View source for Neuston

{{Short description|Organisms that live at the surface of a body of water}}
[[File:Wasserläufer bei der Paarung crop.jpg|right|thumb|upright=1.3|{{center|The [[water strider]], a common freshwater neuston}}]]

'''Neuston''', also called '''pleuston''', are [[organism]]s that live at the [[ocean surface ecosystem|surface]] of a [[body of water]], such as an [[ocean]], [[estuary]], [[lake]], [[river]], [[wetland]] or [[pond]]. Neuston can live on top of the water surface or submersed just below the water surface. In addition, [[Marine microorganism|microorganisms]] can exist in the [[sea surface microlayer|surface microlayer]] that forms between the top- and the under-side of the water surface. Neuston has been defined as "organisms living at the air/water interface of freshwater, estuarine, and marine habitats or referring to the biota on or directly below the water’s surface layer."<ref name= Kennish2016>{{cite book | editor-last=Kennish | editor-first=Michael J. | title=Encyclopedia of Earth Sciences Series | chapter=Encyclopedia of Estuaries | publisher=Springer Netherlands | publication-place=Dordrecht | year=2016 | isbn=978-94-017-8800-7 | issn=1388-4360 | doi=10.1007/978-94-017-8801-4 | s2cid=129770661 | ref={{sfnref | Springer Netherlands | 2016}}}}</ref>

Neustons can be informally separated into two groups: the '''phytoneuston''', which are [[autotroph]]s floating at the water surface including [[cyanobacteria]], filamentous [[algae]] and [[aquatic plant#Free-floating|free-floating aquatic plant]] (e.g. [[mosquito fern]], [[duckweed]] and [[water lettuce]]); and the '''zooneuston''', which are floating [[heterotroph]]s such as [[protist]]s (e.g. [[ciliate]]s) and [[metazoan]]s ([[aquatic animal]]s). This article mainly concerns with metazoan zooneustons.

The word "neuston" comes from [[Greek language|Greek]] ''neustos'', meaning "swimming", and the [[noun]] [[suffix]] ''-on'' (as in "[[plankton]]").<ref>Merriam-Webster Dictionary: [https://www.merriam-webster.com/dictionary/neuston neuston]. Accessed 18 December 2021.</ref> This term first appears in the biological literature in 1917.<ref name=Naumann2017 /> The alternative term pleuston comes from the Greek ''plein'', meaning "to sail or float". The first known use of this word was in 1909, before the first known use of neuston.<ref>Merriam-Webster Dictionary: [https://www.merriam-webster.com/dictionary/pleuston pleuston]. Accessed 18 December 2021.</ref> In the past various authors have attempted distinctions between neuston and pleuston, but these distinctions have not been widely adopted. As of 2021, the two terms are usually used somewhat interchangeably, and neuston is used more often than pleuston.

==Overview==
[[File:Portuguese Man-O-War (Physalia physalis).jpg|thumb|upright=1.2|left| {{center|[[Portuguese man o' war]]<br />emblematic figure of the marine pleuston}}]]

[[File:Neuston, Plankton, Nekton, Benthos.jpg|thumb|upright=1.3| {{center|Marine neuston (organisms that live at the ocean surface) can be contrasted with [[plankton]] (organisms that drift with water currents), [[nekton]] (organisms that can swim against water currents) and [[benthos]] (organisms that live at the ocean floor)}}]]

The neuston of the surface layer is one of the lesser known aquatic ecological groups.<ref name=Albuquerque2021 /> The term was first used in 1917 by Naumann to describe species associated with the surface layer of freshwater habitats.<ref name=Naumann2017>Naumann, E. (1917) "Beiträge zur Kenntnis des Teichnannoplanktons. II. Über das Neuston des Süsswassers", ''Biologisches Zentralblatt'', '''37''': 98–106.</ref>  Later in 1971, Zaitsev identified neuston composition in marine waters.<ref name= Zaitsev1971>Zaitsev, Y. P. (1971) "Marine Neustonology". ''National Marine Fisheries Service'', ''NOAA'' and ''NSF'', Washington DC.</ref> These populations would include microscopic species, plus various plant and animal taxa, such as [[phytoplankton]] and [[zooplankton]], living in this region.<ref name= Zaitsev1971 /><ref name= Marshall2005 /> In 2002, Gladyshev further characterised the major physical and chemical dynamics of the surface layer influencing the composition and relationships with various neustonic populations"<ref name= Gladyshev2002>{{cite book | last=Gladyshev | first=Michail | title=Biophysics of the surface microlayer of aquatic ecosystems | publisher=IWA | publication-place=London | url=https://books.google.com/books?id=rG8KCoOm-TwC | year=2002 | isbn=978-1-900222-17-4 | oclc=49871862}}</ref><ref name= Marshall2005 />

