Neuston

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The water strider, a common freshwater neuston Wasserlaufer bei der Paarung crop.jpg
The water strider, a common freshwater neuston

Neuston, also called pleuston, are organisms that live at the 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, microorganisms can exist in the 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." [1]

Contents

Neustons can be informally separated into two groups: the phytoneuston, which are autotrophs floating at the water surface including cyanobacteria, filamentous algae and free-floating aquatic plant (e.g. mosquito fern, duckweed and water lettuce); and the zooneuston, which are floating heterotrophs such as protists (e.g. ciliates) and metazoans (aquatic animals). This article mainly concerns with metazoan zooneustons.

The word "neuston" comes from Greek neustos, meaning "swimming", and the noun suffix -on (as in "plankton"). [2] This term first appears in the biological literature in 1917. [3] 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. [4] 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

Portuguese man o' war emblematic figure of the marine pleuston Portuguese Man-O-War (Physalia physalis).jpg
Portuguese man o' war
emblematic figure of the marine pleuston
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) Neuston, Plankton, Nekton, Benthos.jpg
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. [5] The term was first used in 1917 by Naumann to describe species associated with the surface layer of freshwater habitats. [3] Later in 1971, Zaitsev identified neuston composition in marine waters. [6] These populations would include microscopic species, plus various plant and animal taxa, such as phytoplankton and zooplankton, living in this region. [6] [7] In 2002, Gladyshev further characterised the major physical and chemical dynamics of the surface layer influencing the composition and relationships with various neustonic populations" [8] [7]

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. [7] [9] [10] Furthermore, the neuston provides a food source to the zooplankton migrating from deeper layers to the surface, [11] as well as to seabirds roaming over the oceans. [12] For these reasons, the neustonic community is believed to play a critical role on the structure and function of marine food webs. Yet, research on neuston communities to date focused predominantly on geographically limited regions of the ocean  [13] [11] [14] [15] [10] or coastal areas. [16] [17] [18] 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, [10] and global scale analyses are yet lacking. [5]

Types

Neuston net Fis01007 (27889419550).jpg
Neuston net

There are different ways neuston can be categorised. Kennish divides them by their physical position into two groups: [1]

To this can be added the organisms living in the microlayer at the interface between air and water:

Marshall and Burchardt divide neuston into three ecological categories: [7] [5]

Freshwater neuston

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

Floods

RIFA Raft After Heavy Rain.jpg
Raft of red fire ants floating over a pond bank submerged by heavy rain
Nuvola apps kaboodle.svg Fire Ants Turn Into a Stinging Life Raft
– YouTube

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. [19]

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. [20] [21] [22] The brood is transported to the highest surface. [23] 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. [23] 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. [24]

Marine neuston

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. [5]

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 flux between superficial and deep layers of the ocean. [25] [26] [27] 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. [28] [29] [30] Copepods constitute the most abundant zooplankton taxon in terms of biomass and diversity worldwide. [31] [32] Consequently, changes in their community composition can impact the biogeochemical cycles   [33] and might be indicative of climate variability impacts on ecosystem functioning. [34] [5]

Historically, zooplankton assemblages research has focused mainly on taxonomic studies and those related to community structure. [35] However, recently, research has veered toward an alternative trait-based approach, [35] [29] [36] providing a perspective more focused on groups of species with analogous functional traits. This allows individuals to be classified into types characterized by the presence/absence of certain alleles of a gene, into size classes, ecological guilds, or functional groups (FGs). [37] Functional traits are phenotypes affecting organism fitness, growth, survival, and reproductive ability. [38] [30] These are regulated by the expression of genes within species, and the expression of traits regulate, in turn, the species fitness under contrasting biotic and abiotic circumstances. [39] Moreover, a specific functional trait can also develop from the interactions between other traits and environmental conditions, [31] 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, physiology, behaviour, or life history. [28] [40] [41] Particularly, feeding strategies and trophic groups are relevant to establish feeding efficiency and associated predation risk. [42] Additionally, they facilitate the understanding of ecosystem services associated with zooplankton, such as the distribution of fisheries or biogeochemical cycling  [43] while also allowing the positioning of zooplankton taxa in the food web. [29] [44] [5]

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

Related Research Articles

<span class="mw-page-title-main">Plankton</span> Organisms living in water or air that are drifters on the current or wind

Plankton are the diverse collection of organisms that drift in water but are unable to actively propel themselves against currents. The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucial source of food to many small and large aquatic organisms, such as bivalves, fish, and baleen whales.

