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=== Blooms === {{Main|Jellyfish bloom}} [[File:Jellyfish population trends by LME.jpg|thumb|Map of population trends of native and invasive jellyfish.<ref name="brotz">{{cite journal |journal=Hydrobiologia |title=Increasing jellyfish populations: trends in Large Marine Ecosystems |year=2012 |volume=688 |author1=Brotz, Lucas |author2=Cheung, William W. L. |author3=Kleisner, Kristin |author4=Pakhomov, Evgeny |author5=Pauly, Daniel |display-authors=3|doi=10.1007/s10750-012-1039-7 |pages=3β20|doi-access=free }}</ref> <br />Circles represent data records; larger circles denote higher certainty of findings.<br /> {{legend|#E51A1D|Increase (high certainty)}}{{legend|#F1A341|Increase (low certainty)}} {{legend|#4DAF4A|Stable/variable}}{{legend|#377CB5|Decrease}}{{legend|#CCCCCA|No data}}]] Jellyfish form large masses or blooms in certain environmental conditions of [[ocean current]]s, [[nutrient]]s, sunshine, temperature, season, prey availability, reduced predation and [[oxygen]] concentration. Currents collect jellyfish together, especially in years with unusually high populations. Jellyfish can detect [[marine current]]s and swim against the current to congregate in blooms.<ref>{{cite news |last1=Gill |first1=Victoria |title=Jellyfish 'can sense ocean currents' |url=https://www.bbc.co.uk/news/science-environment-30936192 |access-date=26 January 2015 |agency=BBC News |url-status=live |archive-url=https://web.archive.org/web/20150126020804/http://www.bbc.co.uk/news/science-environment-30936192 |archive-date=26 January 2015}}</ref><ref>{{cite journal | last=Hays | first=Graeme C. | title=Ocean currents and marine life | journal=Current Biology | volume=27 | issue=11 | year=2017 | doi=10.1016/j.cub.2017.01.044 | pages=R470βR473|pmid=28586681| doi-access=free | bibcode=2017CBio...27.R470H | hdl=10536/DRO/DU:30100274 | hdl-access=free }}</ref> Jellyfish are better able to survive in nutrient-rich, oxygen-poor water than competitors, and thus can feast on plankton without competition. Jellyfish may also benefit from saltier waters, as saltier waters contain more [[iodine]], which is necessary for polyps to turn into jellyfish. Rising sea temperatures caused by [[climate change]] may also contribute to jellyfish blooms, because many species of jellyfish are able to survive in warmer waters.<ref>{{cite web |last=Shubin |first=Kristie |access-date =19 November 2009 |date =10 December 2008 |title =Anthropogenic Factors Associated with Jellyfish Blooms β Final Draft II |url=http://jrscience.wcp.muohio.edu/fieldcourses08/PapersMarineEcologyArticles/AnthropogenicFactorsAssocA.html |url-status=live |archive-url=https://web.archive.org/web/20100614184618/http://jrscience.wcp.muohio.edu/fieldcourses08/PapersMarineEcologyArticles/AnthropogenicFactorsAssocA.html |publisher= Tropical Field Courses: Western Program: Miami University|archive-date=14 June 2010 }}</ref> Increased nutrients from agricultural or urban [[Runoff (water)|runoff]] with nutrients including nitrogen and phosphorus compounds increase the growth of phytoplankton, causing [[eutrophication]] and [[algal bloom]]s. When the phytoplankton die, they may create [[Dead zone (ecology)|dead zones]], so-called because they are [[Hypoxia (environmental)|hypoxic]] (low in oxygen). This in turn kills fish and other animals, but not jellyfish,<ref>{{cite web|url=https://oceanservice.noaa.gov/facts/deadzone.html|title=What is a dead zone?|publisher=National Ocean Service|access-date=3 October 2018}}</ref> allowing them to bloom.