Springtails (Collembola) form the largest of the three lineages of modern hexapods that are no longer considered insects (the other two are the Protura and Diplura). The three orders are sometimes grouped together in a class called Entognatha because they have internal mouthparts, but they do not appear to be more closely related to one another than to insects, which have external mouthparts.
DNA sequence studies suggest that Collembola are a separate evolutionary line from the other Hexapoda. Consequently their taxonomic rank has changed according to need: 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.
Members of Collembola are normally less than 6 mm
long, have six or fewer abdominal segments
and possess a tubular appendage
or ventral tube) with eversible vesicles, projecting ventrally from the first abdominal segment. Most species have an abdominal, tail-like appendage, the furcula
, that is folded beneath the body to be used for jumping when the animal is threatened. It is held under tension by a small structure called the retinaculum
and when released, snaps against the substrate, flinging the springtail into the air.
The Poduromorpha and Entomobryomorpha have an elongated body, while the Symphypleona and Neelipleona have a globular body.
Systematics and evolution
Traditionally, the springtails were divided into the orders Arthropleona
and occasionally also Neelipleona. The Arthropleona were divided into two superfamilies
, the Entomobryoidea and the Poduroidea. But actually, these two and the Symphypleona form three lineages, each of which is equally distant from the other two. Thus, the "Arthropleona" are abolished in modern classifications, and its superfamilies are raised in rank accordingly, being now the Entomobryomorpha
and the Poduromorpha
. Technically, the Arthropleona are thus a partial junior synonym
of the Collembola. The term "Neopleona" is essentially synonymous with Symphypleona + Neelipleona.
The Neelipleona, which are monotypic, were originally seen as particular advanced lineage of Symphypleona, based on the shared global body shape. But the global body of Neelipleona is realised in a completely different way than in Symphypleona. Currently, the Neelipleona are considered as being derived from the Entomobryomorpha.
Springtails are attested to since the Early Devonian. The fossil from 400 million years ago, Rhyniella praecursor, is the 'oldest' terrestrial arthropod found in the famous Rhynie chert of Scotland. Given its morphology resembles current species quite well, the radiation of the Hexapoda can be situated in the Silurian, 420 million years ago or more.
Springtails are cryptozoa
frequently found in leaf litter
and other decaying material, where they are primarily detritivores
and microbivores, and one of the main biological agents responsible for the control and the dissemination of microorganisms
. In sheer numbers, they are reputed to be one of the most abundant of all macroscopic animals, with estimates of 100,000 individuals per cubic meter of topsoil, essentially everywhere on Earth where soil and related habitats (moss
cushions, fallen wood
nests) occur; only nematodes
, and mites
are likely to have global populations of similar magnitude, and each of those groups except mites is more inclusive: though taxonomic rank cannot be used for absolute comparisons, it is notable that nematodes are a phylum
and crustaceans a subphylum
. Most springtails are small and difficult to see by casual observation, but one springtail, the so-called snow flea
(such as Hypogastrura nivicola
), is readily observed on warm winter days when it is active and its dark color contrasts sharply with a background of snow.
In addition, a few species routinely climb trees and form a dominant component of canopy faunas, where they may be collected by beating or insecticide fogging. These tend to be the larger (>2mm) species, mainly in the genera Entomobrya, Orchesella and Lepidocyrtus, though the densities on a per square metre basis are typically 1-2 orders of magnitude lower than soil populations of the same species. A very few species (e.g. Anurophorus spp., Entomobrya albocincta) are almost exclusively arboreal.
The main ecological factor driving locally the distribution of species is the vertical stratification of the environment: in woodlands a continuous change in species assemblages can be observed from tree canopies to ground vegetation then to plant litter down to deeper soil horizons. This a complex factor embracing both nutritional and physiological requirements, together with probable species interactions. Some species have been shown to exhibit negative or positive gravitropism, which adds a behavioural dimension to this still poorly understood vertical segregation.
