Rasberry crazy ant

Last updated

Nylanderia fulva
Tawny crazy ant (Nylanderia fulva) female worker.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Formicinae
Genus: Nylanderia
Species:
N. fulva
Binomial name
Nylanderia fulva
Mayr 1862 [1]
Synonyms

Prenolepis fulva

The tawny crazy ant [2] [3] [4] or Rasberry crazy ant, [2] Nylanderia fulva, is an ant originating in South America. Like the longhorn crazy ant (Paratrechina longicornis), this species is called "crazy ant" because of its quick, unpredictable movements (the related N. pubens is known as the "Caribbean crazy ant"). It is sometimes called the "Rasberry crazy ant" in Texas after the exterminator Tom Rasberry, who noticed that the ants were increasing in numbers in 2002. [5] [6] Scientists have reorganised the genera taxonomy within this clade of ants, and now it is identified as Nylanderia fulva. [7]

Contents

In 2014, it was discovered that the ant produces and covers itself with formic acid as an antidote to the fire ant's venom. [8] It is the first known example of an insect being able to neutralize another insect's venom, an ability speculated to have evolved in South America where the two species share the same native range. Colonies have multiple queens, which also contributes to their survival. [9]

As of 2012, the ants have established colonies [3] [4] in all states of the Gulf Coast of the United States including at least 27 counties in Southeast Texas.[ citation needed ]

Description

The ant is about 3 mm (or about 1/8 inches) long, thus smaller than the red imported fire ant, Solenopsis invicta . It is covered with reddish-brown hairs. Their larvae are plump and hairy, with a specific conformation of mouthparts and unique mandible morphology that allows for precise species identification. [10] The colonies live under stones or piles; they have no centralized nests, beds, or mounds. [2] They tend aphids for honeydew, feed on small insects and vertebrates, and forage on plants, especially for sweet materials. The ants appear to prefer the warmth and moisture of the coast. [11]

N. fulva has been a pest in rural and urban areas of Colombia, and South America, where it displaced all other ant species. There, small poultry such as chickens have died of asphyxiation while larger animals, such as cattle, have been attacked around the eyes, nostrils, and hooves. Grasslands have dried out because of the increase in plant-sucking insect pests (hemipterans), which the ants cultivate to feed on the sugary "honeydew" that they excrete. [2]

When attacked, these ants, like other formicine ants, can bite (but not sting) and excrete formic acid through a hairy circle or acidopore on the end of the abdomen, using it as a venom, [12] which causes a minute pain that quickly fades. Formic acid was named after the Latin word formica (ant), because it was first distilled from ants in the 17th century. [13] Uniquely, the tawny ant also uses formic acid as an antidote against the venom alkaloids of the fire ant (known as solenopsins). The venom alkaloids of fire ants have been demonstrated to be strongly paralytic against competitor species, [14] thus the tawny crazy ant may have developed a resistance by acid-immobilisation of the venom toxins.

Tawny crazy ants were found to displace other ant species in their native Argentina and later the US, including the red imported fire ant. [6] This was first thought to be due to exploitative and interference competition. [15]

Formic acid as an antidote to fire ant venom

In March 2014, researchers concluded that formic acid helped tawny crazy ants survive fire ant venom in ant fights 98% of the time; when the gland ducts were blocked with nail polish in an experiment, crazy ants had only a 48% chance of surviving fights with fire ants. [8] After exposure to fire ant venom, N. fulva retreats, covers itself with formic acid [16] and returns to the fight. [8] This is the first known example of an insect detoxifying another insect's venom, and the first discovery of an ionic liquid in nature which results from mixing of formic acid with venom from S. invicta.

