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Title: Springtail  
Author: World Heritage Encyclopedia
Language: English
Subject: Diplura, Insect, Isotomiella, Asian house martin, Arthropods
Collection: Arthropods, Articles Containing Video Clips, Collembola, Early Devonian First Appearances
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Temporal range: Early Devonian–Recent
Orchesella cincta
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Hexapoda
Class: Entognatha (but see text)
Subclass: Collembola
Lubbock, 1870
Synonyms [1]
  • Oligentoma
  • Oligoentoma

Springtails (Collembola) form the largest of the three lineages of modern hexapods that are no longer considered insects (the other two are the Protura and Diplura). Although the three orders are sometimes grouped together in a class called Entognatha because they have internal mouthparts, they do not appear to be any more closely related to one another than they all are to insects, which have external mouthparts.

Some DNA sequence studies[2][3][4] suggest that Collembola represent a separate evolutionary line from the other Hexapoda, but others disagree;[5] this seems to be caused by widely divergent patterns of molecular evolution among the arthropods.[6] The adjustments of traditional taxonomic rank for springtails reflects 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 (biology) status.

Collembolans are

  • Checklist of the Collembola of the World
  • Maps of Collembola (Britain and Ireland)
  • Maps from 2006 of UK Collembola, plus Photolibrary
  • General information on Collembola
  • The Biology of the Collembola
  • Tree of Life
  • North American Collembola
  • Springtails, Kansas State University