The neustonic community structure is conditioned by sunlight and an array of [[endogenous]] (organic matter, respiratory, photosynthetic, decompositional processes) and [[exogenous]] (atmospheric deposition, inorganic matter, winds, wave action, precipitation, UV radiation, oceanic currents, surface temperature) variables and processes affecting nutrient inputs and recycling.<ref name=Marshall2005 /><ref>{{cite journal |doi = 10.3318/bioe.2005.105.2.107|title = Temporal Variation in Diversity and Community Structure of a Semi-Isolated Neuston Community|year = 2005|last1 = Barnes|first1 = D. K.A.|last2 = Davenport|first2 = J.|last3 = Rawlinson|first3 = K. A.|journal = Biology and Environment: Proceedings of the Royal Irish Academy|volume = 105|issue = 2|pages = 107–122| s2cid=84508946 }}</ref><ref name=Rezai2019>{{cite journal |doi = 10.1016/j.rsma.2018.100473|title = Neustonic zooplankton in the northeastern Persian Gulf|year = 2019|last1 = Rezai|first1 = Hamid|last2 = Kabiri|first2 = Keivan|last3 = Arbi|first3 = Iman|last4 = Amini|first4 = Nafiseh|journal = Regional Studies in Marine Science|volume = 26|page = 100473| bibcode=2019RSMS...2600473R |s2cid = 135269465}}</ref> Furthermore, the neuston provides a food source to the [[zooplankton]] migrating from deeper layers to the surface,<ref name=Hempel1972>Hempel, G. and Weikert, H. (1972) "The neuston of the subtropical and boreal North-eastern Atlantic Ocean. A review". ''Marine Biology'', '''13'''(1): 70–88.</ref> as well as to [[seabird]]s roaming over the oceans.<ref>Cheng, L., Spear, L. and AINLEY, D.G. (2010) "Importance of marine insects (Heteroptera: Gerridae, Halobates spp.) as prey of eastern tropical Pacific seabirds". ''Marine Ornithology'', '''38'''": 91–95.</ref> For these reasons, the neustonic community is believed to play a critical role on the structure and function of [[marine food web]]s. Yet, research on neuston communities to date focused predominantly on geographically limited regions of the ocean{{hsp}}<ref>Zaitsev, Y. P. (1971). Marine Neustonology. ed. K. A. Vinogradov (Jerulasem: Israel program for scientific translations).</ref><ref name=Hempel1972 /><ref>{{cite journal |doi = 10.1007/BF00393027|title = A comparative survey of neuston: Geographical and temporal distribution patterns|year = 1983|last1 = Holdway|first1 = P.|last2 = Maddock|first2 = L.|journal = Marine Biology|volume = 76|issue = 3|pages = 263–270| bibcode=1983MarBi..76..263H |s2cid = 84587026}}</ref><ref>Ebberts, B. D., and Wing, B. L. (1997). "Diversity and abundance of neustonic zooplankton in the North Pacific subarctic frontal zone". U.S. Department of Commerce, NOAA Technical Memorandum NMFS-AFSC-70.</ref><ref name=Rezai2019 /> or coastal areas.<ref>{{cite journal |doi = 10.1139/z89-284|title = Neustonic feeding by juvenile salmonids in coastal waters of the Northeast Pacific|year = 1989|last1 = Brodeur|first1 = Richard D.|journal = Canadian Journal of Zoology|volume = 67|issue = 8|pages = 1995–2007}}</ref><ref>{{cite journal |doi = 10.3354/meps075185|title = Neustonic niche for cirripede larvae as a possible adaptation to long-range dispersal|year = 1991|last1 = Le Fèvre|first1 = J.|last2 = Bourget|first2 = E.|journal = Marine Ecology Progress Series|volume = 75|pages = 185–194|bibcode = 1991MEPS...75..185L}}</ref><ref>Padmavati, G. and Goswami, S.C. (1996). "Zooplankton distribution in neuston and water column along west coast of India from Goa to Gujarat". ''Indian J. Mar. Species'', '''25''': 85–90.</ref> Consequently, neuston complexity is still poorly understood as studies on the community structure and the taxonomical composition of organisms inhabiting this ecological niche remain few,<ref name=Rezai2019 /> and global scale analyses are yet lacking.<ref name=Albuquerque2021 />

