Archerfish

{{Short description|Family of fishes}} {{for|ships with the name Archerfish, Archer-Fish|USS Archerfish}} {{Automatic taxobox | fossil_range = {{fossil range|Eocene|present}} | image = Toxotes jaculatrix.jpg | image_caption = ''Toxotes jaculatrix'' | taxon = Toxotidae | authority = Bleeker, 1859 | type_species = ''Labrus jaculator'' | type_species_authority = Shaw, 1803 | subdivision_ranks = Genera | subdivision = * ''Protoxotes'' Whitley, 1950 * ''Toxotes'' Cloquet, 1816 }}

The '''archerfish''' (also known as '''spinner fish''' or '''archer fish''') or '''Toxotidae''' are a family of percomorph 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 two genera, ''Toxotes'' and ''Protoxotes''.<ref name="CofFF">{{Cof family|family=Toxotidae|access-date=22 December 2025}}</ref> 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>{{cite journal| author1=Maurice Kottelat| author-link=Maurice Kottelat| author2=Tan Heok Hui | year=2018 | title=Three new species of archerfishes from the freshwaters of Southeast Asia (Teleostei: Toxotidae) and notes on Henri Mouhot's fish collections | journal=Ichthyological Exploration of Freshwaters | volume=IEF-952 | pages=1–19 | doi=10.23788/IEF-952 }}</ref><ref>Arthington, A., and McKenzie, F. "[http://www.environment.gov.au/soe/1996/publications/technical/pubs/14fauna.pdf Review of Impacts of Displaced/Introduced Fauna Associated with Inland Waters.] {{webarchive |url=https://web.archive.org/web/20081201161912/http://www.environment.gov.au/soe/1996/publications/technical/pubs/14fauna.pdf |date=December 1, 2008 }}" ''[http://www.environment.gov.au/soe/1996/publications/technical/14fauna.html Environment Australia] {{webarchive |url=https://web.archive.org/web/20090425035311/http://www.environment.gov.au/soe/1996/publications/technical/14fauna.html |date=April 25, 2009 }}'' 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>{{FishBase genus | genus = Toxotes | month = December | year = 2017}}</ref><ref>{{Cite web |title=WoRMS – World Register of Marine Species – Toxotidae Bleeker, 1859 |url=https://www.marinespecies.org/aphia.php?p=taxdetails&id=151464 |access-date=2025-02-01 |website=www.marinespecies.org |language=en}}</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 {{convert|12-18|cm|in|abbr=on|0}}, but ''T. chatareus'' can reach {{convert|40|cm|in|abbr=on}}.<ref name=FishBaseGenus/><ref name=EoF>{{cite book |editor=Paxton, J.R. |editor2=Eschmeyer, W.N.|author1=Johnson, G.D. |author2=Gill, A.C.|year=1998|title=Encyclopedia of Fishes|publisher= Academic Press|location=San Diego|page= 189|isbn= 0-12-547665-5}}</ref>

Archerfish are popular exotic fish for aquaria,<ref>{{cite web|url=http://www.liveaquaria.com/product/prod_display.cfm?c=954+1600&pcatid=1600 |title=Archer Fish (''Toxotes jaculatrix'')|date=2016-03-09|publisher=liveaquaria|archive-date=2016-04-05|url-status=deviated|archive-url=https://web.archive.org/web/20160405064629/http://www.liveaquaria.com/product/prod_display.cfm?c=954+1600&pcatid=1600 }}</ref> but are difficult to feed and maintain by average fishkeepers since they prefer live prey over typical fish foods.

