View source for Stickleback

{{Short description|Family of ray-finned fish}}
{{Other uses}}
{{Automatic taxobox
| name = Sticklebacks
| image = Four species of stickleback (Gasterosteidae).jpg
| image_caption = Four marine species of stickleback from the [[Atlantic Ocean]] coast of North America
| image_upright = 1.1
| taxon = Gasterosteidae
| authority = [[Charles Lucien Bonaparte|Bonaparte]], 1831<ref name = VDLEF>{{cite journal | author1 = Richard van der Laan | author2 = William N. Eschmeyer | author3 = Ronald Fricke | name-list-style = amp | year = 2014 | title = Family-group names of Recent fishes | url = https://www.researchgate.net/publication/268078514 | journal = Zootaxa | volume = 3882 | issue = 2 | pages = 001–230 | doi = 10.11646/zootaxa.3882.1.1 | pmid = 25543675 | doi-access = free }}</ref>
| subdivision_ranks = Genera
| subdivision = see text
}}

The '''sticklebacks''' are a [[family (biology)|family]] of [[ray-finned fish]]es, the '''Gasterosteidae''' which have a [[Holarctic realm|Holarctic]] distribution in fresh, brackish and marine waters.  They were thought to be related to the [[pipefish]] and [[seahorse]]s but are now thought to be more closely related to the [[eelpout]]s and [[sculpin]]s.

==Taxonomy==
The stickleback family, Gasterosteidae, was first proposed as a family by the French [[zoologist]] [[Charles Lucien Bonaparte]] in 1831.<ref name = VDLEF/> It was long thought that the sticklebacks and their relatives comprise a suborder, the [[Gasterosteoidei]], of the [[Order (biology)|order]] Gasterostiformes with the sea horses and pipefishes making up the suborder Syngnathoidei. More recent phylogenetic work has shown that the Gaterosteoidei are more closely related to the [[Zoarcoidei]] and the [[Cottoidei]], which would mean that this taxon would belong in the order [[Scorpaeniformes]].<ref name = Nelson5>{{cite book  |title=Fishes of the World |edition=5th |author1=J. S. Nelson |author2=T. C. Grande |author3=M. V. H. Wilson |year=2016 |pages= 467–495 |publisher=Wiley |isbn= 978-1-118-34233-6 |url=https://sites.google.com/site/fotw5th/ }}</ref> However, in other phylogenetic classifications it is treated as the infraorder Gasterosteales within the suborder [[Cottoidei]] or as a sister clade to the [[Zoarcoidei|Zoarcales]] in the order Zoarciformes.<ref name = Deepfin>{{cite journal | author1 = Ricardo Betancur-R | author2 = Edward O. Wiley | author3 = Gloria Arratia | author4 = Arturo Acero | author5 = Nicolas Bailly | author6 = Masaki Miya | author7 = Guillaume Lecointre | author8 = Guillermo Ortí | display-authors = 3 | title =Phylogenetic classification of bony fishes | journal = BMC Evolutionary Biology | volume = 17 | issue = 162 | year = 2017 | page = 162 | doi = 10.1186/s12862-017-0958-3 | pmid = 28683774 | pmc = 5501477 | doi-access = free | bibcode = 2017BMCEE..17..162B }}</ref>

[[FishBase]] recognises 16 species in the family, grouped in five genera.<ref name = Fishbase>{{FishBase family | family = Gasterosteidae | month = October| year = 2012}}</ref> However, several of the species have a number of recognised subspecies, and the taxonomy of the family is thought to be in need of revision.

===Genera===
The family Gasterosteidae includes the following genera:<ref name = Fishbase/>

* ''[[Apeltes]]'' <small>[[James Ellsworth De Kay|DeKay]], 1842</small>
* ''[[Culaea]]'' <small>[[Gilbert Percy Whitley|Whitley]], 1950</small>
* ''[[Gasterosteus]]'' <small>[[Carl Linnaeus|Linnaeus]], 1758</small>
* ''[[Pungitius]]'' <small>[[Johann Jakob d’Annone|d'Annone]], 1760</small>
* ''[[Spinachia]]'' <small>[[Georges Cuvier|Cuvier]], 1816</small>

==Description==
[[File:Faroe stamp 248 stickleback (gasterosteus aculeatus).gif|thumb|upright=1.1|1994 [[Faroe Islands]] postage stamp with [[three-spined stickleback]]s]]