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==Types==
[[File:Fis01007 (27889419550).jpg|thumb|upright=1.3| {{center|Neuston net}}]]

There are different ways neuston can be categorised. Kennish divides them by their physical position into two groups:<ref name= Kennish2016 /> 
* ''epineuston'': organisms living on the water's surface
* ''hyponeuston'': organisms within a region of specified depth directly below the surface layer 
To this can be added the organisms living in the microlayer at the interface between air and water:
*  ''microlayer neuston'': organisms (microorganisms) living in the [[Sea surface microlayer|surface microlayer]] sandwiched between the upper and under surface.

Marshall and Burchardt divide neuston into three ecological categories:<ref name=Marshall2005>{{cite journal |doi = 10.1127/0003-9136/2005/0164-0429|title = Neuston: Its definition with a historical review regarding its concept and community structure|year = 2005|last1 = Marshall|first1 = Harold G.|last2 = Burchardt|first2 = Lubomira|journal = Archiv für Hydrobiologie|volume = 164|issue = 4|pages = 429–448}}</ref><ref name=Albuquerque2021 /> 
* ''euneuston'': organisms with maximum abundance in the vicinity of the surface on which they reside day and night
* ''facultative neuston'': organisms concentrating at the surface only during certain hours of the day, usually during darkness
* ''pseudoneuston'': organisms with maximum concentrations at deeper layers but reaching the surface layer at least during certain hours.

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==Freshwater neuston==
Freshwater neuston, organisms living at lake or pond surfaces or slow moving parts of rivers and streams, include [[beetle]]s (see [[whirligig beetle]]), [[protozoans]], [[bacteria]] and [[spiders]] (see [[fishing spider]] and [[diving bell spider]]).  [[Springtail]]s in the genera ''[[Podura]]'' and ''[[Sminthurides]]'' are almost exclusively neustonic, while ''[[Hypogastrura]]'' species often aggregate on pond surfaces.  [[Water strider]]s such as ''[[Gerris]]'' are common examples of insects that support their weight on water's [[surface tension]].

<gallery mode="packed" style="float:left" heights="160px">
File:Gyrinus natator.JPG|[[Whirligig beetle]] (''[[Gyrinus natator]]'')
File:Courtship ritual of Sminthurides aquaticus.webm|Water springtail (''[[Sminthurides|Sminthurides aquaticus]]'')
File:Water spider 2.jpg|[[Diving bell spider]] (''Argyroneta aquatica'')
File:Gerris by webrunner.JPG| [[Waterstrider]] (''[[Gerris|Gerris commun]]'')
File:Wolffia-Spirodela.jpg| [[Duckweed]] on a pond<br />By size: [[Spirodela polyrhiza|greater duckweed]], [[Lemna minor|lesser duckweed]] and [[Wolffia arrhiza|rootless dwarf duckweed]]
</gallery>
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===Floods===
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[[File:Nuvola apps kaboodle.svg|16px]] [https://www.youtube.com/watch?v=cfKr6rnpakE&ab_channel=DeepLook Fire Ants Turn Into a Stinging Life Raft]<br />{{align|right|– ''YouTube''}}
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There are different terrestrial environmental factors such as flood pulses and droughts, and these environmental factors affect species such as neuston, whether the effects lead to more or less variations in the species. When flood pulses (an abiotic factor) occur, connectivity between different aquatic environments occur. Species that live in environments with irregular flood patterns tend to have more variations, or even decrease species and variations; similar idea to what happens when droughts occur.<ref>Conceição, E. de O. da, Higuti, J., Campos, R. de, & Martens, K. (2018). Effects of flood pulses on persistence and variability of pleuston communities in a tropical floodplain lake. Hydrobiologia, 807(1), 175–188.</ref>

