World Library  
Flag as Inappropriate
Email this Article


Article Id: WHEBN0008088857
Reproduction Date:

Title: Conus  
Author: World Heritage Encyclopedia
Language: English
Subject: 412th Engineer Command (United States), M16 rifle, United States Army Coast Artillery Corps, Strategic Air Command, Conus
Publisher: World Heritage Encyclopedia


Cone snails
Temporal range: Eocene–Recent
Geography cone, Conus geographus
Conus species eating a small fish, in Guam
Scientific classification
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
(unranked): clade Caenogastropoda
clade Hypsogastropoda
clade Neogastropoda
Superfamily: Conoidea
Family: Conidae
Subfamily: Coninae
Genus: Conus
Linnaeus, 1758[1]
Type species
Conus marmoreus
Linnaeus, 1758
Over 600 species

Africonus Petuch, 1975
Afroconus Petuch, 1975
Ammirales Schepman, 1913
Asperi Schepman, 1913
Asprella Schaufuss, 1869
Chelyconus Mörch, 1842
Cleobula Iredale, 1930
Conasprella Thiele, 1929
Coronaxis Swainson, 1840
Cucullus Röding, 1798
Cylinder Montfort, 1810
Cylindrus Deshayes, 1824
Darioconus Iredale, 1930
Dauciconus Cotton, 1945
Dendroconus Swainson, 1840
Embrikena Iredale, 1937
Endemoconus Iredale, 1931
Eugeniconus da Motta, 1991
Floraconus Iredale, 1930
Gastridium Mödeer, 1793
Hermes Montfort, 1810
Kermasprella Powell, 1958
Lautoconus Monterosato, 1923
Leporiconus Iredale, 1930
Leptoconus Swainson, 1840
Lilliconus Raybaudi Massilia, 1994
Lithoconus Mörch, 1852
Magelliconus da Motta, 1991
Mamiconus Cotton & Godfrey, 1932
Phasmoconus Mörch, 1852
Pionoconus Mörch, 1852
Profundiconus Kuroda, 1956
Purpuriconus da Motta, 1991
Rhizoconus Mörch, 1852
Stephanoconus Mörch, 1852
Strioconus Thiele, 1929
Taranteconus Azuma, 1972
Textilia Swainson, 1840
Thoraconus da Motta, 1991
Tuliparia Swainson, 1840
Turriconus Shikama & Habe, 1968
Virgiconus Cotton, 1945
Virroconus Iredale, 1930

Conus is a large genus of small to large predatory sea snails, marine gastropod molluscs, with the common names of cone snails, cone shells or cones. This genus is placed in the subfamily Coninae within the family Conidae. Geologically speaking, the genus is known from the Eocene to the Recent (Holocene) periods.[3] Conus species have shells that are shaped more or less like geometric cones. Many species have colorful patterning on the shell surface. Conus snails are mostly tropical in distribution.

Because all Conus snails are venomous and capable of "stinging" humans, live ones should be handled with great care or preferably not at all. The species most dangerous to humans are the larger ones which prey on small bottom-dwelling fish; the smaller species mostly hunt and eat marine worms. Cone snails use a hypodermic-like modified radula tooth and a venom gland to attack and paralyze their prey before engulfing it. The tooth is sometimes likened to a dart or a harpoon. It is barbed and can be extended some distance out from the mouth of the snail, at the end of the proboscis.

Cone snail venoms are mainly peptides. The venoms contain many different toxins that vary in their effects; some are extremely toxic. The sting of small cones is no worse than a bee sting, but the sting of a few of the larger species of tropical cone snails can be serious, occasionally even fatal to human beings. Cone snail venom is showing great promise as a source of new, medically important substances.[4][5]

Distribution and habitat

There are over 600 different species of cone snails.[2] This family is typically found in warm and tropical seas and oceans worldwide, and reaches its greatest diversity in the Western Indo-Pacific Region. However, some species of Conus are adapted to temperate environments, such as the Cape coast of South Africa,[6][7] the Mediterranean,[8] or the cool waters of southern California (Conus californicus),[9] and are endemic to these areas.

This genus is found in all tropical and subtropical seas from tidal waters to deeper areas, living on sand or among rocks or coral reefs. When living on sand, these snails will bury themselves with only the siphon protruding from the surface. Many tropical cone snails live in or near coral reefs. Some species are found under rocks in the lower intertidal and shallow subtidal zones.