Sticklebacks are endemic to the temperate zone<ref name=":0">{{Cite book |last1=Foster|first1=Susan A. |last2=Cresko|first2=William A.|last3=Johnson|first3=Kevin P.|last4=Tlusty|first4=Michael U.|last5=Willmott|first5=Harleigh E. |date=1996-01-01 |chapter-url=http://www.sciencedirect.com/science/article/pii/B9780126180305500125 |title=Homoplasy|pages=245–269|editor-last=Sanderson|editor-first=Michael J.|publisher=Academic Press|language=en|doi=10.1016/b978-012618030-5/50012-5|isbn=978-0-12-618030-5|access-date=2020-06-04 |chapter=Patterns of homoplasy in behavioral evolution|editor2-last=Hufford|editor2-first=Larry}}</ref> and are most commonly found in the ocean, but some can be found in fresh water. The freshwater [[taxon|taxa]] were trapped in [[Europe]], [[Asia]], and [[North America]] after the [[Last Glacial Period|Ice Age]] 10,000–20,000 years ago, and have evolved features different from those of the marine species.<ref>{{Cite journal|last=Münzing|first=Joachim|date=1963|title=The evolution of variation and distributional patterns in European populations of the three-spined stickleback, ''Gasterosteus aculeatus'' |url=https://www.jstor.org/stable/2406161|journal=Evolution|volume=17|issue=3|pages=320–332|doi=10.2307/2406161|jstor=2406161 |url-access=subscription}}</ref><ref>{{cite journal |last1=Bell |first1=Michael A. |title=Evolution of phenotypic diversity in ''Gasterosteus aculeatus'' superspecies on the Pacific Coast of North America |journal=Systematic Zoology |date=1976 |volume=25 |issue=3 |pages=211–227 |doi=10.2307/2412489|jstor=2412489 }}</ref>

Sticklebacks are carnivorous, feeding on small animals such as insects, crustaceans and fish larvae.<ref>The Repeater - NYTimes.com</ref><ref name=EoF/>

Sticklebacks are characterised by the presence of strong and clearly isolated spines in their [[dorsal fin]]s.<ref>{{cite journal |last1=Colosimo |first1=Pamela F. |last2=Hosemann |first2=Kim E. |last3=Balabhadra |first3=Sarita |last4=Villarreal |first4=Guadalupe |last5=Dickson |first5=Mark |last6=Grimwood |first6=Jane |last7=Schmutz |first7=Jeremy |last8=Myers |first8=Richard M. |last9=Schluter |first9=Dolph |last10=Kingsley |first10=David M. |title=Widespread parallel evolution in sticklebacks by repeated fixation of ectodysplasin alleles |journal=Science |date=2005 |volume=307 |issue=5717 |pages=1928–33 |doi=10.1126/science.1107239|pmid=15790847 |bibcode=2005Sci...307.1928C |s2cid=1296135 }}</ref> An unusual feature of sticklebacks is that they have no scales, although some species have bony armour plates.

===Sizes===
The maximum size of the best-known species, the [[three-spined stickleback]] (''Gasterosteus aculeatus''), is about 4 inches, but few of them are more than 3 inches long. They mature sexually at a length of about 2 inches.<ref name="GMA">{{cite web|url=http://www.gma.org/fogm/Gasterosteus_aculeatus.htm |title=Three-spined stickleback |publisher=Gma.org |access-date=2012-08-31}}</ref> Most other stickleback species are roughly similar in size or somewhat smaller. The only exception is the far larger [[fifteen-spined stickleback]] (''Spinachia spinachia''), which can reach 22&nbsp;cm (approx. 8.8 inches).<ref>{{FishBase|Spinachia|spinachia|year=2014}}</ref> Body form varies with habitat: sticklebacks in shallow lakes have developed a deep body specialized to enable feeding on benthic invertebrates, whilst those in deep oligotrophic lakes have adapted to feed on plankton and have slimmer bodies.<ref name=":0" />

===Personality===
Research has shown that Sticklebacks display distinct personality traits, specifically in the area of taking a risk, and,  can be considered bold or shy. These personality traits were determined to directly influence if they would lead, and if discouraged, attempt to lead again.<ref>{{Cite web|title=Stickleback fish show initiative, personality and leadership|url=https://phys.org/news/2012-08-stickleback-fish-personality-leadership.html|access-date=2020-06-26|website=phys.org|language=en}}</ref>