[[Red imported fire ant|Red fire ants]] have adapted to contend with both flooding and drought conditions. If the ants sense increased water levels in their nests, they link together and form a ball or raft that floats, with the workers on the outside and the queen inside.<ref>{{cite journal|last1=Mlot|first1=N.J.|last2=Craig|first2=A.T.|last3=Hu|first3=D.L.|title=Fire ants self-assemble into waterproof rafts to survive floods|journal=Proceedings of the National Academy of Sciences|volume=108|issue=19|pages=7669–7673|date=2011|bibcode=2011PNAS..108.7669M|doi=10.1073/pnas.1016658108|pmid=21518911|pmc=3093451|doi-access=free}}</ref><ref>{{cite journal|last1=Morrill|first1=W.L.|title=Dispersal of red imported fire ants by water|journal=The Florida Entomologist|date=1974|volume=57|issue=1|pages=39–42|doi=10.2307/3493830|jstor=3493830}}</ref>{{sfn|Hölldobler|Wilson|1990|p=171}} The brood is transported to the highest surface.<ref name="Adams_2011">{{cite journal|last1=Adams|first1=B.J.|last2=Hooper-Bùi|first2=L.M.|last3=Strecker|first3=R.M.|last4=O'Brien|first4=D.M.|title=Raft formation by the red imported fire ant, ''Solenopsis invicta''|journal=Journal of Insect Science|date=2011|volume=11|issue=171|pages=171|doi=10.1673/031.011.17101|pmid=22950473|pmc=3462402}}</ref> They are also used as the founding structure of the raft, except for the eggs and smaller larvae. Before submerging, the ants will tip themselves into the water and sever connections with the dry land. In some cases, workers may deliberately remove all males from the raft, resulting in the males drowning.

The longevity of a raft can be as long as 12 days. Ants that are trapped underwater escape by lifting themselves to the surface using bubbles which are collected from submerged substrate.<ref name="Adams_2011" /> Owing to their greater vulnerability to predators, red imported fire ants are significantly more aggressive when rafting. Workers tend to deliver higher doses of venom, which reduces the threat of other animals attacking. Due to this, and because a higher workforce of ants is available, rafts are potentially dangerous to those that encounter them.<ref>{{cite journal|last1=Haight|first1=K.L.|title=Defensiveness of the fire ant, ''Solenopsis invicta'', is increased during colony rafting|journal=Insectes Sociaux|date=2006|volume=53|issue=1|pages=32–36|doi=10.1007/s00040-005-0832-y|s2cid=24420242}}</ref>

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==Marine  neuston==
{{further|Ocean surface ecosystem|Sea surface microlayer}}

The marine neuston, organisms living at the ocean surface, are one of the least studied planktonic groups. Neuston occupies a restricted ecological niche and is affected by a wide range of endogenous and exogenous processes while also being a food source to zooplankton and fish migrating from the deep layers and seabirds.<ref name=Albuquerque2021>{{cite journal |doi = 10.3389/fmars.2020.606088|doi-access = free|title = Trophic Structure of Neuston Across Tropical and Subtropical Oceanic Provinces Assessed with Stable Isotopes|year = 2021|last1 = Albuquerque|first1 = Rui|last2 = Bode|first2 = Antonio|last3 = González-Gordillo|first3 = Juan Ignacio|last4 = Duarte|first4 = Carlos M.|last5 = Queiroga|first5 = Henrique|journal = Frontiers in Marine Science|volume = 7|hdl = 10754/667566|hdl-access = free}} [[File:CC-BY icon.svg|50px]] Material was copied from this source, which is available under a [https://creativecommons.org/licenses/by/4.0/ Creative Commons Attribution 4.0 International License].</ref>