External links

  1. ^ Cedric Gillott (2005). "Apterygote hexapods". Entomology (3rd ed.).  
  2. ^ Frédéric Delsuc, Matthew J. Phillips & David Penny (2003). "Hexapod origins: monophyletic or paraphyletic?"Comment on ( 
  3. ^ Francesco Nardi, Giacomo Spinsanti, Jeffrey L. Boore, Antonio Carapelli, Romano Dallai & Francesco Frati (2003). "Hexapod origins: monophyletic or paraphyletic?" ( 
  4. ^ Francesco Nardi, Giacomo Spinsanti, Jeffrey L. Boore, Antonio Carapelli, Romano Dallai & Francesco Frati (2003). "Hexapod origins: monophyletic or paraphyletic?"Response to comment on ( 
  5. ^ a b Yan Gao, Yun Bu & Yun-Xia Luan (2008). "Phylogenetic relationships of basal hexapods reconstructed from nearly complete 18S and 28S rRNA gene sequences" ( 
  6. ^ Alexandre Hassanin (2006). "Phylogeny of Arthropoda inferred from mitochondrial sequences: strategies for limiting the misleading effects of multiple changes in pattern and rates of substitution" ( 
  7. ^ Nyle C. Brady & Ray R. Weil (2009). Elements of the Nature and Properties of Soils (3rd Edition). Prentice Hall.  
  8. ^ Torsten Thimm, Andrea Hoffmann†, Heinz Borkott, Jean Charles Munch & Christoph C. Tebbe (1998). (Collembola) is a frequently changeable but selective habitat and a vector for microorganisms"Folsomia candida"The gut of the soil microarthropod ( 
  9. ^ W. Maldwyn Davies (1927). , Lubb."Sminthurus viridis"On the tracheal system of Collembola, with special reference to that of ( 
  10. ^ a b  
  11. ^ "The incredible shrinking springtail".  
  12. ^ Howell V. Daly, John T. Doyen & Alexander H. Purcell (1998). Introduction to insect biology and diversity (2nd ed.).  
  13. ^ "Hexapoda. Insects, springtails, diplurans, and proturans".  
  14. ^ José A. Mari Mutt (1983). "Collembola in amber from the Dominican Republic" ( 
  15. ^ André Nel, Gaėl De Ploëg, Jacqueline Milliet, Jean-Jacques Menier & Alain Waller (2004). "The French ambers: a general conspectus and the Lowermost Eocene amber deposit of Le Quesnoy in the Paris Basin" ( 
  16. ^ David Penney, Andrew McNeil, Andrew, David I. Green, Robert S. Bradley, Robert S., James E. Jepson, Philip J. Withers & Richard F. Preziosi (2012). "Ancient Ephemeroptera-Collembola symbiosis fossilized in amber predicts contemporary phoretic associations" ( 
  17. ^ Kenneth Christiansen & Paul Nascimbene (2006). "Collembola (Arthropoda, Hexapoda) from the mid Cretaceous of Myanmar (Burma)".  
  18. ^ Koehler, PG; Aparicio, ML; Pfiester, M (July 2011). "Springtails". University of Florida IFAS Extension. Retrieved 2014-06-15. 
  19. ^ a b Stephen P. Hopkin (1997). "The biology of the Collembola (springtails): the most abundant insects in the world" ( 
  20. ^ Jean-François Ponge (1991). "Food resources and diets of soil animals in a small area of Scots pine litter" ( 
  21. ^ Jean-François Ponge, Pierre Arpin, Francis Sondag & Ferdinand Delecour (1997). "Soil fauna and site assessment in beech stands of the Belgian Ardennes" ( 
  22. ^ a b c d e Jean-François Ponge (1993). "Biocenoses of Collembola in atlantic temperate grass-woodland ecosystems" ( 
  23. ^ Island Creek Elementary School. "Hypogastrura nivicola"Snow Flea. . Study of Northern Virginia Ecology. Fairfax County Public Schools. Retrieved June 22, 2014. 
  24. ^ Peter Shaw, Claire Ozanne, Martin Speight & Imogen Palmer (2007). "Edge effects and arboreal Collembola in coniferous plantations" ( 
  25. ^ Jürg Zettel, Ursula Zettel & Beatrice Egger (2000). "Jumping technique and climbing behaviour of the collembolan Ceratophysella sigillata (Collembola: Hypogastruridae)" ( 
  26. ^ a b Wim A. M. Didden (1987). (Collembola) to loose and compact soil: methods and first results"Onychiurus fimatus"Reactions of . Pedobiologia 30 (2): 93–100. 
  27. ^ John Bowden, I. H. Haines & D. Mercer (1976). "Climbing Collembola". Pedobiologia 16: 298–312. 
  28. ^ Julia Nickerl, Ralf Helbig, Hans-Jürgen Schulz, Carsten Werner, & Christoph Neinhuis (2013). "Diversity and potential correlations to the function of Collembola cuticle structures".  
  29. ^ Martin Holmstrup & Mark Bayley (2013). "Protaphorura tricampata, a euedaphic and highly permeable springtail that can sustain activity by osmoregulation during extreme drought". Journal of Insect Physiology 59 (11): 1104–1110.  
  30. ^ Herman A. Verhoef (1984). "Releaser and primer pheromones in Collembola". Journal of Insect Physiology 30 (8): 665–670.  
  31. ^ Joshua B. Benoit, Michael A. Elnitsky, Glen G. Schulte, Richard E. Lee Jr & David L. Denlinger (2009). "Antarctic Collembolans use chemical signals to promote aggregation and egg laying" ( 
  32. ^ Andreas Prinzing, Cyrille A. D'Haese, Sandrine Pavoine & Jean-François Ponge (2014). (Collembola)"Willemia"Species living in harsh environments have low clade rank and are localized on former Laurasian continents: a case study of ( 
  33. ^ Herman A. Verhoef (1981). "Water balance in Collembola and its relation to habitat selection: water content, haemolymph osmotic pressure and transpiration during an instar". Journal of Insect Physiology 27 (11): 755–760.  
  34. ^ Hans Petter Leinaas (1983). "Synchronized moulting controlled by communication in group-living Collembola".  
  35. ^ Jane M. Wilson (1982). "A review of world Troglopedetini (Insecta, Collembola, Paronellidae), including an identification table and descriptions of new species" ( 
  36. ^ José G. Palacios-Vargas & Jane Wilson (1990). , a new genus and species of cave collembolan from Madagascar with notes on its ecology"Troglobius coprophagus" ( 
  37. ^ Rafael Jordana, Enrique Baquero, Sofía Reboleira & Alberto Sendra (2012). "Reviews of the genera Schaefferia Absolon, 1900, Deuteraphorura Absolon, 1901, Plutomurus Yosii, 1956 and the Anurida Laboulbène, 1865 species group without eyes, with the description of four new species of cave springtails (Collembola) from Krubera-Voronya cave, Arabika Massif, Abkhazia" ( 