==Mating==
All stickleback species show similar, unusual mating behaviour. Freshwater males develop a red colouration, and although this may be seen in oceanic and benthic species these tend to remain dull-coloured. The male then constructs a nest from weeds held together by spiggin,<ref name=":0" /> a kidney secretion, then attract females to the nest. Females signal their readiness to mate with solitary rather than shoaling behaviour, a head-up posture; their bellies are also obviously distended with eggs.<ref name=":0" /> Courtship typically involves a zig-zag 'dance' where the male approaches the female in an erratic side-to-side pattern, and dorsal pricking of the female's abdomen.<ref name=":0" />  A female lays her eggs inside the nest, where the male [[fertilise]]s them. The male then guards the eggs until they hatch 7–14 days later (depending on temperature),<ref name=":0" /><ref name=EoF>{{cite book |editor=Paxton, J.R. |editor2=Eschmeyer, W.N.|author1=Orr, James W.  |author2=Pietsch, T.W. |name-list-style=amp |year=1998|title=Encyclopedia of Fishes|publisher= Academic Press|location=San Diego|pages= 171–172|isbn= 0-12-547665-5}}</ref> and may continue to guard the fry after they hatch. This large investment in both the nesting site and guarding of the eggs limits the number of females a male can mate with however males spawn multiple times.<ref name=":0" /> This introduces the ability for selection to favor male [[mate choice]].<ref>{{Cite journal|last1=Sargent|first1=Robert Craig|last2=Gross|first2=Mart R.|last3=Van Den Berghe|first3=Eric P.|date=1986-04-01|title=Male mate choice in fishes|journal=Animal Behaviour|language=en|volume=34|issue=2|pages=545–550|doi=10.1016/S0003-3472(86)80123-3|s2cid=53183473|issn=0003-3472}}</ref> Some males die following spawning.<ref name="GMA" />

=== Mating choice ===
Typically, the sex with the greatest parental investment has the strongest mate preferences.<ref>{{Cite journal|last1=Balshine|first1=S.|last2=Kempenaers|first2=B.|last3=Székely|first3=T.|last4=Kokko|first4=H.|last5=Johnstone|first5=R. A.|date=2002-03-29|title=Why is mutual mate choice not the norm? Operational sex ratios, sex roles and the evolution of sexually dimorphic and monomorphic signalling|journal=Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences|volume=357|issue=1419|pages=319–330|doi=10.1098/rstb.2001.0926|pmc=1692955|pmid=11958700}}</ref> Stickleback species exhibit mutual mate choice in which both the male and female have strong mate preferences. This is due in part to the strong parental investment on behalf of the male in guarding the eggs.<ref name="Kraak 859–866">{{Cite journal|last1=Kraak|first1=Sarah B. M|last2=Bakker|first2=Theo C. M. |date=1998-10-01|title=Mutual mate choice in sticklebacks: attractive males choose big females, which lay big eggs|url=http://www.sciencedirect.com/science/article/pii/S0003347298908223|journal=Animal Behaviour|language=en|volume=56|issue=4|pages=859–866|doi=10.1006/anbe.1998.0822|pmid=9790696|s2cid=21298270 |url-access=subscription}}</ref>

=== Female mate choice ===
Female sticklebacks show a strong preference to male stickleback with bright red coloration under their throats. Females mate both more often with males with brighter red coloration and give on average, larger eggs to be fertilized by these males. This preference has led to brighter red coloring.<ref>{{Cite journal|last1=Barber|first1=Iain|last2=Arnott|first2=Stephen A.|last3=Braithwaite|first3=Victoria A.|last4=Andrew|first4=Jennifer|last5=Huntingford|first5=Felicity A.|date=2001-01-07|title=Indirect fitness consequences of mate choice in sticklebacks: offspring of brighter males grow slowly but resist parasitic infections|journal=Proceedings of the Royal Society of London. Series B: Biological Sciences|volume=268|issue=1462|pages=71–76|doi=10.1098/rspb.2000.1331|pmc=1087602|pmid=12123300}}</ref><ref>{{Cite journal|last1=Bakker|first1=Theo C. M.|last2=Mundwiler|first2=Beat|date=1994-03-01|title=Female mate choice and male red coloration in a natural three-spined stickleback (''Gasterosteus aculeatus'') population|url=https://academic.oup.com/beheco/article/5/1/74/322603|journal=Behavioral Ecology|language=en|volume=5|issue=1|pages=74–80|doi=10.1093/beheco/5.1.74 }}</ref> This association is possible because the red coloration can only be produced by males that are free of parasites. This is referred to in the Hamilton-Zuk hypothesis.<ref>{{Cite journal|last1=Milinski|first1=Manfred|last2=Bakker|first2=Theo C. M.|date=March 1990|title=Female sticklebacks use male coloration in mate choice and hence avoid parasitized males|url=https://www.nature.com/articles/344330a0 |journal=Nature|language=en|volume=344|issue=6264|pages=330–333|doi=10.1038/344330a0|bibcode=1990Natur.344..330M|s2cid=4322443 |url-access=subscription}}</ref>

