is a mental disorder in which individuals have a persistent belief that they are infested with living or nonliving pathogens such as parasites, insects, or bugs, when no such infestation is present. They usually report ...
, 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 Pareidolia (; ) is the tendency for perception to impose a meaningful interpretation on a nebulous stimulus, usually visual, so that one sees an object, pattern, or meaning where there is none. Common examples are perceived images of animals, ...
; 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, 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. In 1952,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
accused the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country Continental United States, primarily located in North America. It consists of 50 U.S. state, states, a Washington, D.C., ...
military of spreading bacteria-laden insects and other objects during the
Korean War {{Infobox military conflict , conflict = Korean War , partof = the Cold War and the Korean conflict , image = Korean War Montage 2.png , image_size = 300px , caption = Clockwise from top:{ ...
by dropping them from
P-51 The North American Aviation P-51 Mustang is an American long-range, single-seat fighter and fighter-bomber used during World War II and the Korean War, among other conflicts. The Mustang was designed in April 1940 by a team headed by James ...
fighters above rebel villages over
North Korea North Korea, officially the Democratic People's Republic of Korea (DPRK), is a country in East Asia. It constitutes the northern half of the Korean Peninsula and shares borders with China and Russia to the north, at the Yalu (Amnok) and T ...
. 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 Biological warfare, also known as germ warfare, is the use of biological toxins or infectious agents such as bacteria, viruses, insects, and fungi with the intent to kill, harm or incapacitate humans, animals or plants as an act of war. ...
effort. The alleged associated diseases included
anthrax Anthrax is an infection caused by the bacterium '' Bacillus anthracis''. It can occur in four forms: skin, lungs, intestinal, and injection. Symptom onset occurs between one day and more than two months after the infection is contracted. The s ...
,
cholera Cholera is an infection of the small intestine by some strains of the bacterium '' Vibrio cholerae''. Symptoms may range from none, to mild, to severe. The classic symptom is large amounts of watery diarrhea that lasts a few days. Vomiting an ...
,
dysentery Dysentery (UK pronunciation: , US: ), historically known as the bloody flux, is a type of gastroenteritis that results in bloody diarrhea. Other symptoms may include fever, abdominal pain, and a feeling of incomplete defecation. Complications ...
, fowl septicemia, paratyphoid,
plague Plague or The Plague may refer to: Agriculture, fauna, and medicine *Plague (disease), a disease caused by ''Yersinia pestis'' * An epidemic of infectious disease (medical or agricultural) * A pandemic caused by such a disease * A swarm of pes ...
, scrub typhus,
small pox Smallpox was an infectious disease caused by variola virus (often called smallpox virus) which belongs to the genus Orthopoxvirus. The last naturally occurring case was diagnosed in October 1977, and the World Health Organization (WHO) ce ...
, and
typhoid Typhoid fever, also known as typhoid, is a disease caused by ''Salmonella'' serotype Typhi bacteria. Symptoms vary from mild to severe, and usually begin six to 30 days after exposure. Often there is a gradual onset of a high fever over several d ...
. 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 The federal government of the United States (U.S. federal government or U.S. government) is the national government of the United States, a federal republic located primarily in North America, composed of 50 states, a city within a feder ...
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. 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''. Captive springtails are often kept in a terrarium as part of a clean-up crew.