<ref>{{cite web|url=http://blogs.discovermagazine.com/notrocketscience/2011/06/06/jellyfish-shift-ocean-food-webs-by-feeding-bacteria-with-mucus-and-excrement/|title=Jellyfish shift ocean food webs by feeding bacteria with mucus and excrement|last1=Yong|first1=Ed|date=6 June 2011|publisher=Discover Magazine|access-date=3 October 2018|archive-date=6 November 2018|archive-url=https://web.archive.org/web/20181106190642/http://blogs.discovermagazine.com/notrocketscience/2011/06/06/jellyfish-shift-ocean-food-webs-by-feeding-bacteria-with-mucus-and-excrement/|url-status=dead}}</ref><ref name="WaPost">{{Cite web|title=Jellyfish blooms could be sign of ailing seas|url=http://www.eurocbc.org/page727.html|access-date=2023-02-10|website=www.eurocbc.org}}</ref> Jellyfish populations may be expanding globally as a result of land runoff and [[overfishing]] of their [[predation|natural predators]].<ref>{{cite journal|last1=Hays|first1=G. C.|last2=Bastian|first2=T.|last3=Doyle|first3=T. K.|last4=Fossette|first4=S.|last5=Gleiss|first5=A. C.|last6=Gravenor|first6=M. B.|last7=Hobson|first7=V. J.|last8=Humphries|first8=N. E.|last9=Lilley|first9=M. K. S.|last10=Pade|first10=N. G.|last11=Sims|first11=D. W.|display-authors=3|year=2011|title=High activity and LΓ©vy searches: jellyfish can search the water column like fish|url=http://www.swan.ac.uk/bs/turtle/reprints/Hays_etal_PRSB_doi_2011.pdf|url-status=live|journal=Proceedings of the Royal Society B|volume=279|issue=1728|pages=465β473|doi=10.1098/rspb.2011.0978|pmc=3234559|pmid=21752825|archive-url=https://web.archive.org/web/20121225074206/http://www.swan.ac.uk/bs/turtle/reprints/Hays_etal_PRSB_doi_2011.pdf|archive-date=25 December 2012}}</ref><ref>{{cite journal|last1=Pauly|first1=D.|last2=Christensen|first2=V.|last3=Dalsgaard|first3=J.|last4=Froese|first4=R.|last5=Torres Jr|first5=F.|display-authors=3|year=1998|title=Fishing down marine food webs|url=http://umanitoba.ca/institutes/natural_resources/pdf/pauly_fishing_down_marine_food_webs.pdf|url-status=live|journal=Science|volume=279|issue=5352|pages=860β863|bibcode=1998Sci...279..860P|doi=10.1126/science.279.5352.860|pmid=9452385|archive-url=https://web.archive.org/web/20120710184954/http://umanitoba.ca/institutes/natural_resources/pdf/pauly_fishing_down_marine_food_webs.pdf|archive-date=10 July 2012}}</ref> Jellyfish are well placed to benefit from disturbance of marine ecosystems. They reproduce rapidly; they prey upon many species, while few species prey on them; and they feed via touch rather than visually, so they can feed effectively at night and in turbid waters.<ref>{{cite journal|last1=Richardson|first1=A. J.|last2=Bakun|first2=A.|last3=Hays|first3=G. C.|last4=Gibbons|first4=M. J.|year=2009|title=The jellyfish joyride: causes, consequences and management responses to a more gelatinous future|url=http://www.swan.ac.uk/bs/turtle/reprints/Richardson_et_al_2009_TREE_-_The_Jellyfish_Joyride.pdf|journal=Trends in Ecology & Evolution|volume=24|issue=6|pages=312β322|doi=10.1016/j.tree.2009.01.010|pmid=19324452|bibcode=2009TEcoE..24..312R }}{{Dead link|date=February 2023 |bot=InternetArchiveBot |fix-attempted=yes }}</ref><ref>{{cite journal|last1=Aksnes|first1=D. L.|last2=Nejstgaard|first2=J.|last3=SΓ¦dberg|first3=E.|last4=SΓΈrnes|first4=T.|year=2004|title=Optical control of fish and zooplankton populations|journal=Limnology and Oceanography|volume=49|issue=1|pages=233β238|bibcode=2004LimOc..49..233A|doi=10.4319/lo.2004.49.1.0233|doi-access=free}}</ref> It may be difficult for [[fish stocks]] to re-establish themselves in marine ecosystems once they have become dominated by jellyfish, because jellyfish feed on plankton, which includes [[fish eggs]] and [[fish larvae|larvae]].<ref>{{cite journal|last1=Lynam|first1=C. P.|last2=Gibbons|first2=M. J.|last3=Axelsen|first3=B. E.|last4=Sparks|first4=C. A. J.|last5=Coetzee|first5=J.|last6=Heywood|first6=B. G.|last7=Brierley|first7=A. S.