  38. ^ Sandrine Salmon, Jean-François Ponge & Nico Van Straalen (2002). "Ionic identity of pore water influences pH preference in Collembola" ( 
  39. ^ Gladys Loranger, Ipsa Bandyopadhyaya, Barbara Razaka & Jean-François Ponge (2001). "Does soil acidity explain altitudinal sequences in collembolan communities?" ( 
  40. ^ Jack H. Faber & E.N.G. Joosse (1993). "Vertical distribution of Collembola in a Pinus nigra organic soil". Pedobiologia 37 (6): 336–350. 
  41. ^ Vassilis Detsis (2000). "Vertical distribution of Collembola in deciduous forests under Mediterranean climatic conditions" ( 
  42. ^ "Isotomurus palustris (Muller, 1776)". Retrieved August 3, 2014. 
  43. ^ Sylvain Pichard (1973). "Contribution à l'étude de la biologie de Podura aquatica (Linné) Collembole". Bulletin Biologique de la France et de la Belgique (in French) 107 (4): 291–299. 
  44. ^ Jean-François Ponge, Servane Gillet, Florence Dubs, Eric Fédoroff, Lucienne Haese, José Paulo Sousa & Patrick Lavelle (2003). "Collembolan communities as bioindicators of land use diversification" ( 
  45. ^ Jean-François Ponge, Florence Dubs, Servane Gillet, Jose Paulo Sousa & Patrick Lavelle (2006). "Decreased biodiversity in soil springtail communities: the importance of dispersal and landuse history in heterogeneous landscapes" ( 
  46. ^ Sara Cristofoli & Grégory Mahy (2010). "Colonisation credit in recent wet heathland butterfly communities". Insect Conservation and Diversity 3 (2): 83–91.  
  47. ^ Charlène Heiniger, Sébastien Barot, Jean-François Ponge, Sandrine Salmon, Léo Botton-Divet, David Carmignac & Florence Dubs (2014). "Effect of habitat spatiotemporal structure on collembolan diversity" ( 
  48. ^ Michael W. Shaw & G. M. Haughs (1983). (L.)"Sminthurus viridis"Damage to potato foliage by .  
  49. ^ Alan L. Bishop, Anne M. Harris, Harry J. McKenzie (2001). (L.) (Collembola: Sminthuridae), in irrigated lucerne in the Hunter dairying region of New South Wales"Sminthurus viridis"Distribution and ecology of the lucerne flea, .  
  50. ^ "Sminthurus viridis"Lucerne Flea .  
  51. ^ A. N. Baker & R. A. Dunning (1975). "Association of populations of onychiurid Collembola with damage to sugar-beet seedlings".  
  52. ^ John N. Klironomos & Peter Moutoglis (1999). "Folsomia candida"Colonization of nonmycorrhizal plants by mycorrhizal neighbours as influenced by the collembolan, . Biology and Fertility of Soils 29 (3): 277–281.  
  53. ^ Maria Agnese Sabatini & Gloria Innocenti (2001). "Effects of Collembola on plant-pathogenic fungus interactions in simple experimental systems". Biology and Fertility of Soils 33 (1): 62–66.  
  54. ^ Hiroyoshi Shiraishi, Yoshinari Enami & Seigo Okano (2003). "Folsomia hidakana (Collembola) prevents damping-off disease in cabbage and Chinese cabbage by Rhizoctonia solani". Pedobiologia 47 (1): 33–38.  
  55. ^ Jean-François Ponge & Marie-José Charpentié (1981). (Packard), Collembole mycophage"Pseudosinella alba"Étude des relations microflore-microfaune: expériences sur ( 
  56. ^ Robert T. Lartey, E. A. Curl, Curt M. Peterson & James D. Harper (1989). (Collembola: Onychiuridae)"Onychiurus encarpatus (Collembola: Isotomidae) and Proisotoma minuta"Mycophagous grazing and food preference of . Environmental Entomology 18 (2): 334–337. 
  57. ^ a b Frans Janssens & Kenneth A. Christiansen (November 22, 2007). "Synanthropic Collembola, springtails in association with man". Checklist of the Collembola. Retrieved December 7, 2014. 
  58. ^ May Berenbaum (2005). "Face Time" ( 
  59. ^ Kenneth Christiansen & Ernest C. Bernard (2008). """Critique of the article "Collembola (Springtails) (Arthropoda: Hexapoda: Entognatha) found in scrapings from individuals diagnosed with delusory parasitosis.  
  60. ^ Michelle T. Fountain & Steve P. Hopkin (2001). (Insecta: Collembola) in a metal exposure test"Folsomia candida"Continuous monitoring of .  
  61. ^ ISO 11267 (2014). ) by soil contaminants"Folsomia candida"Soil quality. Inhibition of reproduction of Collembola (. Geneva:  
  62. ^ Christine Lors, Maite Martínez Aldaya, Sandrine Salmon & Jean-François Ponge (2006). "Use of an avoidance test for the assessment of microbial degradation of PAHs" ( 
  63. ^ ISO 17512-2 (2011). )"Folsomia candida"Soil quality. Avoidance test for determining the quality of soils and effects of chemicals on behaviour. Part 2: Test with collembolans (. Geneva:  
  64. ^ Matthieu Chauvat & Jean-François Ponge (2002). "Colonization of heavy metal-polluted soils by collembola: preliminary experiments in compartmented boxes" ( 
  65. ^ Jean-François Ponge, Ipsa Bandyopadhyaya & Valérie Marchetti (2002). "Interaction between humus form and herbicide toxicity to Collembola (Hexapoda)" ( 
  66. ^ Benjamin Nota, Martijn J.T.N. Timmermans, Oscar Franken, Kora Montagne-Wajer, Janine Mariën, Muriel E. De Boer, Tjalf E. De Boer, Bauke Ylstra, Nico M. Van Straalen & Dick Roelofs (2008). "Gene expression analysis of Collembola in cadmium containing soil".  
  67. ^ Benjamin Nota, Mirte Bosse, Bauke Ylstra, Nico M. Van Straalen & Dick Roelofs (2009). exposed to phenanthrene"Folsomia candida"Transcriptomics reveals extensive inducible biotransformation in the soil-dwelling invertebrate ( 
  68. ^ Elizabeth S. Waldorf (1974). "Sinella curviseta"Sex pheromone in the springtail .  
  69. ^ Marek Wojciech Kozlowski & Shi Aoxiang (2006). (Collembola : Bourletiellidae)"Deuterosminthurus bicinctus"Ritual behaviors associated with spermatophore transfer in .  
  70. ^ Alice B. Czarnetzki & Christoph C. Tebbe (2004). in Collembola"Wolbachia"Detection and phylogenetic analysis of .  
  71. ^ John H. Werren, Wan Zhang & Li Rong Guo (1995). : reproductive parasites of arthropods"Wolbachia"Evolution and phylogeny of ( 
  72. ^ Katelyn Fenn & Mark Blaxter (2004). obligate mutualist symbionts?"Wolbachia"Are filarial nematode .  