Shell description

This genus shows a large variety of colors and patterns, and local varieties and color forms of the same species often occur. This has led to the creation of a large number of known synonyms and probable synonyms, making it difficult to give an exact taxonomic assignment for many snails in this genus. As of 2009, more than 3,200 different species names have been assigned to snails in the genus Conus, with an average of 16 new species names introduced each year.[10]

The shells of Conus species vary in size. The shells are shaped more or less like the geometric shape known as a cone, as one might expect from the popular and scientific name. The shell is many-whorled and in the form of an inverted cone, the anterior end being the narrow end. The protruding parts of the top of the whorls that form the spire are more or less in the shape of another, much more flattened, cone. The aperture is elongated and narrow. The horny operculum is very small. The outer lip is simple, thin, and sharp, is without a callus, and has a notched tip at the upper part. The columella is straight.

The larger species of cone snails can grow up to 23 cm (9.1 in) in length. The shells of cone snails are often brightly colored and have interesting patterns, although in some species the color patterns may be partially or completely hidden under an opaque layer of periostracum. In other species the topmost shell layer is thin periostracum, a transparent yellowish or brownish membrane.

Life habits

Cone snails are carnivorous, and predatory. They hunt and eat prey such as marine worms, small fish, molluscs, and even other cone snails. Because cone snails are slow-moving, they use a venomous harpoon (called a toxoglossan radula) to capture faster-moving prey, such as fish. The venom of a few larger species, especially the piscivorous ones, is powerful enough to kill a human being.

The chitin, along with a venom gland containing neurotoxins. Small species of these cone snails hunt small prey such as marine worms, whereas larger cone snails hunt live fish.

Molecular phylogeny research by Kraus et al. (2010)[11] based on a part of "intron 9" of the gamma-glutamyl carboxylase gene has shown that feeding on fish in Conus has evolved at least twice independently.

Harpoon and venoms

An individual Conus pennaceus attacking one of a cluster of three snails of the species Cymatium nicobaricum, in Hawaii

Cone snails use a radula tooth as a harpoon-like structure for predation. Each of these harpoons is a modified tooth, primarily made of chitin and formed inside the mouth of the snail, in a structure known as the radula. (The radula in most gastropods has rows of many small teeth, and is used for grasping at food and scraping it into the mouth.) Each specialized cone snail tooth is stored in the radula sac (an evaginated pocket in the posterior wall of the buccal cavity), except the tooth that is currently ready to be used.

The tooth is hollow and barbed, and is attached to the tip of the radula in the radular sac, inside the snail's throat. When the snail detects a prey animal nearby, it extends a long flexible tube called a proboscis towards the prey. The radula tooth is loaded with venom from the venom bulb and, still attached to the radula, is fired from the proboscis into the prey by a powerful muscular contraction. The venom paralyzes small fish almost instantly. The snail then retracts the radula, drawing the subdued prey into the mouth. After the prey has been digested, the cone snail will regurgitate any indigestible material, such as spines and scales, along with the then-disposable harpoon. There is always a dart stored in the radular sac. A dart may be used in self-defense when the snail feels threatened.


All cone snail species are equipped with a battery of toxic harpoons which can fire in any direction, even backwards. Some of these toxins can be fatal to humans.[14]

The venom of cone snails contains hundreds of different compounds, and its exact composition varies widely from one species of cone snail to another. The toxins in these various venoms are called conotoxins. These are various peptides, each targeting a specific nerve channel or receptor. Some cone snail venoms also contain a pain-reducing toxin, which the snail uses to pacify the victim before immobilising and then killing it.

Relevance to humans


A live Textile cone, Conus textile, one of the dangerous cones to handle.

The bright colors and patterns of cone snails are attractive to the eye, and therefore people sometimes pick up the live animals and hold them in their hand for a while. This is risky, because the snail often fires its harpoon in these situations. In the case of the larger species of cone snail, the harpoon is sometimes capable of penetrating the skin, even through gloves or wetsuits.

The sting of many of the smallest cone species may be no worse than that of a bee or hornet sting,[15] but in the case of a few of the larger tropical fish-eating species, especially Conus geographus, Conus tulipa and Conus striatus, a sting can sometimes have fatal consequences. Other dangerous species are Conus pennaceus, Conus textile, Conus aulicus, Conus magus and Conus marmoreus.[16] According to Goldfrank's Toxicologic Emergencies, only about 15 human deaths can be confidently attributed to cone snail envenomation.