|display-authors=3|year=2006|title=Jellyfish overtake fish in a heavily fished ecosystem|url=https://www.st-andrews.ac.uk/~perg/Lynam_et_al_Current_Biology_16_2006.pdf|url-status=live|journal=Current Biology|volume=16|issue=13|pages=492β493|doi=10.1016/j.cub.2006.06.018|pmid=16824906|s2cid=62793057|archive-url=https://web.archive.org/web/20110323010524/http://www.st-andrews.ac.uk/~perg/Lynam_et_al_Current_Biology_16_2006.pdf|archive-date=23 March 2011|doi-access=free|bibcode=2006CBio...16.R492L }}</ref><ref>{{cite journal|last1=Pauly|first1=D.|last2=Graham|first2=W.|last3=Libralato|first3=S.|last4=Morissette|first4=L.|last5=Palomares|first5=M. L. D.|display-authors=3|year=2009|title=Jellyfish in ecosystems, online databases, and ecosystem models|url=http://filaman.ifm-geomar.de/home/pages/JellyfishInEcosystems_publication.pdf|url-status=dead|journal=Hydrobiologia|volume=616|pages=67β85|doi=10.1007/s10750-008-9583-x|s2cid=12415790|archive-url=https://web.archive.org/web/20130712214856/http://www.uamshealth.com/?id=11935&sid=1|archive-date=12 July 2013}}</ref><ref name="WaPost" /> [[File:Aurelia aurita (auge24eu).jpg|thumb|[[Aurelia aurita|Moon jellyfishes]] can live in northern hemisphere seas,<ref>{{cite journal |last1=Dawson |first1=M. N. |first2=A. |last2=Sen Gupta |first3=M. H. |last3=England |year=2005 |title=Coupled biophysical global ocean model and molecular genetic analyses identify multiple introductions of cryptogenic species |journal=[[Proceedings of the National Academy of Sciences of the United States of America|Proc. Natl. Acad. Sci. USA]] |volume=102 |issue=34 |pages=11968β73 |doi=10.1073/pnas.0503811102 |pmid=16103373 |pmc=1189321|bibcode=2005PNAS..10211968D |doi-access=free }}</ref><ref>{{cite journal |last=Dawson |first=M. N. |year=2003 |title=Macro-morphological variation among cryptic species of the moon jellyfish, ''Aurelia'' (Cnidaria: Scyphozoa) |journal=[[Marine Biology (journal)|Marine Biology]] |volume=143 |issue=2 |pages=369β79 |doi=10.1007/s00227-003-1070-3 |bibcode=2003MarBi.143..369D |s2cid=189820003 }}</ref> such as the [[Baltic Sea]].<ref name="navandi" /><ref name="haaretz" />]] As suspected at the turn of this century,<ref>{{cite journal |last1=Briand |first1=Frederic |last2=Boero |first2=Ferdinando |title=Gelatinous zooplankton outbreaks - an overview on jellyfish blooms |journal=CIESM Monographs |date=2001 |volume=14 |page=5-17 |url=https://www.researchgate.net/publication/365366204}}</ref><ref>{{cite journal|last1=Mills|first1=C. E.|year=2001|title=Jellyfish blooms: are populations increasing globally in response to changing ocean conditions?|url=http://faculty.washington.edu/cemills/jellyblooms2001.pdf|url-status=live|journal=Hydrobiologia|volume=451|pages=55β68|doi=10.1023/A:1011888006302|s2cid=10927442|archive-url=https://web.archive.org/web/20160303171345/http://faculty.washington.edu/cemills/jellyblooms2001.pdf|archive-date=3 March 2016}}</ref> jellyfish blooms are increasing in frequency. Between 2013 and 2020 the [[Mediterranean Science Commission]] monitored on a weekly basis the frequency of such outbreaks in coastal waters from Morocco to the Black Sea, revealing a relatively high frequency of these blooms nearly all year round, with peaks observed from March to July and often again in the autumn. The blooms are caused by different jellyfish species, depending on their localisation within the Basin: one observes a clear dominance of ''[[Pelagia noctiluca]]'' and ''[[Velella velella]]'' outbreaks in the western Mediterranean, of ''[[Rhizostoma pulmo]]'' and ''[[Rhopilema nomadica]]'' outbreaks in the eastern Mediterranean, and of ''[[Aurelia aurita]]'' and ''[[Mnemiopsis leidyi]]'' outbreaks in the Black Sea.<ref>{{Cite web|title=CIESM GIS|url=https://www.ciesm.org/gis/JW/build/JellyBlooms.php|access-date=2023-02-10|website=www.ciesm.org}}</ref> Some jellyfish populations that have shown clear increases in the past few decades are [[invasive species]], newly arrived from other habitats: examples include the [[Black Sea]], [[Caspian Sea]], [[Baltic Sea]], central and eastern [[Mediterranean Sea|Mediterranean]], Hawaii, and tropical and subtropical parts of the West Atlantic (including the [[Caribbean Sea|Caribbean]], [[Gulf of Mexico]] and Brazil).