How formic acid acts as an antidote against the much more toxic fire ant's venom is unknown. Fire ant venom is a mixture of toxic alkaloids and proteins that presumably enable the alkaloids to enter rival ants’ cells. [13] Each alkaloid in the fire ant's venom, including solenopsin, has a six-membered heterocyclic ring with fat-soluble side chains. [13] The researchers who discovered the antidote property of formic acid in crazy ants speculate that the formic acid denatures the proteins in fire ant venom. [8] Another possibility is that the nitrogen on an alkaloid's heterocyclic ring is protonated, rendering the ionic molecule less lipophilic, thus less likely to penetrate the tawny crazy ant's cells. [13]

Diet

N. fulva eats liquids, [3] [4] including plant nectar [3] [4] and insect honeydew. [3] [4] [17] Calcium, sodium, and potassium are very important to N. fulva: Higher environmental potassium decreased abundance, and higher potassium + sodium decreased abundance even more so. [3] [4] Meanwhile, higher calcium increased abundance. [3] [4]

Attraction to electrical equipment

Infestations of Nylanderia fulva in electrical equipment can cause short circuits, sometimes because the ants chew through insulation and wiring. [3] Overheating, corrosion, and mechanical failures also result from accumulations of dead ants and nest detritus in electrical devices. [18] If an ant is electrocuted, it can release an alarm pheromone in dying, which causes other ants to rush over and search for attackers. If a large enough number of ants gathers, it may short out systems. [19]

It is unclear why colonies of Nylanderia fulva are attracted to electrical equipment. [6] [20] They may sense the magnetic fields that surround wires conducting electric current, or they may prefer the warmth produced by the resistance of the current in the wires. Some argue they simply are searching for food or an attractive place to nest. [21]

Rate of spread

The Texas A&M University research extension service quotes the annual rate of spread by ground migration as about 240 and 360 m per year in neighborhoods and industrial areas, respectively, and 207 m/year in rural landscapes [22] hence spreading more slowly than fire ants. [16] Other sources quote 800 m (0.50 mi) per year. [6] Being carried by people, animals, and vehicles (in trash for example), the observed rate is much higher: the spread from five Texas counties in 2002 to 20 in 2007 yields an accelerated rate of 8 km (5.0 mi) per year, at which rate it would take about 70 years for them to reach New Orleans. However, in 2011, tawny crazy ants were reported in Mississippi, [23] in August 2012 in Port Allen, Louisiana, [24] and in 2013 in Georgia. [25]

Range in the United States

Reported distribution of the Rasberry crazy ant in the United States (2012); actual occurrence is thought to be more widespread Reported distribution of the Rasberry Crazy Ant in the United States - journal.pone.0045314.g001.png
Reported distribution of the Rasberry crazy ant in the United States (2012); actual occurrence is thought to be more widespread

The earliest record of N. fulva presence in the US is from Brownsville, Texas, in 1938. [26] By the early 2000s, the ants spread across the southeastern portion of Texas [7] including at more than 27 counties [22] Large population explosions have been described also on St Croix in the US Virgin Islands; in many cases the ant species was misidentified as its close relative, the hairy crazy ant, Nylanderia pubens. [5] [27] [28] [29] As of 2012, the ants have established colonies in all states of the Gulf Coast of the United States. [6] [7] The ant is considered an invasive species. [2] As of 2021N. fulva establishment is limited to some southern parts of the country. [3]

Control in the US

The ants are not attracted to ordinary ant baits, and are not controlled by over-the-counter pesticides, [6] [30] and are harder to fully exterminate than many other species because their colonies have multiple queens. [9] In June 2008, the United States Environmental Protection Agency granted temporary approval for the use of fipronil, an antitermite agent, to control this ant. [31] Its use is currently restricted to infested counties. [32]

In 2015, the microsporidian parasite Myrmecomorba nylanderiae was found to be a pathogen of the tawny crazy ant. [33] [34] In March 2022, further research indicated that this unicellular fungus may be an effective biological control for the tawny ant. [35] [36]

See also

Related Research Articles

<span class="mw-page-title-main">Alkaloid</span> Class of naturally occurring chemical compounds

Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of similar structure may also be termed alkaloids. In addition to carbon, hydrogen and nitrogen, alkaloids may also contain oxygen or sulfur. More rarely still, they may contain elements such as phosphorus, chlorine, and bromine.

<span class="mw-page-title-main">Ant</span> Family of insects

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 22,000 species have been classified. They are easily identified by their geniculate (elbowed) antennae and the distinctive node-like structure that forms their slender waists.

<span class="mw-page-title-main">Yellow crazy ant</span> Species of ant (Anoplolepis gracilipes)

The yellow crazy ant, also known as the long-legged ant or Maldive ant, is a species of ant, thought to be native to West Africa or Asia. They have been accidentally introduced to numerous places in the world's tropics.