Most of the cone snails that hunt worms rather than fish are probably not a real risk to humans, with the possible exception of larger species such as Conus vexillum or Conus quercinus. One of the fish-eating species, the geography cone, Conus geographus, is also known colloquially as the "cigarette snail," a gallows humor exaggeration implying that, when stung by this creature, the victim will have only enough time to smoke a cigarette before dying.[17][18]

Symptoms of a more serious cone snail sting include intense, localized pain, swelling, numbness and tingling and vomiting. Symptoms can start immediately or can be delayed in onset for days. Severe cases involve muscle paralysis, changes in vision, and respiratory failure that can lead to death.

Medical use of the venom

The appeal of the cone snail's venom for creating pharmaceutical drugs is the precision and speed with which the various components act; many of the compounds target a particular class of receptor, to the exclusion of any other. This means that in isolation, they can reliably and quickly produce a particular effect on the body's systems without side effects; for example, almost instantly reducing heart rate or turning off the signaling of a single class of nerve, such as pain receptors.

The venom of some cone snails such as the magician cone, Conus magus, shows much promise for providing a non-addictive pain reliever 1,000 times as powerful as morphine.[19] (See Ziconotide.)

Many peptides produced by the cone snails show prospects for being potent pharmaceuticals, such as AVC1, isolated from the Australian species, the Queen Victoria cone, Conus victoriae. This has proved very effective in treating postsurgical and neuropathic pain, even accelerating recovery from nerve injury.

The first painkiller derived from cone snail toxins, ziconotide, was approved by the U.S. Food and Drug Administration in December 2004 under the name "Prialt". Other drugs are in clinical and preclinical trials, such as compounds of the toxin that may be used in the treatment of Alzheimer's disease, Parkinson's disease, depression, and epilepsy.[20][21]

Shell collecting

The intricate color patterns of cones have made them one of the most popular collectible shells.[22][23]

Conus gloriamaris, the "Glory of the Seas" cone was, in earlier centuries, one of the most famous and sought-after seashells, with only a few specimens in private collections. This apparent rarity meant that shells of this species fetched very high prices, until finally the habitat for this cone was discovered. Sizable populations were then located, and this brought the price down dramatically.[24]

As jewelry

Naturally-occurring, beachworn cone shell "tops" (the broken-off spire of the shell, which usually end up with a hole worn at the tip) can function as beads without any further modification. In Hawaii, these natural beads were traditionally collected from the beach drift to make puka shell jewelry. Since it is hard to obtain enough naturally-occurring cone tops, almost all modern puka shell jewelry uses cheaper imitations, cut from thin shells of other species of mollusk, or even made of plastic.


Shell of Conus aulicus

The number of valid names of recent species in the genus Conus is over 600[2] and there are, in addition, a large number of fossil species.

Prior to 2009, all species within the family Conidae were still placed in one genus Conus. Testing in order to try to understand the molecular phylogeny of the Conidae was initially begun by Christopher Meyer and Alan Kohn,[25] and is continuing, particularly with the advent of nuclear DNA testing in addition to mDNA testing.

In 2009, J.K. Tucker and M.J. Tenorio proposed a classification system consisting of three distinct families and 82 genera for the living species of cone snails. This classification was based upon shell morphology, radular differences, anatomy, physiology, and cladistics, with comparisons to molecular (DNA) studies.[26] Published accounts of genera within the Conidae that use these new genera include J.K. Tucker & M.J. Tenorio (2009), and Bouchet et al. (2011).[27] Tucker and Tenorio's proposed classification system for the cone shells and their allies (and the other clades of Conoidean gastropods) is shown in Tucker & Tenorio cone snail taxonomy 2009.

Some experts, however, have preferred to keep using the traditional classification, where all species are placed in Conus within the single family Conidae: for example, according to the November 2011 version of the World Register of Marine Species, all species within the family Conidae are in the genus Conus. The binomial names of species in the 82 genera of living cone snails listed in Tucker & Tenorio 2009 are recognized by the World Register of Marine Species as "alternative representations." [28] Debate within the scientific community regarding this issue continues, and additional molecular phylogeny studies are being carried out in an attempt to clarify the issue.[26][29][30][31][32][33][34][35][36][37]

See also

  • ConoServer, a database of cone snail toxins, known as conopeptides.[38] These toxins are of importance to medical research.