The neustonic animals form a subset of the zooplankton community, which plays a pivotal role in the functioning of marine ecosystems. Zooplankton are partially responsible for the active [[Energy flow (ecology)|energy flux]] between superficial and deep layers of the ocean.<ref>{{cite journal |doi = 10.3354/ame027057|title = Zooplankton fecal pellets, marine snow and sinking phytoplankton blooms|year = 2002|last1 = Turner|first1 = JT|journal = Aquatic Microbial Ecology|volume = 27|pages = 57–102|doi-access = free}}</ref><ref>{{cite journal |doi = 10.1073/pnas.1512110112|title = Seasonal copepod lipid pump promotes carbon sequestration in the deep North Atlantic|year = 2015|last1 = Jónasdóttir|first1 = Sigrún Huld|last2 = Visser|first2 = André W.|last3 = Richardson|first3 = Katherine|last4 = Heath|first4 = Michael R.|journal = Proceedings of the National Academy of Sciences|volume = 112|issue = 39|pages = 12122–12126|pmid = 26338976|pmc = 4593097|doi-access = free}}</ref><ref>{{cite journal |doi = 10.1038/s41467-020-19875-7|title = Large deep-sea zooplankton biomass mirrors primary production in the global ocean|year = 2020|last1 = Hernández-León|first1 = S.|last2 = Koppelmann|first2 = R.|last3 = Fraile-Nuez|first3 = E.|last4 = Bode|first4 = A.|last5 = Mompeán|first5 = C.|last6 = Irigoien|first6 = X.|last7 = Olivar|first7 = M. P.|last8 = Echevarría|first8 = F.|last9 = Fernández De Puelles|first9 = M. L.|last10 = González-Gordillo|first10 = J. I.|last11 = Cózar|first11 = A.|last12 = Acuña|first12 = J. L.|last13 = Agustí|first13 = S.|last14 = Duarte|first14 = C. M.|journal = Nature Communications|volume = 11|issue = 1|page = 6048|pmid = 33247160|pmc = 7695708| bibcode=2020NatCo..11.6048H |s2cid = 227191974}}</ref> Zooplankton [[species composition]], biomass, and secondary production influence a wide range of trophic levels in marine communities, as they constitute a link between primary production and secondary consumers.<ref name=Litchman2013>{{cite journal |doi = 10.1093/plankt/fbt019|title = Trait-based approaches to zooplankton communities|year = 2013|last1 = Litchman|first1 = Elena|last2 = Ohman|first2 = Mark D.|last3 = Kiørboe|first3 = Thomas|journal = Journal of Plankton Research|volume = 35|issue = 3|pages = 473–484|doi-access = free|url = https://escholarship.org/content/qt2vj5c3hx/qt2vj5c3hx.pdf?t=qc3atc}}</ref><ref name=Benedetti2016 /><ref name=deOliveira2019>{{cite journal |doi = 10.1007/s40071-019-0233-x|title = How planktonic microcrustaceans respond to environment and affect ecosystem: A functional trait perspective|year = 2019|last1 = Sodré|first1 = Elder de Oliveira|last2 = Bozelli|first2 = Reinaldo Luiz|journal = International Aquatic Research|volume = 11|issue = 3|pages = 207–223|s2cid = 197594398|doi-access = free| bibcode=2019InAqR..11..207S }}</ref> [[Copepod]]s constitute the most abundant zooplankton taxon in terms of [[biomass (ecology)|biomass]] and [[Biodiversity|diversity]] worldwide.<ref name="Kiørboe2011">{{cite journal |doi = 10.1111/j.1469-185X.2010.00148.x|title = How zooplankton feed: Mechanisms, traits and trade-offs|year = 2011|last1 = Kiørboe|first1 = Thomas|journal = Biological Reviews|volume = 86|issue = 2|pages = 311–339|pmid = 20682007|s2cid = 25218654}}</ref><ref>{{cite journal |doi = 10.3389/fmicb.2018.00355|doi-access = free|title = Zooplankton from a Reef System Under the Influence of the Amazon River Plume|year = 2018|last1 = Neumann-Leitão|first1 = Sigrid|last2 = Melo|first2 = Pedro A. M. C.|last3 = Schwamborn|first3 = Ralf|last4 = Diaz|first4 = Xiomara F. G.|last5 = Figueiredo|first5 = Lucas G. P.|last6 = Silva|first6 = Andrea P.|last7 = Campelo|first7 = Renata P. S.|last8 = Melo Júnior|first8 = Mauro de|last9 = Melo|first9 = Nuno F. A. C.|last10 = Costa|first10 = Alejandro E. S. F.|last11 = Araújo|first11 = Moacyr|last12 = Veleda|first12 = Dóris R. A.|last13 = Moura|first13 = Rodrigo L.|last14 = Thompson|first14 = Fabiano|journal = Frontiers in Microbiology|volume = 9|page = 355|pmid = 29545783|pmc = 5838004}}</ref> Consequently, changes in their community composition can impact the [[biogeochemical cycle]]s{{hsp}}<ref>{{cite journal |doi = 10.1002/lno.10219|title = Global patterns of diel vertical migration times and velocities from acoustic data|year = 2016|last1 = Bianchi|first1 = Daniele|last2 = Mislan|first2 = K. A. S.|journal = Limnology and Oceanography|volume = 61|issue = 1|pages = 353–364|bibcode = 2016LimOc..61..353B| s2cid=3400176 |doi-access = free}}</ref> and might be indicative of climate variability impacts on ecosystem functioning.<ref>{{cite journal |doi = 10.4319/lo.2006.51.6.2607|title = Copepod biodiversity as an indicator of changes in ocean and climate conditions of the northern California current ecosystem|year = 2006|last1 = Hooff|first1 = Rian C.|last2 = Peterson|first2 = William T.|journal = Limnology and Oceanography|volume = 51|issue = 6|pages = 2607–2620|bibcode = 2006LimOc..51.2607H| s2cid=16280978 |doi-access = free}}</ref><ref name=Albuquerque2021 />