== Taxonomy == The taxonomic affiliation of the archerfish was previously uncertain, and they were often placed in the order Perciformes. More recently, phylogenetic studies have identified their closest relatives to be the beachsalmons in the family Leptobramidae, with the two comprising the suborder Toxotoidei within an expanded Carangiformes. The Leptobramidae have features in the oral cavity reminiscent of those that allow the Toxotidae to shoot water, affirming their relationship.<ref name=":0" /><ref name=":13">{{Cite web |last1=Fricke |first1=R. |last2=Eschmeyer |first2=W. N. |last3=Van der Laan |first3=R. |date=2025 |title=ESCHMEYER'S CATALOG OF FISHES: CLASSIFICATION |url=https://www.calacademy.org/eschmeyers-catalog-of-fishes-classification |access-date=2025-08-14 |website=California Academy of Sciences |language=en}}</ref><ref>{{Cite journal |last=Girard |first=Matthew G. |last2=Davis |first2=Matthew P. |last3=Smith |first3=W. Leo |date=2020-05-08 |title=The Phylogeny of Carangiform Fishes: Morphological and Genomic Investigations of a New Fish Clade |url=https://bioone.org/journals/copeia/volume-108/issue-2/CI-19-320/The-Phylogeny-of-Carangiform-Fishes--Morphological-and-Genomic-Investigations/10.1643/CI-19-320.full |journal=Copeia |volume=108 |issue=2 |pages=265 |doi=10.1643/CI-19-320 |issn=0045-8511}}</ref>

In the past, ''Toxotes'' was the only genus within the family. However, a 2022 study revived the genus ''Protoxotes'' due to its deep genetic divergence from ''Toxotes''.<ref name=":0" />

=== Species === [[Image:Toxotes Aquarium Liège 30012016 1.jpg|thumb|''Toxotes blythii'']]

There are 9 valid species in the genus ''Toxotes'':<ref name="CofFF" /><ref name="Kottelat2018" /><ref name=":0">{{Cite journal |last1=Girard |first1=M G |last2=Davis |first2=M P |last3=Tan |first3=H H |last4=Wedd |first4=D J |last5=Chakrabarty |first5=P |last6=Ludt |first6=W B |last7=Summers |first7=A P |last8=Smith |first8=W L |year=2022 |title=Phylogenetics of archerfishes (Toxotidae) and evolution of the toxotid shooting apparatus |journal=Integrative Organismal Biology |volume=4 |issue=1 |doi=10.1093/iob/obac013 |pmc=9259087 |pmid=35814192 |article-number=obac013}}</ref><ref>{{Cite web |title=WoRMS - World Register of Marine Species - Toxotidae Bleeker, 1859 |url=https://www.marinespecies.org/aphia.php?p=taxdetails&id=151464 |access-date=2025-02-01 |website=www.marinespecies.org |language=en}}</ref>

*''Protoxotes'' Whitley, 1950 **''Proxotes lorentzi'' (Weber, 1910) – primitive archerfish *''Toxotes'' Cloquet, 1816 ** ''Toxotes blythii'' Boulenger, 1892 – clouded archerfish, zebra archerfish ** ''Toxotes carpentariensis'' Castelnau, 1878<ref name=":0" /> ** ''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<ref name="Kottelat20183">{{cite journal |author1=Maurice Kottelat |author-link=Maurice Kottelat |author2=Tan Heok Hui |year=2018 |title=Three new species of archerfishes from the freshwaters of Southeast Asia (Teleostei: Toxotidae) and notes on Henri Mouhot's fish collections |journal=Ichthyological Exploration of Freshwaters |volume=IEF-952 |pages=1–19 |doi=10.23788/IEF-952}}</ref> A single fossil species, †''Toxotes beauforti'' Sanders, 1934 is known from the Eocene-aged Sangkarewang Formation of Sumatra, Indonesia. It very closely resembles modern archerfish species, aside from having six dorsal spines instead of the four or five present in modern species.<ref>{{Cite journal |last=Allen |first=Gerald R. |date=1978 |title=A REVIEW OF THE ARCHERFISHES (FAMILY TOXOTIDAE) |url=https://museum.wa.gov.au/research/records-supplements/records/review-archerfishes-family-toxotidae |journal=Rec. West. Aust. Mus. |volume=6 |issue=4}}</ref>