  1. ^ Linnaeus C. (1758). Systema Naturae, ed. 10, 712; 1767, ed. 12, 1165.
  2. ^ a b c Linnaeus, 1758Conus.  Retrieved through: World Register of Marine Species on 20 May 2010.
  3. ^ (Czech) Pek I., Vašíček Z., Roček Z., Hajn. V. & Mikuláš R. (1996). Základy zoopaleontologie. Olomouc, 264 pp., ISBN 80-7067-599-3.
  4. ^ Olivera BM, Teichert RW (2007). "Diversity of the neurotoxic Conus peptides: a model for concerted pharmacological discovery". Molecular Interventions 7 (5): 251–60.  
  5. ^ Roger Van Oosten (September 2008). "Nature's brew". Quest online. 
  6. ^ Tenorio, M. J. & Monteiro, A. J. (2008). The Family Conidae. The South African species of Conus. In: Poppe, G. T. & Groh, K. (eds): A Conchological Iconography. Hackenheim: ConchBooks. 47 pp., 60 pls.
  7. ^ Branch, G.M. Griffiths, C.L. Branch, M.L. Beckley, L.E. (2010). Two oceans : a guide to the marine life of Southern Africa. Cape Town: Struik Nature.  
  8. ^ Monteiro, A. J., Tenorio, M. J. & Poppe, G. T. (2004). The Family Conidae. The West African and Mediterranean species of Conus. In: Poppe, G. T. & Groh, K. (eds): A Conchological Iconography. Hackenheim: ConchBooks. 102 pp., 164 pls.
  9. ^ Tenorio, M. J., Tucker, J. K. & Chaney, H. W. (2012). The Families Conilithidae and Conidae. The Cones of the Eastern Pacific. In: Poppe, G. T. & Groh, K. (eds): A Conchological Iconography. Hackenheim: ConchBooks. 112 pp., 88 pls.
  10. ^ The Conus biodiversity website
  11. ^ Kraus N. J., Corneli P. S., Watkins M., Bandyopadhyay P. K., Seger J. & Olivera B. M. (in press, available online 13 December 2010). "Against Expectation: A Short Sequence With High Signal Elucidates Cone Snail Phylogeny". Molecular Phylogenetics and Evolution doi:10.1016/j.ympev.2010.11.020.
  12. ^ National Geographic Cone Snail Profile
  13. ^ Kohn, Alan J. (March 1956). "PISCIVOROUS GASTROPODS OF THE GENUS CONUS". Proc Natl Acad Sci U S A 42 (3): 168–171.  
  14. ^ Dart, RC and Caravati, EM (2004) Medical Toxicology Lippincott Williams. ISBN 978-0-7817-2845-4
  15. ^ Marine wounds and stings
  16. ^ Killer Cones
  17. ^ NIGMS - Findings, September 2002: Secrets of the Killer Snails
  18. ^ Geographic Cone Snail, Geographic Cone Snail Profile, Facts, Information, Photos, Pictures, Sounds, Habitats, Reports, News - National Geographic
  19. ^ ANI (2007). "Sea snail venom paves way for potent new painkiller". Compassionate health care network. Retrieved 2008-11-19. 
  20. ^ Louise Yeoman (2006-03-28). "Venomous snails aid medical science". BBC. Retrieved 2008-11-19. 
  21. ^
  22. ^ Conidae - worldwideconchology
  23. ^ Conus gloriamaris
  24. ^ Conus gloriamaris, Glory of the Seas Cone photos, Phillip Colla Natural History Photography :: Online Photo Search
  25. ^ Interview of Professor Alan Kohn, Professor Emeritus, Zoology
  26. ^ a b Tucker J.K. & Tenorio M.J. (2009), Systematic Classification of Recent and Fossil Conoidean Gastropods, ConchBooks, Hankenheim, Germany, 295 pp.
  27. ^ Bouchet P., Kantor Yu.I., Sysoev A. & Puillandre N. (2011). "A new operational classification of the Conoidea". Journal of Molluscan Studies 77: 273-308.
  28. ^ Classification: Traditionally, all cone shells have been included in the Linnaean genus Conus. Tucker & Tenorio (2009) have recently proposed an alternative shell- and radula-based classification that recognizes 4 families and 80 genera of cones. In WoRMS, we currently still recognize a single family Conidae (following Puillandre et al. 2011), but Tucker & Tenorio's 80 genera classification is presented as "alternative representation". [P. Bouchet, 14 Aug. 2011]
  29. ^ C.M.L. Afonso & M.J. Tenorio (August 2011), A new, distinct endemic Africonus species (Gastropoda, Conidae) from Sao Vicente Island, Cape Verde Archipelago, West Africa, Gloria Maris 50(5): 124-135
  30. ^ P. Bouchet, Yu I. Kantor, A. Sysoev, and N. Puillandre (March 2011), A New Operational Classification of the Conoidea, Journal of Molluscan Studies 77:273-308, at p. 275.
  31. ^ N. Puillandre, E. Strong, P. Bouchet, M. Boisselier, V. Couloux, & S. Samadi (2009), Identifying gastropod spawn from DNA barcodes: possible but not yet practicable, Molecular Ecology Resources 9:1311-1321.
  32. ^ P.K. Bandyopadhyay, B.J. Stevenson, J.P. Ownby, M.T. Cady, M. Watkins, & B. Olivera (2008), The mitochondrial genome of Conus textile, coxI-conII intergenic sequences and conoidean evolution. Mollecular Phylogenetics and Evolution 46: 215-223.
  33. ^ S.T. Williams & T.F. Duda, Jr. (2008), Did tectonic activity stimulate Oligo-Miocene speciation in the Indo-West Pacific? Evolution 62:1618-1634.
  34. ^ R.L. Cunha, R. Castilho, L. Ruber, & R. Zardoya (2005), Patterns of cladogenesis in the venomous marine gastropod genus Conus from the Cape Verde Islands Systematic Biology 54(4):634-650.
  35. ^ T.F. Duda, Jr. & A.J. Kohn (2005), Species-level phylogeography and evolutionary history of the hyperdiverse marine gastropod genus Conus, Molecular Phylogenetics and Evolution 34:257-272.
  36. ^ T.F. Duda, Jr. & E. Rolan (2005), Explosive radiation of Cape Verde Conus, a marine species flock, Molecular Ecology 14:267-272.
  37. ^ B. Vallejo, Jr. (2005), Inferring the mode of speciation in the Indo-West Pacific Conus (Gastropoda: Conidae), Journal of Biogeography 32:1429-1439.
  38. ^ Kaas, Quentin; Yu Rilei; Jin Ai-Hua; Dutertre Sébastien; Craik David J (Jan 2012). "ConoServer: updated content, knowledge, and discovery tools in the conopeptide database". Nucleic Acids Res. (England) 40 (Database issue): D325–30.  