<gallery mode=packed style=float:left heights=170px>
File:Portuguese man-o-war Physalia sp.png|[[Portuguese man-o-war]] ''Physalia'' sp.
File:By-the-wind sailor Velella sp.png|By-the-wind sailor ''[[Velella]]'' sp.
File:Blue button Porpita sp.png|Blue button ''[[Porpita]]'' sp.
File:Flying fish from the family Exocoetidae.png|Flying fish from the family [[Exocoetidae]]
File:Buoy barnacle Dosima fascicularis.png|[[Buoy barnacle]] ''Dosima fascicularis''
File:Blue sea dragons Glaucus sp.png|Blue sea dragons ''[[Glaucus (gastropod)|Glaucus]]'' sp.
</gallery>
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Historically, zooplankton assemblages research has focused mainly on taxonomic studies and those related to community structure.<ref name=Pomerleau2015>{{cite journal |doi = 10.1093/plankt/fbv045|title = Evaluation of functional trait diversity for marine zooplankton communities in the Northeast subarctic Pacific Ocean|year = 2015|last1 = Pomerleau|first1 = Corinne|last2 = Sastri|first2 = Akash R.|last3 = Beisner|first3 = Beatrix E.|journal = Journal of Plankton Research|volume = 37|issue = 4|pages = 712–726|doi-access = free}}</ref> However, recently, research has veered toward an alternative [[Phenotypic trait|trait-based approach]],<ref name=Pomerleau2015/><ref name=Benedetti2016 /><ref>{{cite journal |doi = 10.1016/j.ecolind.2017.08.018|title = Ecological indicators and functional groups of copepod assemblages|year = 2017|last1 = Campos|first1 = C.C.|last2 = Garcia|first2 = T.M.|last3 = Neumann-Leitão|first3 = S.|last4 = Soares|first4 = M.O.|journal = Ecological Indicators|volume = 83|pages = 416–426| bibcode=2017EcInd..83..416C | url=http://www.repositorio.ufc.br/handle/riufc/66703 }}</ref> providing a perspective more focused on groups of species with analogous [[functional trait]]s. This allows individuals to be classified into types characterized by the presence/absence of certain [[alleles]] of a [[gene]], into size classes, [[ecological guild]]s, or [[functional group (ecology)|functional group]]s (FGs).<ref>{{cite journal |doi = 10.1111/j.1600-0587.2009.05880.x|title = A diversity of beta diversities: Straightening up a concept gone awry. Part 1. Defining beta diversity as a function of alpha and gamma diversity|year = 2010|last1 = Tuomisto|first1 = Hanna|journal = Ecography|volume = 33| issue=1 |pages = 2–22| bibcode=2010Ecogr..33....2T }}</ref> Functional traits are [[phenotype]]s affecting organism fitness, growth, survival, and reproductive ability.<ref>{{cite journal |doi = 10.1111/j.2007.0030-1299.15559.x|title = Let the concept of trait be functional!|year = 2007|last1 = Violle|first1 = Cyrille|last2 = Navas|first2 = Marie-Laure|last3 = Vile|first3 = Denis|last4 = Kazakou|first4 = Elena|last5 = Fortunel|first5 = Claire|last6 = Hummel|first6 = Irène|last7 = Garnier|first7 = Eric|journal = Oikos|volume = 116|issue = 5|pages = 882–892}}</ref><ref name=deOliveira2019 /> These are regulated by the expression of genes within species, and the expression of traits regulate, in turn, the species fitness under contrasting [[biotic factor|biotic]] and [[abiotic factor|abiotic circumstances]].<ref>{{cite journal |doi = 10.1111/ele.12063|title = The biogeography of marine plankton traits|year = 2013|last1 = Barton|first1 = Andrew D.|last2 = Pershing|first2 = Andrew J.|last3 = Litchman|first3 = Elena|last4 = Record|first4 = Nicholas R.|last5 = Edwards|first5 = Kyle F.|last6 = Finkel|first6 = Zoe V.|last7 = Kiørboe|first7 = Thomas|last8 = Ward|first8 = Ben A.