Archerfish
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The archerfish (also known as spinner fish or archer fish) or Toxotidae are a monotypic family (although some include a second genus) of perciform tropical fish known for their unique predation technique of "shooting down" land-based insects and other small prey with jets of water spit from their specialized mouths. The family is small, consisting of ten species in a single genus, Toxotes. Most archerfish live in freshwater streams, ponds and wetlands, but two or three species are euryhaline, inhabiting both fresh and brackish water habitats such as estuaries and mangroves.<ref name=Kottelat2018>Template:Cite journal</ref><ref>Arthington, A., and McKenzie, F. "Review of Impacts of Displaced/Introduced Fauna Associated with Inland Waters. Template:Webarchive" Environment Australia Template:Webarchive Australia: State of the Environment Technical Paper Series (Inland Waters), Series 1, 1997. Accessed 2009-05-24.</ref> They can be found from India, Bangladesh and Sri Lanka, through Southeast Asia, to Melanesia and Northern Australia.<ref name=FishBaseGenus>{{#invoke:Cite taxon|main|fishbase|genus=|species=|subspecies=}}</ref><ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>
Archerfish have deep and laterally compressed bodies, with the dorsal fin and the profile a straight line from dorsal fin to mouth. The mouth is protractile, and the lower jaw juts out. Sizes are fairly small, typically up to about Template:Convert, but T. chatareus can reach Template:Convert.<ref name=FishBaseGenus/><ref name=EoF>Template:Cite book</ref>
Archerfish are popular exotic fish for aquaria,<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> but are difficult to feed and maintain by average fishkeepers since they prefer live prey over typical fish foods.
Capture of preyEdit
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Archerfish are remarkably accurate in their shooting; an adult fish almost always hits the target on the first shot. Although it is presumed that all archerfish species do this, it has only been confirmed from T. blythii, T. chatareus and T. jaculatrix.<ref name=Kottelat2018/> They can bring down insects and other prey<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> up to Template:Convert above the water's surface.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> This is partially due to their good eyesight, but also to their ability to compensate for the refraction of light as it passes through the air-water interface when aiming at their prey.<ref name="Schuster animal cognition">Template:Cite journal</ref> They typically spit at prey at a mean angle of about 74° from the horizontal but can still aim accurately when spitting at angles between 45° and 110°.<ref>Temple, S. E. "Effect of salinity on the refracive index of water: considerations for archer fish aerial vision" 'Journal of Fish Biology 1269.,74”’(1629 2007.</ref>
When an archerfish selects its prey, it rotates its eye so that the image of the prey falls on a particular portion of the eye in the ventral temporal periphery of the retina,<ref>Temple, S.E., Hart, N. S., and Colin, S. P. "A spitting image: visual specializations of the arsherfish (Toxotes chatareus)" 'Brain Behaviour and Evolution' Vol. 73, 309 2009.</ref> and its lips just break the surface, squirting a jet of water at its victim. The archerfish does this by forming a small groove in the roof of its mouth and its tongue into a narrow channel. It then fires by contracting its gill covers and forcing water through the channel, shooting a stream that, shaped by its mouth parts, travels faster at the rear than at the front. This speed differential causes the stream to become a blob directly before impact as the slower leading water is overtaken by the faster trailing water, and it is varied by the fish to account for differences in range. It also makes this one of the few animals that both make and use tools, as they both utilise the water and shape it to make it more useful to them.<ref name="SN Oct 4 2014">Template:Cite journal</ref><ref>Template:Cite magazine</ref> They are persistent and will make multiple shots if the first one fails.<ref>Template:Cite journal</ref>
Young archerfish start shooting when they are about Template:Convert long but are inaccurate at first and must learn from experience.Template:Citation needed During this learning period, they hunt in small schools. This way, the probability is enhanced that at least one jet will hit its target.Template:Citation needed A 2006 experimental study found that archerfish appear to benefit from observational learning by watching a performing group member shoot, without having to practice: <templatestyles src="Template:Blockquote/styles.css" />
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{{#invoke:Check for unknown parameters|check|unknown=Template:Main other|preview=Page using Template:Blockquote with unknown parameter "_VALUE_"|ignoreblank=y| 1 | 2 | 3 | 4 | 5 | author | by | char | character | cite | class | content | multiline | personquoted | publication | quote | quotesource | quotetext | sign | source | style | text | title | ts }}However, little of their social behaviour is currently known beyond that archerfish are sensitive to, and make changes to their shooting behaviour, when conspecifics are visible to them.<ref>Jones, N. A. R., Webster, M. M., Templeton, C. N., Schuster, S., & Rendell, L. (2018). Presence of an audience and consistent interindividual differences affect archerfish shooting behaviour. Animal Behaviour, 141, 95–103. https://doi.org/10.1016/j.anbehav.2018.04.024
</ref> This is probably as a result of the potential threat of kleptoparasitism that other archerfish represent to a shooting fish.<ref>Davis, B. D., & Dill, L. M. (2012). Intraspecific kleptoparasitism and counter-tactics in the archerfish (Toxotes chatareus). Behaviour, 149(13–14), 1367–1394. https://doi.org/10.1163/1568539X-00003026 </ref>
An archerfish will often leap out of the water and grab an insect in its mouth if it happens to be within reach. Individuals typically prefer to remain close to the surface of the water.<ref name="SN Oct 4 2014" />
New research has found that archerfish also use jets to hunt underwater prey, such as those embedded in silt. It is not known whether they learned aerial or underwater shooting first, but the two techniques may have evolved in parallel, as improvements in one can be adapted to the other.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> This makes it an example of exaptation.
SpeciesEdit
There are 9 valid species, 8 in the genus Toxotes:<ref name=Kottelat2018/><ref name=FishBaseGenus/><ref name=FishBaseFamily>{{#invoke:Cite taxon|main|fishbase|genus=|species=|subspecies=}}</ref><ref>Template:Cite journal</ref><ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>
- Protoxotes lorentzi Weber, 1910 - primitive archerfish
- Toxotes blythii Boulenger, 1892 - clouded archerfish, zebra archerfish
- Toxotes carpentariensis, Castelnau, 1878
- Toxotes chatareus (Hamilton, 1822) - largescale archerfish, common archerfish
- Toxotes jaculatrix (Pallas, 1767) - banded archerfish
- Toxotes kimberleyensis Allen, 2004 - Kimberley archerfish, western archerfish
- Toxotes microlepis Günther, 1860 - smallscale archerfish
- Toxotes oligolepis Bleeker, 1876 - big scale archerfish
- Toxotes sundaicus Kottelat & Tan, 2018
TimelineEdit
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