Ecotoxicology laboratory animals

Springtails are currently used in laboratory tests for the early detection of
soil pollution Soil contamination, soil pollution, or land pollution as a part of land degradation is caused by the presence of xenobiotic (human-made) chemicals or other alteration in the natural soil environment. It is typically caused by industrial activit ...
.
Acute Acute may refer to: Science and technology * Acute angle ** Acute triangle ** Acute, a leaf shape in the glossary of leaf morphology * Acute (medicine), a disease that it is of short duration and of recent onset. ** Acute toxicity, the adverse ef ...
and
chronic toxicity Chronic toxicity, the development of adverse effects as a result of long term exposure to a contaminant or other stressor, is an important aspect of aquatic toxicology. Adverse effects associated with chronic toxicity can be directly lethal but are ...
tests have been performed by researchers, mostly using the
parthenogenetic Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and development ...
isotomid ''Folsomia candida''. These tests have been standardized. Details on a ringtest, on the biology and
ecotoxicology Ecotoxicology is the study of the effects of toxic chemicals on biological organisms, especially at the population, community, ecosystem, and biosphere levels. Ecotoxicology is a multidisciplinary field, which integrates toxicology and ecolog ...
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. Care should be taken that different strains of the same species may be conducive to different results. 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 move actively far from pollution spots. 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
earthworm An earthworm is a terrestrial invertebrate that belongs to the phylum Annelida. They exhibit a tube-within-a-tube body plan; they are externally segmented with corresponding internal segmentation; and they usually have setae on all segments. T ...
s, and like many insects and molluscs, Collembola are very sensitive to
herbicide Herbicides (, ), also commonly known as weedkillers, are substances used to control undesired plants, also known as weeds.EPA. February 201Pesticides Industry. Sales and Usage 2006 and 2007: Market Estimates. Summary in press releasMain page f ...
s 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 Genomics is an interdisciplinary field of biology focusing on the structure, function, evolution, mapping, and editing of genomes. A genome is an organism's complete set of DNA, including all of its genes as well as its hierarchical, three-dim ...
model organism for soil toxicology. With
microarray A microarray is a multiplex lab-on-a-chip. Its purpose is to simultaneously detect the expression of thousands of genes from a sample (e.g. from a tissue). It is a two-dimensional array on a solid substrate—usually a glass slide or silicon ...
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.


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. There are negative, positive and neutral responses reported. Neutral responses to experimental warming have also been reported in studies of non-polar regions. 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. 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.


Reproduction

Sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote th ...
occurs through the clustered or scattered deposition of spermatophores by
male Male (symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or ovum, in the process of fertilization. A male organism cannot reproduce sexually without access to ...
adults. Stimulation of
spermatophore A spermatophore or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especially salamanders and arthropods, and transferred in entirety to the female's ovipore during reproduction. Spermatophore ...
deposition by female
pheromones A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
has been demonstrated in '' Sinella curviseta''. Mating behavior can be observed in Symphypleona. Among Symphypleona, males of some
Sminthuridae Sminthuridae is a family of springtails of the order Symphypleona. Sminthurids are commonly referred to as globular springtails. Description Like other Symphypleona, Sminthuridae are globular in shape and have a furcula that allows them to ju ...
use a clasping organ located on their
antenna Antenna ( antennas or antennae) may refer to: Science and engineering * Antenna (radio), also known as an aerial, a transducer designed to transmit or receive electromagnetic (e.g., TV or radio) waves * Antennae Galaxies, the name of two collid ...
. Many collembolan species, mostly those living in deeper soil horizons, are parthenogenetic, which favors
reproduction Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. Reproduction is a fundamental feature of all known life; each individual or ...
to the detriment of
genetic diversity Genetic diversity is the total number of genetic characteristics in the genetic makeup of a species, it ranges widely from the number of species to differences within species and can be attributed to the span of survival for a species. It is dis ...
and thereby to
population Population typically refers to the number of people in a single area, whether it be a city or town, region, country, continent, or the world. Governments typically quantify the size of the resident population within their jurisdiction usi ...
tolerance of
environmental hazards An environmental hazard is a substance, state or event which has the potential to threaten the surrounding natural environment or adversely affect people's health, including pollution and natural disasters such as storms and earthquakes. It can i ...
.
Parthenogenesis Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and developmen ...
(also called thelytoky) is under the control of symbiotic
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
of the genus ''
Wolbachia ''Wolbachia'' is a genus of intracellular bacteria that infects mainly arthropod species, including a high proportion of insects, and also some nematodes. It is one of the most common parasitic microbes, and is possibly the most common reproducti ...
'', which live, reproduce and are carried in female reproductive organs and eggs of Collembola. Feminizing ''
Wolbachia ''Wolbachia'' is a genus of intracellular bacteria that infects mainly arthropod species, including a high proportion of insects, and also some nematodes. It is one of the most common parasitic microbes, and is possibly the most common reproducti ...
'' species are widespread in
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s and
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant- parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a bro ...
s, where they co-evolved with most of their lineages.


See also

*
Hexapoda The subphylum Hexapoda (from Greek for 'six legs') comprises most species of arthropods and includes the insects as well as three much smaller groups of wingless arthropods: Collembola, Protura, and Diplura (all of these were once considere ...


References


External links

*
Checklist of the Collembola of the World

Summary information about the distribution and ecology of Collembola (springtails) in the UK and Ireland





A small lecture from Steve Hopkin


* ttp://collembola.co.za/ Collembola species currently recorded from South Africa {{Authority control Articles containing video clips Extant Early Devonian first appearances Taxa named by John Lubbock, 1st Baron Avebury