==Capture of prey== thumb|Illustration of an archerfish shooting water at a bug on a hanging branch|left thumb|Video of an archerfish shooting at prey 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>{{cite web|url=http://www98.griffith.edu.au/dspace/bitstream/10072/4430/1/29443.pdf |title=River and wetland food webs in Australia's wet-dry tropics: general principles and implications for management|author1=Douglas, M.M. |author2=Bunn, S.E. |author3=Davies, P.M. |name-list-style=amp|date=2005-06-03|publisher=Marine and Freshwater Research Vol. 56, No. 3, 329–342|access-date=2009-05-24}}</ref> up to {{convert|3|m|ft|abbr=on|0}} above the water's surface.<ref>{{cite web|url=https://www.telegraph.co.uk/news/uknews/1401098/Plastic-flies-help-spitting-archer-fish-regain-aim.html |title="Plastic flies help spitting archer fish regain aim" Telegraph.co.uk|date=2002-07-11|publisher=The Telegraph|access-date=2009-05-24}}</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">{{cite journal|url=https://www.nbb.cornell.edu/neurobio/department/reprints/Schuster%20et%20al.%202006.pdf |title=Animal Cognition:, How Archer Fish Learn to Down Rapidly Moving Targets |author1=Schuster, S. |author2=Wöhl, S. |author3=Griebsch, M. |author4=Klostermeier, I. |journal=Current Biology |name-list-style=amp|date=2006-02-21 |volume=16 |issue=4 |pages=378–383 |publisher=Current Biology Vol. 16, No. 4, 378–383 |doi=10.1016/j.cub.2005.12.037 |pmid=16488871 |bibcode=2006CBio...16..378S |s2cid=1139246 |access-date=2014-05-31 |archive-url=https://web.archive.org/web/20100614133100/http://www.nbb.cornell.edu/neurobio/department/reprints/Schuster%20et%20al.%202006.pdf |archive-date=2010-06-14 }}</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">{{cite journal | last1 = Milius | first1 = Susan | last2 = October | year = 2014 | title = Archerfish mouth reveals spit secret | journal = Science News | volume = 186 | issue = 7| page = 8 | doi=10.1002/scin.2014.186007005}}</ref><ref>{{cite magazine|url=https://www.wired.com/2013/11/spit-decision-how-archerfish-decide/|title=Spit Decision: How Archerfish Decide|date=19 November 2013|magazine=WIRED}}</ref> They are persistent and will make multiple shots if the first one fails.<ref>{{Cite journal|url = http://booksandjournals.brillonline.com/content/journals/10.1163/156854200x00162|title = Prey Catching in the Archer Fish: Marksmanship, and Endurance of Squirting At an Aerial Target|last = Timmermans|first = P.J.A|date = 2000|journal = Netherlands Journal of Zoology| volume=50 | issue=4 | pages=411–423 |access-date = 2014-10-06|doi = 10.1163/156854200X00162|url-access = subscription}}</ref>

Young archerfish start shooting when they are about {{convert|2.5|cm|in|abbr=on|0}} long but are inaccurate at first and must learn from experience.{{Citation needed|date=October 2020}} During this learning period, they hunt in small schools. This way, the probability is enhanced that at least one jet will hit its target.{{Citation needed|reason=This is rather speculative and is not shown in any primary literature as far as I am aware.|date=October 2020}} A 2006 experimental study found that archerfish appear to benefit from observational learning by watching a performing group member shoot, without having to practice: {{blockquote|This instance of social learning in a fish is most remarkable as it could imply that observers can ''change their viewpoint,'' mapping the perceived shooting characteristics of a distant team member into angles and target distances that they later must use to hit.<ref name="Schuster animal cognition" />}}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>{{cite web|url=https://www.newscientist.com/article/2118582-spitting-archerfish-shoot-at-prey-above-and-beneath-the-water/|title=New Scientist: Spitting archerfish shoot at prey above and beneath the water}}</ref> This makes it an example of exaptation.

==References== {{Reflist}}

==External links== * [http://educatedearth.net/video.php?id=3938 Archerfish Shooting Various Insects] * [https://web.archive.org/web/20150318000633/http://www.maniacworld.com/Archerfish-vs-fly.html Archerfish capturing prey] * [http://mangrove.nus.edu.sg/guidebooks/text/2105.htm Guide to the Mangroves of Singapore: Archerfish] * [http://www.naturia.per.sg/buloh/verts/archer_fish.htm Information and Photos on Archerfish] * [https://web.archive.org/web/20061010225403/http://www.livescience.com/animalworld/061009_archerfish.html Archerfish Can Adjust Their Deadly Shots Based on the Size of the Prey] – LiveScience.com

Category:Archerfish Category:Articles containing video clips Category:Brackish water organisms Category:Taxa named by Pieter Bleeker Category:Fish of Indonesia Category:Fish of Australia Category:Fish of the Pacific Ocean