<ref name="navandi">{{cite journal |author1=Abed-Navandi, D. |author2=Kikinger, R. |year=2007 |title=First record of the tropical scyphomedusa ''Phyllorhiza punctata'' von Lendenfeld, 1884 (Cnidaria: Rhizostomeae) in the Central Mediterranean Sea |journal=Aquatic Invasions |volume=2 |issue=4 |pages=391β394 |url=http://elnais.ath.hcmr.gr/PDF/Abel_Navanti_Phylorizapunctata.pdf |doi=10.3391/ai.2007.2.4.7 |url-status=live |archive-url=https://web.archive.org/web/20120917090718/http://elnais.ath.hcmr.gr/PDF/Abel_Navanti_Phylorizapunctata.pdf |archive-date=17 September 2012|doi-access=free }}</ref><ref name="haaretz">{{cite news |url=https://www.haaretz.com/1.5065394 |title=World's Most Invasive Jellyfish Spreading Along Israel Coast |author=Rinat, Zafrir |date=15 June 2009 |newspaper=Haaretz |access-date=13 October 2018}}</ref> Jellyfish blooms can have significant impact on community structure. Some carnivorous jellyfish species prey on zooplankton while others graze on primary producers.<ref>{{Cite journal|last1=Purcell|first1=J.|last2=Arai|first2=M.|date=2001|title=Interactions of pelagic cnidarians and ctenophores with fish: a review|journal=Hydrobiologia|volume=541|pages=27β44|doi=10.1007/978-94-010-0722-1|isbn=978-94-010-3835-5|s2cid=27615539|editor1-last=Purcell|editor1-first=J. E|editor2-last=Graham|editor2-first=W. M|editor3-last=Dumont|editor3-first=H. J}}</ref> Reductions in zooplankton and [[ichthyoplankton]] due to a jellyfish bloom can ripple through the trophic levels. High-density jellyfish populations can outcompete other predators and reduce fish recruitment.<ref>{{Cite journal|last1=Brodeur|first1=Richard D.|last2=Link|first2=Jason S.|last3=Smith|first3=B.E.|last4=Ford|first4=M.D.|last5=Kobayashi|first5=D.R.|last6=Jones|first6=T.T.|display-authors=3|date=2016|title=Ecological and Economic Consequences of Ignoring Jellyfish: A Plea for Increased Monitoring of Ecosystems|journal=Fisheries|volume=41|issue=11|pages=630β637|doi=10.1080/03632415.2016.1232964|bibcode=2016Fish...41..630B }}</ref> Increased grazing on primary producers by jellyfish can also interrupt energy transfer to higher trophic levels.<ref>{{Cite journal|last1=Ruzicka|first1=J.J.|last2=Brodeur|first2=R.D.|last3=Emmett|first3=R.L.|last4=Steele|first4=J.H.|last5=Zamon|first5=J.E.|last6=Morgan|first6=C.A.|last7=Thomas|first7=A.C.|last8=Wainwright|first8=T.C.|display-authors=3|date=2012|title=Interannual variability in the Northern California Current food web structure: changes in energy flow pathways and the role of forage fish, euphausiids, and jellyfish|journal=Progress in Oceanography|volume=102|pages=19β41|doi=10.1016/j.pocean.2012.02.002|bibcode=2012PrOce.102...19R}}</ref> During blooms, jellyfish significantly alter the nutrient availability in their environment. Blooms require large amounts of available organic nutrients in the water column to grow, limiting availability for other organisms.<ref name="Pitt-2009">{{Cite journal|last1=Pitt|first1=Kylie|last2=Welsh|first2=David|last3=Condon|first3=Robert|date=January 2009|title=Influence of jellyfish blooms on carbon, nitrogen and phosphorus cycling and plankton production|journal=Hydrobiologia|volume=616|pages=133β149|doi=10.1007/s10750-008-9584-9|s2cid=22838905}}</ref> Some jellyfish have a symbiotic relationship with single-celled dinoflagellates, allowing them to assimilate inorganic carbon, phosphorus, and nitrogen creating competition for phytoplankton.<ref name="Pitt-2009" /> Their large biomass makes them an important source of dissolved and particulate organic matter for microbial communities through excretion, mucus production, and decomposition.<ref name="brotz" /><ref>{{Cite journal|last1=Condon|first1=Robert H.|last2=Duarte|first2=Carlos M.