<span class="mw-page-title-main">Formicinae</span> Subfamily of ants

The Formicinae are a subfamily within the Formicidae containing ants of moderate evolutionary development.

<span class="mw-page-title-main">Fire ant</span> Genus of red ants

Fire ants are several species of ants in the genus Solenopsis, which includes over 200 species. Solenopsis are stinging ants, and most of their common names reflect this, for example, ginger ants and tropical fire ants. Many of the names shared by this genus are often used interchangeably to refer to other species of ant, such as the term red ant, mostly because of their similar coloration despite not being in the genus Solenopsis. Both Myrmica rubra and Pogonomyrmex barbatus are common examples of non-Solenopsis ants being termed red ants.

<span class="mw-page-title-main">Red imported fire ant</span> Invasive ant species

Solenopsis invicta, the fire ant, or red imported fire ant (RIFA), is a species of ant native to South America. A member of the genus Solenopsis in the subfamily Myrmicinae, it was described by Swiss entomologist Felix Santschi as a variant of S. saevissima in 1916. Its current specific name invicta was given to the ant in 1972 as a separate species. However, the variant and species were the same ant, and the name was preserved due to its wide use. Though South American in origin, the red imported fire ant has been accidentally introduced in Australia, New Zealand, several Asian and Caribbean countries, Europe and the United States. The red imported fire ant is polymorphic, as workers appear in different shapes and sizes. The ant's colours are red and somewhat yellowish with a brown or black gaster, but males are completely black. Red imported fire ants are dominant in altered areas and live in a wide variety of habitats. They can be found in rainforests, disturbed areas, deserts, grasslands, alongside roads and buildings, and in electrical equipment. Colonies form large mounds constructed from soil with no visible entrances because foraging tunnels are built and workers emerge far away from the nest.

<i>Paratrechina</i> Genus of ants

Paratrechina is one of seven ant genera in the Prenolepis genus-group from the subfamily Formicinae. Six species are included in Paratrechina; one of which, the longhorn crazy ant, is a widespread, pantropical pest.

Fulvous is a colour, sometimes described as dull orange, brownish-yellow or tawny; it can also be likened to a variation of buff, beige or butterscotch. As an adjective it is used in the names of many species of birds, and occasionally other animals, to describe their appearance. It is also used as in mycology to describe fungi with greater colour specificity, specifically the pigmentation of the surface cuticle, the broken flesh and the spores en masse.

<i>Nylanderia pubens</i> Species of ant

Nylanderia pubens is a species of ant of the genus Nylanderia, commonly called the Caribbean crazy ant. It was originally described as Paratrechina pubens from Saint Vincent, Lesser Antilles. This species was moved from Paratrechina to the genus Nylanderia in 2010.

<i>Lasius neglectus</i> Species of ant

Lasius neglectus is a polygynous, sometimes invasive, ant of the genus Lasius. The ant was identified in 1990 after establishing a colony in Budapest, Hungary. Superficially, they are similar in appearance to the common black garden ant, Lasius niger, but have significantly different behavioural patterns, particularly in the social structure within colonies.

<span class="mw-page-title-main">Ant venom</span> Medical condition

Ant venom is any of, or a mixture of, irritants and toxins inflicted by ants. Most ants spray or inject a venom, the main constituent of which is formic acid only in the case of subfamily Formicinae.

<i>Solenopsis saevissima</i> Species of ant

Solenopsis saevissima, commonly known in Brazil as formiga de fogo, formiga-vermelha, or formiga-lava-pes, is one of more than 185 species in the genus Solenopsis. It, along with 13 other species, is also a member of the Solenopsis saevissima species group which are popularly known as fire ants.

<span class="mw-page-title-main">Longhorn crazy ant</span> Species of ant

The longhorn crazy ant, also known as the black crazy ant, is a species of small, dark-coloured insect in the family Formicidae. These ants are commonly called "crazy ants" because instead of following straight lines, they dash around erratically. They have a broad distribution, including much of the tropics and subtropics, and are also found in buildings in more temperate regions, making them one of the most widespread ant species in the world. This species, as well as all others in the ant subfamily Formicinae, cannot sting. However, this species can fire/shoot a formic acid spray from its abdomen when under attack by other insects or attacking other insects. When the longhorn crazy ant bends its abdomen while aiming at an enemy insect, it is typically shooting its hard-to-see acid.