As a group, springtails are highly sensitive to desiccation, because of their tegumentary respiration: they lack trachea (exceptions exist, such as in Sminthuridae), which forces them to respire through a porous cuticle. The gregarious behaviour of Collembola, mostly driven by the attractive power of pheromones excreted by adults, gives more chance to every juvenile or adult individual to find suitable, better protected places, where desiccation could be avoided and reproduction rate could be kept at an optimum. Sensitivity to dryness varies from species to species and increases during ecdysis. Given that springtails are moulting repeatedly during their entire life (an ancestral character in Hexapoda) they spend much time in concealed micro-sites where they can find protection against desiccation and predation.
The horizontal distribution of springtail species is affected by environmental factors which act at the landscape scale, such as soil acidity, moisture and light. Requirements for pH can be reconstructed experimentally. Altitudinal changes in species distribution can be at least partly explained by increased acidity at higher elevation. Moisture requirements explain why some species cannot live aboveground (vertical stratification), but also why some epigeal springtails are always found in the vicinity of ponds and lakes, such as the hygrophilous Isotomurus palustris. Adaptive features, such as the presence of a fan-like wettable mucro, allow some species to move at the surface of water (Sminthurides aquaticus, Sminthurides malmgreni). Podura aquatica, a unique representative of the family Poduridae (and one of the first springtails to have been described by Linnaeus), spends its entire life at the surface of water, its wettable eggs dropping in water until the non-wettable first instar hatches then surfaces.
In a variegated landscape, made of a patchwork of closed (woodland) and open (meadows, cereal crops) environments, most soil-dwelling species are not specialized and can be found everywhere, but most epigeal and litter-dwelling species are attracted to a particular environment, either forested of not. As a consequence of dispersal limitation, landuse change, when too rapid, may cause the local disappearance of slow-moving, specialized species.
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
. Mating behaviour
can be observed in Symphypleona
. Among Symphypleona, males of Sminthurididae use a clasping organ located on their antenna
. Many collembolan species, mostly those living in deeper soil horizons, are parthenogenetic
, which favours reproduction
to the detriment of genetic diversity
and thereby to population
tolerance of environmental hazards
(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. Feminizing Wolbachia
species are widespread in arthropods
where they co-evolved with most of their lineages
Relationship with humans
Various sources and publications have suggested that some springtails may parasitize humans, but this is entirely inconsistent with their biology, and no such phenomenon has ever been scientifically confirmed, though it has been documented that the scales or hairs from collembolans can cause irritation when rubbed onto the skin. They can sometimes be abundant indoors in damp places such as bathrooms and basements, and under such circumstances may be found on one's person, but this is only accidental.
Claims of persistent human skin infection by springtails may indicate a neurological problem, or else delusory parasitosis, a psychological rather than entomological problem. Researchers themselves may be subject to psychological phenomena. For example, a publication in 2004 claiming that springtails had been found in skin samples was later determined to be a case of pareidolia; that is, no springtail specimens were actually recovered, but the researchers had digitally enhanced photos of sample debris to create images resembling small arthropod heads, which they then claimed to be springtail remnants. However, Hopkin reports one instance of an entomologist aspirating an Isotoma species and in the process accidentally inhaling some of their eggs, which hatched in his nasal cavity and made him quite ill until they were flushed out.
More serious and well-documented are the records of springtails as pests
for some agricultural crops. Sminthurus viridis
, the 'lucerne flea', has been shown to cause severe damage to agricultural crops and is considered as a pest in Australia
- (2005): Face Time. American Entomologist 51: 68-69.
- (2001): Distribution and ecology of the lucerne flea, Sminthurus viridis (L.) (Collembola: Sminthuridae), in irrigated lucerne in the Hunter dairying region of New South Wales. Australian Journal of Entomology 40(1):49-55. HTML abstract
- (1976): Climbing Collembola. Pedobiologia 16: 298–312.
- (2004): Detection and phylogenetic analysis of Wolbachia in Collembola. Environmental Microbiology 6: 35-44. HTML abstract
- (2003): Comment on "Hexapod Origins: Monophyletic or Paraphyletic?" Science 301(5639): 1482. PDF fulltext
- (1987): Reactions of Onychiurus fimatus (Collembola) to loose and compact soil methods and first results. Pedobiologia 30(2): 93-100. HTML abstract
- (2004): Are filarial nematode Wolbachia obligate mutualist symbionts? Trends Ecol. Evol. 19(4): 163-166. (HTML abstract)
- (2008): Mikko's Phylogeny Archive - Collembola Version of 2008-MAR-11. Retrieved 2008-JUL-11.