|last3=Pitt|first3=Kylie A.|last4=Robinson|first4=Kelly L.|last5=Lucas|first5=Cathy H.|last6=Sutherland|first6=Kelly R.|last7=Mianzan|first7=Hermes W.|last8=Bogeberg|first8=Molly|last9=Purcell|first9=Jennifer E.|last10=Decker|first10=Mary Beth|last11=Uye|first11=Shin-ichi|display-authors=3|date=2013-01-15|title=Recurrent jellyfish blooms are a consequence of global oscillations|journal=Proceedings of the National Academy of Sciences|volume=110|issue=3|pages=1000β1005|doi=10.1073/pnas.1210920110|pmid=23277544|pmc=3549082|bibcode=2013PNAS..110.1000C|doi-access=free}}</ref> The microbes break down the organic matter into inorganic ammonium and phosphate. However, the low carbon availability shifts the process from production to respiration creating low oxygen areas making the dissolved inorganic nitrogen and phosphorus largely unavailable for primary production. These blooms have very real impacts on industries. Jellyfish can outcompete fish by utilizing open niches in over-fished fisheries.<ref>{{Cite journal|last1=Lynam|first1=Christopher P.|last2=Gibbons|first2=Mark J.|last3=Axelsen|first3=BjΓΈrn E.|last4=Sparks|first4=Conrad A. J.|last5=Coetzee|first5=Janet|last6=Heywood|first6=Benjamin G.|last7=Brierley|first7=Andrew S.|display-authors=3|date=2006-07-11|title=Jellyfish overtake fish in a heavily fished ecosystem|journal=Current Biology |volume=16|issue=13|pages=R492β493|doi=10.1016/j.cub.2006.06.018|pmid=16824906|s2cid=62793057|doi-access=free|bibcode=2006CBio...16.R492L }}</ref> Catch of jellyfish can strain fishing gear and lead to expenses relating to damaged gear. Power plants have been shut down due to jellyfish blocking the flow of cooling water.<ref>{{Cite journal|last1=Masilamani|first1=J|last2=Jesudoss|first2=K|last3=Kanavillil|first3=Nandakumar|last4=Satpathy|first4=K.K.|last5=Nair|first5=K|last6=Azariah|first6=J|display-authors=3|date=2000-09-10|title=Jellyfish ingress: A threat to the smooth operation of coastal power plants|url=https://www.researchgate.net/publication/236844536|journal=Current Science|volume=79|pages=567β569}}</ref> Blooms have also been harmful for tourism, causing a rise in stings and sometimes the closure of beaches.<ref name="Purcell-2007">{{Cite journal|last1=Purcell|first1=Jennifer E.|last2=Uye|first2=Shin-ichi|last3=Lo|first3=Wen-Tseng|date=2007-11-22|title=Anthropogenic causes of jellyfish blooms and their direct consequences for humans: a review|journal=Marine Ecology Progress Series|volume=350|pages=153β174|doi=10.3354/meps07093|bibcode=2007MEPS..350..153P|doi-access=free}}</ref> Jellyfish form a component of [[jelly-falls]], events where gelatinous [[zooplankton]] fall to the seafloor, providing food for the [[benthic]] organisms there.<ref>{{Cite journal |last1=Sweetman |first1=Andrew K. |last2=Smith |first2=Craig R. |last3=Dale |first3=Trine |last4=Jones |first4=Daniel O. B. |date=2014 |title=Rapid scavenging of jellyfish carcasses reveals the importance of gelatinous material to deep-sea food webs |journal=Proceedings of the Royal Society B: Biological Sciences |volume=281 |issue=1796 |page=20142210 |doi=10.1098/rspb.2014.2210 |pmid=25320167 |pmc=4213659}}</ref> In temperate and subpolar regions, jelly-falls usually follow immediately after a bloom.<ref>{{Cite journal |last1=Lebrato |first1=Mario |last2=Pahlow |first2=Markus |last3=Oschlies |first3=Andreas |last4=Pitt |first4=Kylie A. |last5=Jones |first5=Daniel O. B. |last6=Molinero |first6=Juan Carlos |last7=Condon |first7=Robert H. |display-authors=3 |date=2011 |title=Depth attenuation of organic matter export associated with jelly falls |journal=Limnology and Oceanography |volume=56 |issue=5 |pages=1917β1928 |doi=10.4319/lo.2011.56.5.1917|url=http://oceanrep.geomar.de/12648/1/1917.pdf |bibcode=2011LimOc..56.1917L |hdl=10072/43275 |s2cid=3693276 |hdl-access=free }}</ref>
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