<span class="mw-page-title-main">Red imported fire ants in the United States</span>

The red imported fire ant, or simply RIFA, is one of over 280 species in the widespread genus Solenopsis. It is native to South America but it has become both a pest and a health hazard in the southern United States as well as a number of other countries.

<i>Nylanderia</i> Genus of ants

Nylanderia is a large genus of ants in the subfamily Formicinae. The genus has a nearly cosmopolitan distribution with species inhabiting a wide array of habitats in almost all geographic regions. Nylanderia, currently containing over 110 species, is an ecologically important genus, with some species reported as being invasive. The ants are small to medium in size and range in color from pale yellow to black.

<i>Paratrechina zanjensis</i> Species of ant

Paratrechina zanjensis is an African species of ant in the genus Paratrechina. It is one of two species in the genus, the other being the longhorn crazy ant.

Macrodinychus multispinosus is a mite that lives as an ectoparasitoid on the invasive Paratrechina longicornis. These mites complete their development on a single host, sucking all of its body content and therefore killing it. This is the ninth species of Macrodinychus reported as ant parasite, and the third known as parasitoid of invasive ants, confirming a unique habit in the evolution of mite feeding strategies and suggesting that the entire genus might be parasitic on ants.

The toxicology of fire ant venom is relatively well studied. The venom plays a central role in the biology of Red imported fire ants, such as in capturing prey, and in defending itself from competitors, assailants, and diseases. Some 14 million people are stung annually in the United States, suffering reactions that vary from mild discomfort, to pustule formation, swelling, and in rare cases, systemic reactions followed by anaphylactic shock. Fire ant venoms are mainly composed (>95%) of a complex mixture of insoluble alkaloids added to a watery solution of toxic proteins. For the Red imported fire ant Solenopsis invicta Buren there are currently 46 described proteins, of which four are well-characterised as potent allergens.

In ecology, a tramp species is an organism that has been spread globally by human activities. The term was coined by William Morton Wheeler in the bulletin of the American museum of natural history in 1906, used to describe ants that “have made their way as well known tramps or stow-aways[sic] to many islands The term has since widened to include non-ant organisms, but remains most popular in myrmecology. Tramp species have been noted in multiple phyla spanning both animal and plant kingdoms, including but not limited to arthropods, mollusca, bryophytes, and pteridophytes. The term "tramp species" was popularized and given a more set definition by Luc Passera in his chapter of David F William's 1994 book Exotic Ants: Biology, Impact, And Control Of Introduced Species.

<span class="mw-page-title-main">Ant supercolony</span> Exceptionally large ant colony

An ant supercolony is an exceptionally large ant colony, consisting of a high number of spatially separated but socially connected nests of a single ant species, spread over a large area without territorial borders. Supercolonies are typically polygynous, containing many egg-laying females. Workers and queens from different nests within the same supercolony can freely move among the nests, and all workers cooperate indiscriminately with each other in collecting food and care of the brood, and show no apparent mutual aggressive behavior.