|journal = Ecology Letters|volume = 16|issue = 4|pages = 522–534|pmid = 23360597| bibcode=2013EcolL..16..522B }}</ref> Moreover, a specific functional trait can also develop from the interactions between other traits and environmental conditions,<ref name="Kiørboe2011" /> leading to a given trait grouping being favoured under certain conditions. Zooplankton traits can be classified in accordance to ecological functions – feeding, growth, reproduction, survival, and other characteristics such as [[Morphology (biology)|morphology]], [[physiology]], behaviour, or life history.<ref name=Litchman2013 /><ref>{{cite journal |doi = 10.1016/j.dsr2.2014.10.023|title = A coupled stable isotope-size spectrum approach to understanding pelagic food-web dynamics: A case study from the southwest sub-tropical Pacific|year = 2015|last1 = Hunt|first1 = B.P.V.|last2 = Allain|first2 = V.|last3 = Menkes|first3 = C.|last4 = Lorrain|first4 = A.|last5 = Graham|first5 = B.|last6 = Rodier|first6 = M.|last7 = Pagano|first7 = M.|last8 = Carlotti|first8 = F.|journal = Deep Sea Research Part II: Topical Studies in Oceanography|volume = 113|pages = 208–224|bibcode = 2015DSRII.113..208H}}</ref><ref>{{cite journal |doi = 10.1111/ele.12688|title = Trait biogeography of marine copepods - an analysis across scales|year = 2016|last1 = Brun|first1 = Philipp|last2 = Payne|first2 = Mark R.|last3 = Kiørboe|first3 = Thomas|journal = Ecology Letters|volume = 19|issue = 12|pages = 1403–1413|pmid = 27726281| bibcode=2016EcolL..19.1403B | s2cid=216070768 | url=https://backend.orbit.dtu.dk/ws/files/139594757/Postprint.pdf }}</ref> Particularly, [[Aquatic feeding mechanisms|feeding strategies]] and [[trophic level|trophic groups]] are relevant to establish feeding efficiency and associated predation risk.<ref>{{cite journal |doi = 10.5194/essd-9-99-2017|title = A trait database for marine copepods|year = 2017|last1 = Brun|first1 = Philipp|last2 = Payne|first2 = Mark R.|last3 = Kiørboe|first3 = Thomas|journal = Earth System Science Data|volume = 9|issue = 1|pages = 99–113|bibcode = 2017ESSD....9...99B| s2cid=55732646 |doi-access = free|url = https://essd.copernicus.org/articles/9/99/2017/essd-9-99-2017.pdf}}</ref> Additionally, they facilitate the understanding of [[ecosystem service]]s associated with zooplankton, such as the distribution of fisheries or biogeochemical cycling{{hsp}}<ref>{{cite journal |doi = 10.1002/lno.11067|title = Biogeography of zooplankton feeding strategy|year = 2019|last1 = Prowe|first1 = A. E. Friederike|last2 = Visser|first2 = André W.|last3 = Andersen|first3 = Ken H.|last4 = Chiba|first4 = Sanae|last5 = Kiørboe|first5 = Thomas|journal = Limnology and Oceanography|volume = 64|issue = 2|pages = 661–678|bibcode = 2019LimOc..64..661P|s2cid = 91541174|doi-access = free}}</ref> while also allowing the positioning of zooplankton taxa in the food web.<ref name=Benedetti2016>{{cite journal |doi = 10.1093/plankt/fbv096|title = Identifying copepod functional groups from species functional traits|year = 2016|last1 = Benedetti|first1 = Fabio|last2 = Gasparini|first2 = Stéphane|last3 = Ayata|first3 = Sakina-Dorothée|journal = Journal of Plankton Research|volume = 38|issue = 1|pages = 159–166|pmid = 26811565|pmc = 4722884}}</ref><ref>{{cite journal |doi = 10.1111/jbi.13166|title = Do functional groups of planktonic copepods differ in their ecological niches?|year = 2018|last1 = Benedetti|first1 = Fabio|last2 = Vogt|first2 = Meike|last3 = Righetti|first3 = Damiano|last4 = Guilhaumon|first4 = François|last5 = Ayata|first5 = Sakina-Dorothée|journal = Journal of Biogeography|volume = 45|issue = 3|pages = 604–616| bibcode=2018JBiog..45..604B | s2cid=90358144 | url=https://hal.science/hal-01826722/file/Benedetti%20et%20al._revised_JBio_HAL%20%281%29.pdf }}</ref><ref name=Albuquerque2021 />