However, there is also evidence that attractive male red coloration may be a faulty signal of male quality.  Male sticklebacks that are more attractive to females due to carotenoid colorants may under-allocate carotenoids to their germline cells.<ref name = Kim2020>{{cite journal |vauthors=Kim SY, Velando A |title=Attractive male sticklebacks carry more oxidative DNA damage in the soma and germline |journal=J Evol Biol |volume=33 |issue=1 |pages=121–6 |date=January 2020 |pmid=31610052 |doi=10.1111/jeb.13552 }}</ref>   Since carotinoids are beneficial antioxidants, their under-allocation to [[germline]] cells can lead to increased oxidative [[DNA damage (naturally occurring)|DNA damage]] to these cells.<ref name = Kim2020/>  Therefore, female sticklebacks may risk [[fertility]] and the viability of their offspring by choosing redder, but more deteriorated partners with reduced [[sperm]] quality.

Female mate choice has also been seen to be condition dependent. Females are almost always the more choosy sex in most species. Female sticklebacks though, have been found to be less choosy of mates when in poor physical condition and inversely, more choosy in good condition.<ref>{{Cite journal|last1=Bakker|first1=Theo C. M.|last2=Künzler|first2=Reto|last3=Mazzi|first3=Dominique|date=September 1999|title=Condition-related mate choice in sticklebacks|journal=Nature|language=en|volume=401|issue=6750|pages=234|doi=10.1038/45727|s2cid=205043554 |doi-access=free}}</ref>

=== Male mate choice ===
In some species, such as the three-spined stickleback, the large investment in both nesting site and guarding of eggs by males limits the number of females a male can mate with.<ref>{{Cite book|title=Encyclopedia of fishes|date=1998|publisher=Academic Press |editor=Paxton, J.R. |editor2=Eschmeyer, W.N.|isbn=0-12-547665-5|edition=2nd|location=San Diego, CA|oclc=39641701}}</ref> This introduces the ability for selection to favor male mate choice. Male mate choice is rarely studied or observed in many species but multiple studies have confirmed male mate choice within stickleback species. Males show a choosiness similar to females as to what female they are willing to court and mate. Male sticklebacks have been observed to show preference towards female sticklebacks that are larger and longer. This is believed to be because larger females on average produce larger eggs, which leads to a greater offspring survival and fitness.<ref name="Kraak 859–866"/> In addition, male sticklebacks have also been observed to prefer females with more distended or bloated stomachs. The benefits of this is also due to larger eggs and thus offspring survival and fitness<ref>{{Cite journal|last=Rowland|first=William J.|date=1982-11-01|title=Mate choice by male sticklebacks, ''Gasterosteus aculeatus'' |url=http://www.sciencedirect.com/science/article/pii/S0003347282801991|journal=Animal Behaviour|language=en|volume=30|issue=4|pages=1093–1098|doi=10.1016/S0003-3472(82)80199-1|s2cid=54356731 |url-access=subscription}}</ref>

===Inbreeding avoidance===

Female three-spined sticklebacks adjust their courting behaviour to the risk of [[inbreeding]].<ref name = Frommen2006>{{cite journal |vauthors=Frommen JG, Bakker TC |title=Inbreeding avoidance through non-random mating in sticklebacks |journal=Biol Lett |volume=2 |issue=2 |pages=232–5 |date=June 2006 |pmid=17148370 |pmc=1618905 |doi=10.1098/rsbl.2005.0432 }}</ref>  When gravid females are given the choice between a courting unfamiliar non-sibling and a familiar brother, they prefer to mate with the non-sibling and thus avoid the disadvantages that accompany [[incest]].<ref name = Frommen2006/>  Eggs from inbred matings compared to eggs from outbred matings have a lower rate of fertilization and hatching, and fewer progeny survive to reproductive age.<ref name = Frommen2006/>