- (2006): Phylogeny of Arthropoda inferred from mitochondrial sequences: Strategies for limiting the misleading effects of multiple changes in pattern and rates of substitution. Mol. Phylogenet. Evol. 38(1): 100-116. (HTML abstract)
- (1997): The Biology of the Collembola (Springtails): The Most Abundant Insects in the World. Natural History Museum, London. PDF fulltext
- (2007): Checklist of the Collembola - Synanthropic Collembola, Springtails in Association with Man Version of 2007-NOV-22. Retrieved 2008-JUL-11.
- (2006): Ritual behaviors associated with spermatophore transfer in Deuterosminthurus bicinctus (Collembola : Bourletiellidae). Journal of Ethology 24(2): 103-110. (HTML abstract)
- (2001): Does soil acidity explain altitudinal sequences in collembolan communities? Soil Biology and Biochemistry 33(3): 381–393. (HTML abstract)
- } (2003a): Hexapod Origins: Monophyletic or Paraphyletic? Science 299(5614): 1887-1889. Supporting Online Material
- } (2003b): Response to Comment on "Hexapod Origins: Monophyletic or Paraphyletic?" Science 301(5639): 1482. PDF fulltext
- (1991): Food resources and diets of soil animals in a small area of Scots pine litter. Geoderma 49(1-2): 33–62. (HTML abstract)
- (1993): Biocenoses of Collembola in atlantic temperate grass-woodland ecosystems. Pedobiologia 37(4): 223-244.
- (2006): Decreased biodiversity in soil springtail communities: the importance of dispersal and landuse history in heterogeneous landscapes. Soil Biology & Biochemistry 38: 1158-1161. (HTML abstract)
- (1983): Damage to potato foliage by Sminthurus viridis (L.). Plant Pathology 32(4): 465-466.
- (1997): The development of a bioindicator system for soil acidity based on arthropod pH preferences.Journal of Applied Ecology 34(1): 217–232.
- (1984): Releaser and primer pheromones in Collembola. Journal of Insect Physiology 30(8): 665-670.
- (1974): Sex pheromone in the springtail Sinella curviseta. Environmental Entomology 3: 916–918.
- (1995): Evolution and phylogeny of Wolbachia: reproductive parasites of arthropods. Proceedings of the Royal Society of London, Series B, Biological Sciences 261(1360): 55–63. PMID 7644549 HTML abstract and first page image
See complete list
- Bengtsson Göran
- Berg Matty
- Chagnon Madeleine
- Chauvat Matthieu
- Christian Erhard
- Christiansen Kenneth A.
- Cortet Jérôme
- Crouau Yves
- Deharveng Louis
- Dunger Wolfram
- Eisenbeis Gerhard
- Filser Juliane
- Fountain Michelle
- Frati Francesco
- Fjellberg Arne
- Frampton Geoffrey
- Gers Charles
- Greenslade Penelope
- Gruia Magdalena
- Hågvar Sigmund
- Hedlund Katarina
- Holmstrup Martin
- Huhta Veikko
- Janssens Frans
- Jordana Rafael
- Kampichler Christian
- Krogh Paul Henning
- Lee Byung Hoon
- Mari Mutt José A.
- Mateos Eduardo
- Najt Judith
- Palacios-Vargas José G.
- Pomorski Romuald Jacek
- Ponge Jean-François
- Potapov Mikhail
- Rusek Josef
- Salmon Sandrine
- Shaw Peter
- Simon Benito José Carlos
- Snider Richard J.
- Sousa Jorge Paulo
- Takeda Hiroshi
- Thibaud Jean-Marc
- Van Straalen Nico M.
- Verhoef Herman A.
- Weiner Wanda
- Wolters Volkmar
- Zettel Jürg