Diving bell spider

{{Short description|Genus of spiders}} {{Speciesbox | name = Diving bell spider | image = Argyroneta aquatica Paar.jpg | image_caption = Female (on left) and male | image2 = | image2_caption = | fossil_range = {{Fossil range|Neogene|present}} | taxon = Argyroneta aquatica | authority = (Clerck, 1758) | synonyms = *''Aranea amphibia'' Müller, 1776<ref name=Müll1776 /> *''Aranea aquatica'' Linnaeus, 1758<ref name=Linn1758 /> *''Araneus aquaticus'' Clerck, 1757<ref name=Cler1757 /> *''Aranea urinatoria'' Poda, 1761<ref name=Poda1761 /> *''Clubiona fallax'' Walckenaer, 1837<ref name=Walc1837 /> | synonyms_ref = <ref name=wsc /> | parent_authority = Latreille, 1804 }}

The '''diving bell spider''' or '''water spider''' ('''''Argyroneta aquatica''''') is the only species of spider known to live almost entirely under water. It is the only member of the genus '''''Argyroneta''''' and the type genus of the family Argyronetidae.<ref name=wsc /> When out of the water, the spider ranges in colour from mid to dark brown, although the hairs on the abdomen give it a dark grey, velvet-like appearance.<ref name=DanmarksEdderkopper /> It is native to freshwater habitats in Europe and Asia.<ref name=wsc /><ref name="Schütz2003" />

==Uniqueness of aquatic behavior==

thumb|Although prey is usually consumed underwater in the diving bell, it is occasionally brought to the surface.

''A. aquatica'' is the only known species of spider that spends almost all its life underwater, including resting, catching and eating prey, mating, egg laying, and overwintering. It only briefly surfaces to replenish its oxygen supply and occasionally will bring prey to the surface.<ref name=Thorp2015 /><ref name=Hillyard2007 /><ref name=BAS /><ref name=Schutz2007 />

Several other spiders are semiaquatic, either periodically living underwater or willing to dive.<ref name="swierk2022" /> For example, certain ''Desis'' species spend the high tide in an air-filled underwater retreat made from silk and forage on land in the intertidal zone during low tide.<ref name=Hillyard2007 /><ref name="baehr2017" /><ref name="arau" /> Some spiders living in periodically flooded habitats can survive for an extended period underwater by entering a coma-like state, up to 16–36 hours in ''Arctosa fulvolineata''.<ref name="petillon2009" /> Numerous species, including some ''Ancylometes'', ''Dolomedes'', ''Megadolomedes'', ''Pardosa'', ''Pirata'', ''Thalassius'' and others, live above water at the surface, but may actively submerge for a prolonged period, are strong swimmers and will catch underwater prey.<ref name=Thorp2015 /><ref name=Hillyard2007 /><ref name="nyffeler2014" /> Several of these, as well as a few others, may dive into the water to avoid larger predators.<ref name=Thorp2015 /><ref name=Henaut2018 />

==Distribution and habitat== ''A. aquatica'' is found in clean freshwater habitats with aquatic vegetation, such as lakes, ponds, canals, marshes and slow-moving streams.<ref name=BAS /><ref name="filoramo2012" /> It ranges through much of mainland Europe (no records from Portugal, Greece and Albania), the British Isles and central to northern Asia ranging as far south as Iran and as far north as Siberia, up to latitude 62°N.<ref name="Schütz2003" /><ref name=SpidersEurope /> Most of the range is inhabited by the nominate subspecies, but Japan has its own subspecies, the very similar ''A. a. japonica''.<ref name=DanmarksEdderkopper /><ref name="ono2002" />

thumb|Habitat at Yeoncheon Eundaeri Water Spider Habitat

South Korea has a protected area, ''Yeoncheon Eundaeri water spider habitat'',<ref name="enckor" /><ref name="protec" /> for this endangered species in that country.<ref name="nie" />

==Ecology== As with other spiders it breathes air; when submerged in water, an air bubble is trapped by a dense layer of hydrophobic hairs on its abdomen and legs,<ref name="Schütz2003" /> giving the abdomen a silvery appearance. The spider lives for about two years in captivity.<ref name="Schütz2003" />