References

  1. ↑ Mayr, G. 1862. Myrmecologische Studien.[ dead link ] Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien 12:649-776.
  2. 1 2 3 4 5 "Tawny (Rasberry) Crazy Ant. Nylanderia fulva". Texas A&M, Department of Entomology, Center for Urban & StructuralEntomology. Retrieved 2013-05-21.
  3. 1 2 3 4 5 6 7 8 9 Reihart, Ryan (2021-01-21). "Invasive tawny crazy ants have an intense craving for calcium – with implications for their spread in the US". The Conversation. Retrieved 2021-01-23.
  4. 1 2 3 4 5 6 7 Reihart, Ryan W.; Angelos, Kiersten P.; Gawkins, Kaitlin M.; Hurst, Shania E.; Montelongo, Denise C.; Laws, Angela N.; Pennings, Steven C.; Prather, Chelse M. (2021-01-11). "Crazy ants craving calcium: macronutrients and micronutrients can limit and stress an invaded grassland brown food web". Ecology . 102 (2). Wiley: e03263. doi: 10.1002/ecy.3263 . ISSN   0012-9658. PMID   33314072. S2CID   229178510.
  5. 1 2 Ayres, Chris (2008-05-16). "Billions of electronic-eating 'crazy Rasberry ants' invade Texas". Times UK.
  6. 1 2 3 4 5 6 Main, Douglas (17 May 2013). "'Crazy' Ants Driving Out Fire Ants in Southeast". LiveScience.com. Retrieved 2013-05-18.
  7. 1 2 3 Gotzek, D.; Brady, S. N. G.; Kallal, R. J.; Lapolla, J. S. (2012). Moreau, Corrie S (ed.). "The Importance of Using Multiple Approaches for Identifying Emerging Invasive Species: The Case of the Rasberry Crazy Ant in the United States". PLOS ONE. 7 (9): e45314. Bibcode:2012PLoSO...745314G. doi: 10.1371/journal.pone.0045314 . PMC   3462614 . PMID   23056657.
  8. 1 2 3 4 LeBrun, Edward G.; Nathan T. Jones; Lawrence E. Gilber (28 February 2014). "Chemical Warfare Among Invaders: A Detoxification Interaction Facilitates an Ant Invasion". Science. 343 (6174): 1014–1017. Bibcode:2014Sci...343.1014L. doi: 10.1126/science.1245833 . PMID   24526314. S2CID   45087292.
  9. 1 2 Can Ants Eat Your Computer: Why the "crazy rasberry" ant infests electronic devices., Slate , 20 May 2008.
  10. ↑ Correa Bueno, Odair; Rossi, Monica Lanzoni; Solis, Daniel Russ; Fox, Eduardo Gonçalves Paterson (2018-01-02). "Morphological Studies on the Mature Worker Larvae of Paratrechina fulva (Hymenoptera, Formicidae)". doi:10.6084/m9.figshare.5746644.v1.{{cite journal}}: Cite journal requires |journal= (help)
  11. ↑ Ralph Blumenthal, A Pest Without a Name, Becoming Known to Ever More, The New York Times , May 16, 2008
  12. ↑ Touchard, Axel; Aili, Samira; Fox, Eduardo; Escoubas, Pierre; Orivel, Jérôme; Nicholson, Graham; Dejean, Alain (2016-01-20). "The Biochemical Toxin Arsenal from Ant Venoms". Toxins. 8 (1): 30. doi: 10.3390/toxins8010030 . ISSN   2072-6651. PMC   4728552 . PMID   26805882.
  13. 1 2 3 4 Everts, Sarah (3 March 2014). "An Ant's Acid Antidote". Chemical & Engineering News. 92 (9): 44–45. doi:10.1021/cen-09209-scitech3 . Retrieved 29 April 2014.
  14. ↑ Fox, Eduardo G.P.; Wu, Xiaoqing; Wang, Lei; Chen, Li; Lu, Yong-Yue; Xu, Yijuan (February 2019). "Queen venom isosolenopsin A delivers rapid incapacitation of fire ant competitors". Toxicon. 158: 77–83. doi:10.1016/j.toxicon.2018.11.428. PMID   30529381. S2CID   54481057.
  15. ↑ Danny Lee McDonald (December 2012). "Investigation of an invasive ant species: Nylanderia fulva colony extraction, management, diet preference, fecundity, and mechanical vector potential" (PDF). aglifesciences.tamu.edu. Archived from the original (PDF) on 22 September 2013. Retrieved 19 November 2018.
  16. 1 2 University of Texas at Austin (February 13, 2014). "Crazy ants dominate fire ants by neutralizing their venom". ScienceDaily. Retrieved 16 February 2014.