See also

. lineages where they co-evolved with most of their [72],nematodes and [71]arthropods species are widespread in Wolbachia Feminizing [70]


exposed to environmental toxicants allow fast and sensitive detection of pollution, and additionally clarifies molecular mechanisms causing toxicology. Folsomia candida technology the expression of thousands of genes can be measured in parallel. The gene expression profiles of microarray With [67][66] Springtails are currently used in laboratory tests for the early detection of

Ecotoxicology laboratory animals

Various sources and publications have suggested that some springtails may delusory parasitosis, a psychological rather than entomological problem. Researchers themselves may be subject to psychological phenomena. For example, a publication in 2004 claiming that springtails had been found in skin samples was later determined to be a case of pareidolia; that is, no springtail specimens were actually recovered, but the researchers had digitally enhanced photos of sample debris to create images resembling small arthropod heads, which then were claimed to be springtail remnants.[57][58][59] 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.[19]

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,[48] and is considered as a pest in Australia.[49][50] Also Onychiuridae are known to feed on tubers and to damage them to some extent.[51] 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.[52] They also contribute to controlling plant fungal diseases through their active consumption of mycelia and spores of damping-off and pathogenic fungi.[53][54] It has been suggested that they could be reared to be used for the control of pathogenic fungi in greenhouses and other indoor cultures.[55][56]

Tomocerus sp. from Germany

Relationship with humans

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.[22][44] As a consequence of dispersal limitation, landuse change, when too rapid, may cause the local disappearance of slow-moving, specialist species,[45] a phenomenon the measure of which was recently called colonisation credit.[46][47]

The horizontal distribution of springtail species is affected by environmental factors which act at the landscape scale, such as soil acidity, moisture and light.[22] Requirements for pH can be reconstructed experimentally.[38] Altitudinal changes in species distribution can be at least partly explained by increased acidity at higher elevation.[39] Moisture requirements, among other ecological and behavioural factors, explain why some species cannot live aboveground,[40] or retreat in the soil during dry seasons,[41] but also why some epigeal springtails are always found in the vicinity of ponds and lakes, such as the hygrophilous Isotomurus palustris.[42] 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.[43]