<span class="mw-page-title-main">Phytoplankton</span> Autotrophic members of the plankton ecosystem

Phytoplankton are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words φυτόν, meaning 'plant', and πλαγκτός, meaning 'wanderer' or 'drifter'.

<span class="mw-page-title-main">Zooplankton</span> Heterotrophic protistan or metazoan members of the plankton ecosystem

Zooplankton are the animal component of the planktonic community, having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequently, they drift or are carried along by currents in the ocean, or by currents in seas, lakes or rivers.

<span class="mw-page-title-main">Biological pump</span> Carbon capture process in oceans

The biological pump (or ocean carbon biological pump or marine biological carbon pump) is the ocean's biologically driven sequestration of carbon from the atmosphere and land runoff to the ocean interior and seafloor sediments. In other words, it is a biologically mediated process which results in the sequestering of carbon in the deep ocean away from the atmosphere and the land. The biological pump is the biological component of the "marine carbon pump" which contains both a physical and biological component. It is the part of the broader oceanic carbon cycle responsible for the cycling of organic matter formed mainly by phytoplankton during photosynthesis (soft-tissue pump), as well as the cycling of calcium carbonate (CaCO3) formed into shells by certain organisms such as plankton and mollusks (carbonate pump).

The mesopelagiczone, also known as the middle pelagic or twilight zone, is the part of the pelagic zone that lies between the photic epipelagic and the aphotic bathypelagic zones. It is defined by light, and begins at the depth where only 1% of incident light reaches and ends where there is no light; the depths of this zone are between approximately 200 to 1,000 meters below the ocean surface.

The oxygen minimum zone (OMZ), sometimes referred to as the shadow zone, is the zone in which oxygen saturation in seawater in the ocean is at its lowest. This zone occurs at depths of about 200 to 1,500 m (700–4,900 ft), depending on local circumstances. OMZs are found worldwide, typically along the western coast of continents, in areas where an interplay of physical and biological processes concurrently lower the oxygen concentration and restrict the water from mixing with surrounding waters, creating a "pool" of water where oxygen concentrations fall from the normal range of 4–6 mg/L to below 2 mg/L.

<span class="mw-page-title-main">Holoplankton</span>

Holoplankton are organisms that are planktic for their entire life cycle. Holoplankton can be contrasted with meroplankton, which are planktic organisms that spend part of their life cycle in the benthic zone. Examples of holoplankton include some diatoms, radiolarians, some dinoflagellates, foraminifera, amphipods, krill, copepods, and salps, as well as some gastropod mollusk species. Holoplankton dwell in the pelagic zone as opposed to the benthic zone. Holoplankton include both phytoplankton and zooplankton and vary in size. The most common plankton are protists.

<span class="mw-page-title-main">Marine ecosystem</span> Ecosystem in saltwater environment

Marine ecosystems are the largest of Earth's aquatic ecosystems and exist in waters that have a high salt content. These systems contrast with freshwater ecosystems, which have a lower salt content. Marine waters cover more than 70% of the surface of the Earth and account for more than 97% of Earth's water supply and 90% of habitable space on Earth. Seawater has an average salinity of 35 parts per thousand of water. Actual salinity varies among different marine ecosystems. Marine ecosystems can be divided into many zones depending upon water depth and shoreline features. The oceanic zone is the vast open part of the ocean where animals such as whales, sharks, and tuna live. The benthic zone consists of substrates below water where many invertebrates live. The intertidal zone is the area between high and low tides. Other near-shore (neritic) zones can include mudflats, seagrass meadows, mangroves, rocky intertidal systems, salt marshes, coral reefs, lagoons. In the deep water, hydrothermal vents may occur where chemosynthetic sulfur bacteria form the base of the food web.