  17. ↑ Sharma, Shweta; Oi, David H.; Buss, Eileen A. (2013). "Honeydew-Producing Hemipterans in Florida Associated with Nylanderia fulva (Hymenoptera: Formicidae), an Invasive Crazy Ant". The Florida Entomologist . 96 (2). Florida Entomological Society: 538–547. doi: 10.1653/024.096.0219 . ISSN   0015-4040. S2CID   54668879.
  18. ↑ NASA moves to save computers from swarming ants | Computerworld, Computerworld, 15 May 2008.
  19. ↑ McConnaughey, Janet. "'Hairy crazy ants' invade from Florida to Texas - Technology & science - Science - NBCNews.com". NBC News. Retrieved 2012-08-13.[ dead link ]
  20. ↑ Opam, Kwane (26 September 2011). "These Ants Terrorize Everything—Even Gadgets". gizmodo.com. Retrieved 2 October 2011.
  21. ↑ Mooallem, Jon (2013-12-05). "There's a Reason They Call Them 'Crazy Ants'". The New York Times.
  22. 1 2 "Tawny (Rasberry) Crazy Ant". University of Texas A & M University, Center for Urban and Structural Entomology. Retrieved 16 February 2014.
  23. ↑ Joe A. MacGown. Tawny crazy Ants invade Mississippi. YouTube video. Posted 2011-09-29.
  24. ↑ Kiran Chawla. Army of hard to kill ants marches into south Louisiana. WAFB, Port Allen, LA. Posted 2012-08-14.
  25. ↑ Sharon Dowdy. Invasive tawny crazy ant found in Georgia. University of Georgia. Posted 2013-09-17.
  26. ↑ http://gap.entclub.org/taxonomists/Trager/1984b.pdf%5B%5D
  27. ↑ Wetterer, James K.; Keularts, Jozef L. W. (September 2008). "Population explosion of the hairy crazy ant, Paratrechina pubens (hymenoptera: formicidae), on St. Croix, US Virgin Islands". Florida Entomologist. 91 (3): 423–427. doi: 10.1653/0015-4040(2008)91[423:peothc]2.0.co;2 . S2CID   86286910.
  28. ↑ Robert Lee (20 September 2013). "'Crazy Ants': The ants that destroy electronics march into Georgia". www.wsbtv.com. Archived from the original on 21 September 2013. Retrieved 15 December 2018– via web.archive.org.
  29. ↑ Arjan van den Bosch. "Ccrazy Rasberry ant". Ant-maps.com. Archived from the original on 1 August 2009. Retrieved 15 December 2018.
  30. ↑ Ants swarm over Houston area, fouling electronics, Yahoo News, 15 May 2008. Archived 19 May 2008 at the Wayback Machine
  31. ↑ Pesticide for SE Texas 'crazy' ants approved by EPA, Houston Chronicle, 2008-07-01, retrieved 2008-07-01
  32. ↑ FIFRA Sec. 18 Quarantine Exemption Use Directions, Texas Department of Agriculture, 2015, retrieved 2016-06-14
  33. ↑ Plowes, RM; Becnel, JJ; LeBrun, EG; Oi, DH; Valles, SM; Jones, NT; Gilbert, LE (July 2015). "Myrmecomorba nylanderiae gen. et sp. nov., a microsporidian parasite of the tawny crazy ant Nylanderia fulva". Journal of Invertebrate Pathology. 129: 45–56. doi:10.1016/j.jip.2015.05.012. PMID   26031565.
  34. ↑ Wang, Z; Moshman, L; Kraus, EC; Wilson, BE; Acharya, N; Diaz, R (15 December 2016). "A Review of the Tawny Crazy Ant, Nylanderia fulva, an Emergent Ant Invader in the Southern United States: Is Biological Control a Feasible Management Option?". Insects. 7 (4): 77. doi: 10.3390/insects7040077 . PMC   5198225 . PMID   27983690.
  35. ↑ Stokstad, Eric (28 March 2022). "Invasive crazy ants could meet their match in a mysterious, funguslike pathogen". Science. AAAS.
  36. ↑ LeBrun, Edward G.; Jones, Melissa; Plowes, Robert M.; Gilbert, Lawrence E. (5 April 2022). "Pathogen-mediated natural and manipulated population collapse in an invasive social insect". Proceedings of the National Academy of Sciences. 119 (14): e2114558119. Bibcode:2022PNAS..11914558L. doi:10.1073/pnas.2114558119. PMC   9168452 . PMID   35344435. S2CID   247778033.

Further reading

This page is based on this Wikipedia article
Text is available under the CC BY-SA 4.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.