''A. aquatica'' is able to remain submerged for prolonged periods of time due to the silk-based structure it constructs in order to retain an oxygen supply, named after the diving bell structure it resembles. The species range in size, although the size of females may be limited as they put more energy into building and maintaining their larger bells.<ref name="Schütz2003" /> Males are more active and on average almost 30% larger than females,<ref name="Schütz2003" /> measuring {{convert|10-15|mm|in|abbr=on}} in head-and-body length compared to {{convert|8-12|mm|in|abbr=on}}.<ref name=DanmarksEdderkopper /> This size differential favoring males is unusual for spiders, where sexual dimorphism is usually in favour of larger females. Theories suggest that the male's more active hunting style requires greater strength to overcome water resistance and counteract the buoyancy of their mobile air supplies. This larger body size is also associated with longer front legs, shown to affect diving ability and giving the males superiority in diving over the more sessile females. In addition, females build larger air bells than males, as the bell is also used to care for the offspring, forcing them to collect air from the surface much more frequently. And the larger the body, the more air is trapped on their abdomen, which means more buoyancy to overcome. Larger females therefore have higher energy costs than males of the same size and smaller females, which could limit the number of offspring they are able to produce.<ref name="Schütz2003" />

The spiders prey on aquatic insects and crustaceans such as mosquito larvae and ''Daphnia''.<ref name="Seymour2011" /> The spiders themselves fall prey to frogs and fish.<ref name="Waterspider" />

==Diving bell== The appearance of the diving bell gave rise to the genus name ''Argyroneta'', from the Greek "argyros" (ἄργυρος), meaning "silver", and "neta", a neologism (perhaps for *νητής) derived from the verb "neo" (νέω) "spin", intended to mean "spinner of silver".<ref name="OES" /> Both sexes build diving bell webs which are used for digesting prey, although only the female's larger bell is used for mating and raising offspring. Females spend most of their time within their bells, darting out to catch prey animals that touch the bell or the silk threads that anchor it and occasionally surfacing to replenish the air within the web. The bells built by males are typically smaller than females' and are replenished less often. It is thought that prior to mating, the male constructs a diving bell adjacent to the female's then spins a tunnel from his bell, breaking into hers to gain entrance.<ref name="Waterspider" /> Mating takes place in the female's bell.<ref name="schutz2005" /> The female spider then constructs an egg sac within her bell, laying between 30 and 70 eggs.<ref name="Waterspider" /> Where this species moults is less clear, with some sources stating that it occurs below water in the diving bell<ref name=Schutz2007 /> and others that it occurs out of water.<ref name=BAS />

Diving bells are irregularly constructed sheets of silk and an unknown protein-based hydrogel<ref name="neumann2013" /> which is spun between submerged water plants then inflated with air brought down from the surface by the builder. Studies have considered gas diffusion between the diving bell and the spiders' aquatic environment. The silk is waterproof but allows gas exchange with the surrounding water. There is net diffusion of oxygen into the bell and net diffusion of carbon dioxide out. This process is driven by differences in partial pressure. The production of carbon dioxide and use of oxygen by the spider maintains the concentration gradient, required for diffusion. However, there is net diffusion of nitrogen out of the bell, resulting in a gradually shrinking air bubble which must be regularly replenished by the spider.<ref name="Seymour2011" />

Larger spiders are able to produce larger bubbles which have a consequently higher oxygen conductance, but all spiders of this species are able to enlarge their bells in response to increased oxygen demands in low aquatic P(O2) environments. These spiders voluntarily tolerate internal conditions of low oxygen, enlarging their bells with air when the P(O2) drops below 1 kPa; this replenishment process may not need to occur for several days, in some cases.<ref name="Seymour2011" /> This system has been referred to as "the water spider's aqua-lung of air bubbles", though an aqua-lung lacks gas exchange with the surroundings;<ref name="flynn2008" /> this system is more properly regarded as an inorganic form of gill.

==Bite== Their bite is often described as being very painful to humans and as causing localised inflammation, vomiting, and slight feverishness that disappears after 5–10 days.<ref name="ross2007" /><ref name="foelix2011" /> However, solid evidence is lacking,<ref name="white2008" /> with information being based on old and unverified reports because recent confirmed and published reports are lacking,<ref name=SpidersEurope /><ref name="nentwig2013" /> leading some sources to refer to its bite as reputedly painful.<ref name=DanmarksEdderkopper />

==References== {{Reflist|refs= <ref name="wsc">{{cite journal| title=Gen. 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==External links== *[https://web.archive.org/web/20080915164452/http://www.arkive.org/species/ARK/invertebrates_terrestrial_and_freshwater/Argyroneta_aquatica/ Water spider pictures] on Arkive.org *{{cite web|url=http://mmem.spschools.org/grade5science/wetland/waterspider.html |archive-url=https://web.archive.org/web/20100507064818/http://mmem.spschools.org/grade5science/wetland/waterspider.html |title=Water Spider |access-date=2014-01-25 |archive-date=2010-05-07}} *[https://wwwnews.live.bbc.co.uk/nature/13614742 Diving bell spiders use bubble webs 'like gills']

Category:Argyronetidae Category:Spiders of Asia Category:Spiders of Europe Category:Palearctic spiders Category:Taxa named by Pierre André Latreille