<span class="mw-page-title-main">Diel vertical migration</span> A pattern of daily vertical movement characteristic of many aquatic species

Diel vertical migration (DVM), also known as diurnal vertical migration, is a pattern of movement used by some organisms, such as copepods, living in the ocean and in lakes. The adjective "diel" comes from Latin: diēs, lit. 'day', and refers to a 24-hour period. The migration occurs when organisms move up to the uppermost layer of the water at night and return to the bottom of the daylight zone of the oceans or to the dense, bottom layer of lakes during the day. DVM is important to the functioning of deep-sea food webs and the biologically-driven sequestration of carbon.

<span class="mw-page-title-main">Paradox of the plankton</span> The ecological observation of high plankton diversity despite competition for few resources

In aquatic biology, the paradox of the plankton describes the situation in which a limited range of resources supports an unexpectedly wide range of plankton species, apparently flouting the competitive exclusion principle, which holds that when two species compete for the same resource, one will be driven to extinction.

<span class="mw-page-title-main">Sea surface microlayer</span> Boundary layer where all exchange occurs between the atmosphere and the ocean

The sea surface microlayer (SML) is the boundary interface between the atmosphere and ocean, covering about 70% of Earth's surface. With an operationally defined thickness between 1 and 1,000 μm (1.0 mm), the SML has physicochemical and biological properties that are measurably distinct from underlying waters. Recent studies now indicate that the SML covers the ocean to a significant extent, and evidence shows that it is an aggregate-enriched biofilm environment with distinct microbial communities. Because of its unique position at the air-sea interface, the SML is central to a range of global marine biogeochemical and climate-related processes.

<span class="mw-page-title-main">Gelatinous zooplankton</span> Fragile and often translucent animals that live in the water column

Gelatinous zooplankton are fragile animals that live in the water column in the ocean. Their delicate bodies have no hard parts and are easily damaged or destroyed. Gelatinous zooplankton are often transparent. All jellyfish are gelatinous zooplankton, but not all gelatinous zooplankton are jellyfish. The most commonly encountered organisms include ctenophores, medusae, salps, and Chaetognatha in coastal waters. However, almost all marine phyla, including Annelida, Mollusca and Arthropoda, contain gelatinous species, but many of those odd species live in the open ocean and the deep sea and are less available to the casual ocean observer. Many gelatinous plankters utilize mucous structures in order to filter feed. Gelatinous zooplankton have also been called Gelata.

<span class="mw-page-title-main">Ecosystem of the North Pacific Subtropical Gyre</span> Major circulating ecosystem of ocean currents

The North Pacific Subtropical Gyre (NPSG) is the largest contiguous ecosystem on earth. In oceanography, a subtropical gyre is a ring-like system of ocean currents rotating clockwise in the Northern Hemisphere and counterclockwise in the Southern Hemisphere caused by the Coriolis Effect. They generally form in large open ocean areas that lie between land masses.

<span class="mw-page-title-main">Marine habitat</span> Habitat that supports marine life

A marine habitat is a habitat that supports marine life. Marine life depends in some way on the saltwater that is in the sea. A habitat is an ecological or environmental area inhabited by one or more living species. The marine environment supports many kinds of these habitats.

<span class="mw-page-title-main">Planktivore</span> Aquatic organism that feeds on planktonic food

A planktivore is an aquatic organism that feeds on planktonic food, including zooplankton and phytoplankton. Planktivorous organisms encompass a range of some of the planet's smallest to largest multicellular animals in both the present day and in the past billion years; basking sharks and copepods are just two examples of giant and microscopic organisms that feed upon plankton.