==Use in science==
[[Niko Tinbergen]]'s studies of the [[behaviour]] of this fish were important in the early development of [[ethology]] as an example of a [[fixed action pattern]]. More recently, the fish have become a favourite system for studying the molecular genetics of evolutionary change in wild populations<ref>{{cite book |last1=Kingsley |first1=D.M. |last2=Peichel |first2=C.L. |chapter=The molecular genetics of evolutionary change in sticklebacks |chapter-url=https://www.taylorfrancis.com/chapters/mono/10.1201/9781420004830-6/molecular-genetics-evolutionary-change-sticklebacks-sara-ostlund-nilsson-ian-mayer-felicity-anne-huntingford |editor-last=Ostlund-Nillson |editor-first=S. |editor2-last=Mayer |editor2-first=I. |editor3-last=Huntingford |editor3-first=F.A. |title=Biology of the three-spine stickleback |publisher=CRC Press |location= |date=2007 |isbn=978-1-4200-0483-0 |pages=41–81 |doi=10.1201/9781420004830-6 }}</ref> and a powerful  "supermodel" for combining evolutionary studies at molecular, developmental, population genetic, and ecological levels.<ref>{{cite journal|url=https://www.science.org/doi/full/10.1126/science.1109835 |title=The synthesis and evolution of a supermodel |journal=[[Science (journal)|Science]] |publisher=AAAS |date=2005-03-25 |doi=10.1126/science.1109835 |access-date=2012-08-31|last1=Gibson |first1=Greg |volume=307 |issue=5717 |pages=1890–1 |pmid=15790836 |s2cid=82257399 |url-access=subscription }}</ref> The nearly complete genome sequence of a reference freshwater stickleback was described in 2012, along with set of genetic variants commonly found in 21 marine and freshwater populations around the world. Some variants, and several chromosome inversions, consistently distinguish marine and freshwater populations, helping identify a genome-wide set of changes contributing to repeated adaptation of sticklebacks to marine and freshwater environments.<ref>{{cite journal|title=The genomic basis of adaptive evolution in threespine sticklebacks |date=2012-04-04 | volume=484|issue=7392 |doi=10.1038/nature10944|pmid=22481358 |journal=Nature|pages=55–61|pmc=3322419|last1=Jones |first1=Felicity C. |last2=Grabherr |first2=Manfred G. |last3=Chan |first3=Yingguang Frank |last4=Russell |first4=Pamela |last5=Mauceli |first5=Evan |last6=Johnson |first6=Jeremy |last7=Swofford |first7=Ross |last8=Pirun |first8=Mono |last9=Zody |first9=Michael C. |last10=White |first10=Simon |last11=Birney |first11=Ewan |last12=Searle |first12=Stephen |last13=Schmutz |first13=Jeremy |last14=Grimwood |first14=Jane |last15=Dickson |first15=Mark C. |last16=Myers |first16=Richard M. |last17=Miller |first17=Craig T. |last18=Summers |first18=Brian R. |last19=Knecht |first19=Anne K. |last20=Brady |first20=Shannon D. |last21=Zhang |first21=Haili |last22=Pollen |first22=Alex A. |last23=Howes |first23=Timothy |last24=Amemiya |first24=Chris |last25=Lander |first25=Eric S. |last26=Di Palma |first26=Federica |last27=Lindblad-Toh |first27=Kerstin |last28=Kingsley |first28=David M. |last29=Kingsley |first29=D. M. |bibcode=2012Natur.484...55. }}</ref> The adaptations seen in oceanic threespine sticklebacks make them an ideal organism for the study of parallel evolution.<ref>{{Cite journal |last1=Reid |first1=Kerry |last2=Bell |first2=Michael A. |last3=Veeramah |first3=Krishna R. |date=2021-08-31 |title=Threespine Stickleback: A Model System For Evolutionary Genomics |journal=Annual Review of Genomics and Human Genetics |volume=22 |pages=357–383 |doi=10.1146/annurev-genom-111720-081402 |pmc=8415275 |pmid=33909459}}</ref>

==In culture==
[[File:Monument blockaded fish kolushka.JPG|thumb|{{ill|Monument to the siege stickleback|ru|Памятник блокадной колюшке}}]]
There is a sculpture in [[Kronstadt]] dedicated to stickleback, which saved thousands of city residents from starvation during the [[Leningrad Siege]] of [[World War II]].<ref>{{Cite web |last=hecksinductionhour |date=2022-02-01 |title=Monument to a Tiny Fish |url=https://therussianreader.com/2022/02/01/stickleback-kronstadt-mikov/ |access-date=2025-03-09 |website=The Russian Reader |language=en-US}}</ref>

==References==
{{Reflist}}

==External links==
{{Commons category|Gasterosteidae}}
{{Wiktionary}}

*{{UCSC genomes|gasAcu1}}
*{{Cite EB1911|wstitle=Stickleback}}

{{Taxonbar|from=Q215420}}
{{Authority control}}

[[Category:Gasterosteidae|*]]