Anurida maritima on water

As a group, springtails are highly sensitive to desiccation, because of their tegumentary respiration.[28] although some species with thin, permeable cuticles have been shown to resist severe drought by regulating the osmotic pressure of their body fluid.[29] The gregarious behaviour of Collembola, mostly driven by the attractive power of pheromones excreted by adults,[30] 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.[31] Sensitivity to drought varies from species to species[32] and increases during ecdysis.[33] Given that springtails are moulting repeatedly during their entire life (an ancestral character in Hexapoda) they spend much time in concealed micro-sites where they can find protection against desiccation and predation during ecdysis, an advantage reinforced by synchronized moulting.[34] The high humidity environment of many caves also favours springtails and there are numerous cave adapted species,[35][36] including one, Plutomurus ortobalaganensis living 1,980 metres (6,500 ft) down the Krubera Cave.[37]

Dicyrtomina sp. on leaf

The main ecological factor driving the local distribution of species is the vertical stratification of the environment: 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.[22] This is a complex factor embracing both nutritional and physiological requirements, together with behavioural trends[26] and probable species interactions. Some species have been shown to exhibit negative[27] or positive[26] gravitropism, which adds a behavioural dimension to this still poorly understood vertical segregation.

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.[24][25] These tend to be the larger (>2 mm) species, mainly in the genera Entomobrya and Orchesella, though the densities on a per square meter basis are typically 1–2 orders of magnitude lower than soil populations of the same species. In temperate regions, a few species (e.g. Anurophorus spp., Entomobrya albocincta, Xenylla xavieri, Hypogastrura arborea) are almost exclusively arboreal.[22] In tropical regions a single square meter of canopy habitat can support many species of Collembola.[10]

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,[21] essentially everywhere on Earth where soil and related habitats (moss cushions, fallen wood, grass tufts, ant and termite nests) occur.[22] Only nematodes, crustaceans, and mites are likely to have global populations of similar magnitude, and each of those groups except mites is more inclusive: though taxonomic rank cannot be used for absolute comparisons, it is notable that nematodes are a phylum and crustaceans a subphylum. Most springtails are small and difficult to see by casual observation, but one springtail, the so-called snow flea (Hypogastrura nivicola), is readily observed on warm winter days when it is active and its dark color contrasts sharply with a background of snow.[23]

"Snow flea"

[20] Springtails are



There are about 3,600 different species.[18]

Fossil collembola are rare. Instead, most are found in amber.[14] 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,[15] Miocene of Central America,[16] and the mid-Cretaceous of Burma and Canada.[17] They display some unusual characteristics: first, all but one of the fossils from the Cretaceous belong to extinct genera, whereas none of the specimens from the Eocene or the Miocene are of extinct genera; second, the species from Burma are more similar to the modern fauna of Canada than are the Canadian Cretaceous specimens.

Springtails are attested to since the Early Devonian.[12] The fossil from , Rhyniella praecursor, is the oldest terrestrial arthropod, and was found in the famous Rhynie chert of Scotland. Given its morphology resembles extant species quite closely, the radiation of the Hexapoda can be situated in the Silurian, or more.[13]

The Neelipleona were originally seen as a particular advanced lineage of Symphypleona, based on the shared global body shape. But the global body of Neelipleona is realised in a completely different way than in Symphypleona. Subsequently, the Neelipleona were considered as being derived from the Entomobryomorpha. But analysis of 18S and 28S rRNA sequence data suggests that they form the most ancient lineage of springtails, which would explain their peculiar apomorphies.[5]

Traditionally, the springtails were divided into the orders Arthropleona, Symphypleona 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 their 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.

Allacma fusca (Symphypleona) on rotting wood

Systematics and evolution

[11] Springtails also possess the ability to reduce their body size by as much as 30% through subsequent

Most species have an abdominal, tail-like appendage, the furcula, that is folded beneath the body to be used for jumping when the animal is threatened. It is held under tension by a small structure called the retinaculum and when released, snaps against the substrate, flinging the springtail into the air. All of this takes place in as little as 18 milliseconds.[10]

Members of 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 eversible vesicles, projecting ventrally from the first abdominal segment. The Poduromorpha and Entomobryomorpha have an elongated body, while the Symphypleona have a globular body. Collembola lack a tracheal respiration system, which forces them to respire through a porous cuticle, to the notable exception of Sminthuridae which exhibit a rudimentary, although fully functional, tracheal system.[9]

Isotoma with visible furcula



  • Description 1
  • Systematics and evolution 2
  • Ecology 3
    • Distribution 3.1
    • Relationship with humans 3.2
    • Ecotoxicology laboratory animals 3.3
  • Reproduction 4
  • See also 5
  • References 6
  • External links 7

meaning piston or peg. embolon meaning glue and colle The word "collembola" is from the Greek [8] and the control of soil microbial communities.[7]

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