Further reading

  • Goldfrank's Toxicologic Emergencies, 8th Edition, Edited by Neal E Flomenbaum, Lewis R Goldfrank, Robert S Hoffman, Mary Ann Howland, Neal A Lewin, and Lewis S Nelson. Published by McGraw-Hill, New York, ISBN 978-0-07-143763-9
  • Gmelin, J. F. 1791. Systema naturae per regna tria naturae. Editio decima tertia. Systema Naturae, 13th ed., vol. 1(6): 3021-3910. Lipsiae.
  • Bruguière, [J.-G.] 1792. Encyclopédie Méthodique. Histoire Naturelle des Vers. Encyclopédie Méthodique. Histoire Naturelle des Vers 1: 345-757. Panckoucke: Paris.
  • Sowerby, G. B., II. 1833. Conus. Conchological Illustrations pls. 36-37
  • (French) Bernardi A. C. (1858). Monographie du genre Conus.
  • Reeve, L. 1844. Monograph of the genus Conus. Conchologia Iconica 1: pls. 40-47
  • Kiener, L. C. 1845. Genre Cone. (Conus, Lin.). Spécies Général et Iconographie des Coquilles Vivantes 2: pls. 1-111
  • Clench, W. J. 1942. The Genus Conus in the Western Atlantic. Johnsonia 1(6): 1-40
  • Van Mol, J. J., B. Tursch and M. Kempf. 1967. Mollusques prosobranches: Les Conidae du Brésil. Étude basée en partie sur les spécimens recueillis par la Calypso. Annales de l'Institut Océanographique 45: 233-254, pls. 5-10
  • Vink, D. L. N. and R. von Cosel. 1985. The Conus cedonulli complex: Historical review, taxonomy and biological observations. Revue Suisse de Zoologie 92: 525-603
  • Petuch, E. J. 1986. New South American gastropods in the genera Conus (Conidae) and Latirus (Fasciolariidae). Proceedings of the Biological Society of Washington 99: 8-14.
  • Petuch, E. J. 1987. New Caribbean molluscan faunas. [v] + 154 + A1-A4, 29 pls. Coastal Education & Research Foundation: Charlottesville, Virginia
  • Petuch, E. J. 1988. Neogene history of tropical American mollusks. [vi] + 217, 39 pls. Coastal Education & Research Foundation: Charlottesville, Virginia
  • Petuch, E. J. 1990. A new molluscan faunule from the Caribbean coast of Panama. Nautilus 104: 57-70
  • Petuch, E. J. 1992. Molluscan discoveries from the tropical Western Atlantic region. Part II. New species of Conus from the Bahamas Platform, Central American and northern South American coasts, and the Lesser Antilles. La Conchiglia 24(265): 10-15.
  • Petuch, E. J. 2000. A review of the conid subgenus Purpuriconus da Motta, 1991, with the descriptions of two new Bahamian species. Ruthenica 10: 81-87
  • Petuch, E. J. 2004. Cenozoic Seas. xvi + 308 pp. CRC Press: Boca Raton
  • Tenorio, M. J., Tucker, J. K. & Chaney, H. W. (2012). The Families Conilithidae and Conidae. The Cones of the Eastern Pacific. In: Poppe G.T. & Groh K. (eds): A Conchological Iconography. Hackenheim: ConchBooks. 112 pp., 88 pls.
  • Coltro, J., Jr. 2004. New species of Conidae from northeastern Brazil (Mollusca: Gastropoda). Strombus 11: 1-16
  • García, E. F. 2006. Conus sauros, a new Conus species (Gastropoda: Conidae) from the Gulf of Mexico. Novapex 7: 71-76
  • Franklin JB, Subramanian KA, Fernando SA, & Krishnan KS 2009. Diversity and Distribution of Conidae from the Tamil Nadu Coast of India (Mollusca: Caenogastropoda: Conidae). Zootaxa 2250: 63 pp.
  • J. Benjamin Franklin, S. Antony Fernando, B. A. Chalke, K. S. Krishnan 2007. (Gastropoda: Caenogastropoda: Conidae) from IndiaConusRadular morphology of . Molluscan Research 27(3): 111–122. ISSN 1323-5818