<span class="mw-page-title-main">Mycoplankton</span> Fungal members of the plankton communities of aquatic ecosystems

Mycoplankton are saprotrophic members of the plankton communities of marine and freshwater ecosystems. They are composed of filamentous free-living fungi and yeasts that are associated with planktonic particles or phytoplankton. Similar to bacterioplankton, these aquatic fungi play a significant role in heterotrophicmineralization and nutrient cycling. Mycoplankton can be up to 20 mm in diameter and over 50 mm in length.

<span class="mw-page-title-main">Particulate organic matter</span>

Particulate organic matter (POM) is a fraction of total organic matter operationally defined as that which does not pass through a filter pore size that typically ranges in size from 0.053 millimeters (53 μm) to 2 millimeters.

<span class="mw-page-title-main">Lipid pump</span>

The lipid pump sequesters carbon from the ocean's surface to deeper waters via lipids associated with overwintering vertically migratory zooplankton. Lipids are a class of hydrocarbon rich, nitrogen and phosphorus deficient compounds essential for cellular structures. This lipid carbon enters the deep ocean as carbon dioxide produced by respiration of lipid reserves and as organic matter from the mortality of zooplankton.

<span class="mw-page-title-main">Marine food web</span> Marine consumer-resource system

A marine food web is a food web of marine life. At the base of the ocean food web are single-celled algae and other plant-like organisms known as phytoplankton. The second trophic level is occupied by zooplankton which feed off the phytoplankton. Higher order consumers complete the web. There has been increasing recognition in recent years that marine microorganisms.

<span class="mw-page-title-main">Ocean surface ecosystem</span>

Organisms that live freely at the ocean surface, termed neuston, include keystone organisms like the golden seaweed Sargassum that makes up the Sargasso Sea, floating barnacles, marine snails, nudibranchs, and cnidarians. Many ecologically and economically important fish species live as or rely upon neuston. Species at the surface are not distributed uniformly; the ocean's surface provides habitat for unique neustonic communities and ecoregions found at only certain latitudes and only in specific ocean basins. But the surface is also on the front line of climate change and pollution. Life on the ocean's surface connects worlds. From shallow waters to the deep sea, the open ocean to rivers and lakes, numerous terrestrial and marine species depend on the surface ecosystem and the organisms found there.

References

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  40. ↑ Hunt, B.P.V.; Allain, V.; Menkes, C.; Lorrain, A.; Graham, B.; Rodier, M.; Pagano, M.; Carlotti, F. (2015). "A coupled stable isotope-size spectrum approach to understanding pelagic food-web dynamics: A case study from the southwest sub-tropical Pacific". Deep Sea Research Part II: Topical Studies in Oceanography. 113: 208–224. Bibcode:2015DSRII.113..208H. doi:10.1016/j.dsr2.2014.10.023.
  41. ↑ Brun, Philipp; Payne, Mark R.; Kiørboe, Thomas (2016). "Trait biogeography of marine copepods - an analysis across scales" (PDF). Ecology Letters. 19 (12): 1403–1413. Bibcode:2016EcolL..19.1403B. doi:10.1111/ele.12688. PMID   27726281. S2CID   216070768.
  42. ↑ Brun, Philipp; Payne, Mark R.; Kiørboe, Thomas (2017). "A trait database for marine copepods" (PDF). Earth System Science Data. 9 (1): 99–113. Bibcode:2017ESSD....9...99B. doi: 10.5194/essd-9-99-2017 . S2CID   55732646.
  43. ↑ Prowe, A. E. Friederike; Visser, André W.; Andersen, Ken H.; Chiba, Sanae; Kiørboe, Thomas (2019). "Biogeography of zooplankton feeding strategy". Limnology and Oceanography. 64 (2): 661–678. Bibcode:2019LimOc..64..661P. doi: 10.1002/lno.11067 . S2CID   91541174.
  44. ↑ Benedetti, Fabio; Vogt, Meike; Righetti, Damiano; Guilhaumon, François; Ayata, Sakina-Dorothée (2018). "Do functional groups of planktonic copepods differ in their ecological niches?" (PDF). Journal of Biogeography. 45 (3): 604–616. Bibcode:2018JBiog..45..604B. doi:10.1111/jbi.13166. S2CID   90358144.
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