<gallery mode=packed style=float:left heights=170px>
File:Paper nautilus Argonauta sp.png|[[Paper nautilus]] ''Argonauta'' sp.
File:Sargassum sp. seaweed.png|''[[Sargassum]]'' sp. seaweed
File:Hippolytidae shrimp.png|[[Hippolytidae]] shrimp
File:Marine snail Recluzia sp.png|Marine snail ''[[Recluzia]]'' sp.
File:Violet snail Janthina sp.png|Violet snail ''[[Janthina]]'' sp.
File:Floating anemone Actinecta sp.png|Floating [[Sea anemone|anemone]] ''Actinecta'' sp.
</gallery>
{{clear}}

[[Hermatobates|Coral-treaders]] are a genus of quite rare wingless marine bugs known only from coral reefs in the Indo-Pacific region. During low tide they move over water surfaces around coral atolls and reefs similar to the more familiar water-striders, staying submerged in reef crevices during high tide.

==See also==
{{Portal|Biology}}
* [[Bacterioneuston]]
* [[Virioneuston]]

==References==
{{reflist}}

==External links==
* {{cite web |url=http://www.britannica.com/eb/article-9055398/neuston |title=neuston - Britannica Online |access-date=2007-11-13 |publisher=Encyclopædia Britannica}}

{{aquatic ecosystem topics}}
{{aquatic organisms}}
{{Authority control}}

[[Category:Aquatic ecology]]
[[Category:Ecology terminology]]