External links

  • SEM images of its radula can be found at Thompson; Bebbington (1973). Malacologia 14: 147–165. .
  • Tucker J.K. (2009). Recent cone species database. September 4th 2009 Edition
  • Filmer R.M. (2001). A Catalogue of Nomenclature and Taxonomy in the Living Conidae 1758 - 1998. Backhuys Publishers, Leiden. 388pp
  • Bouchet, P.; Fontaine, B. (2009). List of new marine species described between 2002-2006. Census of Marine Life.
  • shellsConusNatural History Museum Rotterdam - photos of
  • Cone snail and conotoxins page
  • Biodiversity websiteConusThe
  • ConoServer
  • Conidae from Scroll down for many photographs.
  • Pain-killer comes out of its shell (The Age news article)
  • Venomous snails aid medical science (BBC News Article).
  • ConeShell Collection Giancarlo Paganelli
  • Cone Shells - Knights of the Sea. Alexander Medvedev's collection
  • Cone Snail Video - Hunting Footage and Physiology
  • Deadly Critters That Might Save Your Life (CNN)
  • Baldomero "Toto" Olivera's short talk: Conus Peptides
  • Zonatus Gallery
  • Miller, John A. (1989). "The toxoglossan proboscis: structure and function". Journal of Molluscan Studies 55 (2): 167–181.   [1]
  • BBC Nature Video Cone snails are silent assassins of the sea, drugging sleeping fish before poisoning them
This article was sourced from Creative Commons Attribution-ShareAlike License; additional terms may apply. World Heritage Encyclopedia content is assembled from numerous content providers, Open Access Publishing, and in compliance with The Fair Access to Science and Technology Research Act (FASTR), Wikimedia Foundation, Inc., Public Library of Science, The Encyclopedia of Life, Open Book Publishers (OBP), PubMed, U.S. National Library of Medicine, National Center for Biotechnology Information, U.S. National Library of Medicine, National Institutes of Health (NIH), U.S. Department of Health & Human Services, and, which sources content from all federal, state, local, tribal, and territorial government publication portals (.gov, .mil, .edu). Funding for and content contributors is made possible from the U.S. Congress, E-Government Act of 2002.
Crowd sourced content that is contributed to World Heritage Encyclopedia is peer reviewed and edited by our editorial staff to ensure quality scholarly research articles.
By using this site, you agree to the Terms of Use and Privacy Policy. World Heritage Encyclopedia™ is a registered trademark of the World Public Library Association, a non-profit organization.

Copyright © World Library Foundation. All rights reserved. eBooks from Hawaii eBook Library are sponsored by the World Library Foundation,
a 501c(4) Member's Support Non-Profit Organization, and is NOT affiliated with any governmental agency or department.