View source for Dytiscidae

{{Short description|Family of beetles}}
{{Redirect|Water tiger|the mythical Chinese creature|shuihu}}
{{Automatic taxobox
| fossil_range = {{fossil range|Late Jurassic|Recent}}
| image = Cybister lateralimarginalis (Dytiscidae) - (imago), Berkenwoude, the Netherlands.jpg
| image_alt = "Cybister lateralimarginalis"
| image_caption = ''[[Cybister lateralimarginalis]]''
| display_parents = 2
| taxon = Dytiscidae
| authority = [[William Elford Leach|Leach]], 1815
| subdivision_ranks = Subfamilies
| subdivision = * [[Agabinae]] <small>Thomson, 1867</small>
* [[Colymbetinae]] <small>Erichson, 1837</small>
* [[Copelatinae]] <small>Branden, 1885</small>
* [[Coptotominae]] <small>Branden, 1885</small>
* [[Cybistrinae]] <small>Sharp, 1880</small>
* [[Dytiscinae]] <small>Leach, 1815</small>
* [[Hydrodytinae]] <small>Miller, 2001</small>
* [[Hydroporinae]] <small>Aubé, 1836</small>
* [[Laccophilinae]] <small>Gistel, 1856</small>
* [[Lancetinae]] <small>Branden, 1885</small>
* [[Matinae]] <small>Branden, 1885</small>
* † [[Liadytiscinae]] <small>Prokin & Ren, 2010</small>
}}
The '''Dytiscidae''' – based on the Greek ''dytikos'' (δυτικός), "able to dive" – are the '''predaceous diving beetles''', a [[Family (biology)|family]] of [[water beetle]]s. They occur in virtually any freshwater habitat around the world, but a few species live among [[leaf litter]].<ref name=Foster2014>{{cite book| author1=G.N. Foster | author2=D.T. Bilton | year=2014 | chapter=The Conservation of Predaceous Diving Beetles: Knowns, Unknowns and Anecdotes | editor=D.A. Yee | title=Ecology, Systematics, and the Natural History of Predaceous Diving Beetles (Coleoptera: Dytiscidae) | pages=437–462 | publisher=Springer | isbn=978-94-017-9109-0 }}</ref> The adults of most are between {{convert|1|and(-)|2.5|cm|in|abbr=on|1}} long, though much variation is seen between species. The European ''[[Dytiscus latissimus]]'' and Brazilian ''[[Megadytes ducalis]]'' are the largest, reaching up to {{convert|4.5|and|4.75|cm|in|abbr=on|1}} respectively.<ref name=Foster2014/><ref>{{cite web |url=http://www.hurdan.com/search?q=dytiscidae |title=Dytiscidae - Hurdan, the answer engine |access-date=2015-05-19 |url-status=dead |archive-url=https://web.archive.org/web/20150521150320/http://www.hurdan.com/search?q=dytiscidae |archive-date=2015-05-21 }}</ref> In contrast, the smallest is likely the Australian ''[[Limbodessus]] atypicali'' of subterranean waters, which only is about {{convert|0.9|mm|in|abbr=on}} long.<ref name=Foster2014/> Most are dark brown, blackish, or dark [[olive (color)|olive]] in color with golden highlights in some [[subfamilies]]. The [[larva]]e are commonly known as '''water tigers''' due to their voracious appetite.<ref>{{cite book| author=G.C. McGavin | year=2010 | title=Insects | pages=86–87 | publisher=Dorling Kindersley | isbn=978-1-4053-4997-0 }}</ref> They have short, but sharp [[mandible (insect)|mandible]]s and immediately upon biting, they deliver digestive enzymes into prey to suck their liquefied remains. The family includes more than 4,000 described [[species]] in numerous [[genera]].<ref>{{cite web | author=Nilsson, A.N. | year=2013 | url=http://www2.emg.umu.se/projects/biginst/andersn/WCD_20130101.pdf | title=A World Catalogue of the Family Dytiscidae, or the Diving Beetles (Coleoptera, Adephaga) | publisher=University of Umeå | access-date=10 April 2018 | archive-url=https://web.archive.org/web/20180403125519/http://www2.emg.umu.se/projects/biginst/andersn/WCD_20130101.pdf | archive-date=3 April 2018 | url-status=dead }}</ref>

== Habitat ==
Diving beetles are the most diverse beetles in the aquatic environment and can be found in almost every kind of freshwater habitat, from small rock pools to big lakes. Some dytiscid species are also found in brackish water.<ref>{{cite book|last1=Yee|first1=D.A.|chapter=An Introduction to the Dytiscidae: Their Diversity, Historical Importance, Cultural Significance, and Other Musings |title= Ecology, Systematics, and the Natural History of Predaceous Diving Beetles (Coleoptera: Dytiscidae)|year=2014|pages=1–16|doi=10.1007/978-94-017-9109-0_1|isbn=978-94-017-9108-3}}</ref> Diving beetles live in water bodies in various landscapes, including agricultural and urban landscapes.<ref name="Lundkvist">{{cite journal|last1=Lundkvist|first1=E.|last2=Landin|first2=J.|last3=Karlsson|first3=F.|title= Dispersing diving beetles (Dytiscidae) in agricultural and urban landscapes in south-eastern Sweden |journal= Annales Zoologici Fennici |year=2002 }}</ref><ref name="Law1">{{cite journal|last1=Law|first1=A.|last2=Baker|first2=A.|last3=Sayer|first3=C.|last4=Foster|first4=G.|last5=Gunn|first5=I.D.|last6=Taylor|first6=P.|last7=Blaikie|first7=James|last8=Willby|first8=N.J.|title= The effectiveness of aquatic plants as surrogates for wider biodiversity in standing fresh waters |journal= Freshwater Biology |year=2019|volume=64|issue=9|pages=1664–1675|doi=10.1111/fwb.13369|bibcode=2019FrBio..64.1664L |url=https://research.tees.ac.uk/ws/files/7865012/The_effectiveness_of_aquatic_plants_as_surrogates_for_wider_biodiversity_in_standing_fresh_waters.pdf|hdl=1893/30068|s2cid=202032378 |hdl-access=free}}</ref><ref name="Liao1">{{cite journal|last1=Liao|first1=W.|last2=Venn|first2=S.|last3=Niemelä|first3=J.|title=Environmental determinants of diving beetle assemblages (Coleoptera: Dytiscidae) in an urban landscape|journal=Biodiversity and Conservation|year=2020|volume=29|issue=7|pages=2343–2359|doi=10.1007/s10531-020-01977-9|doi-access=free|bibcode=2020BiCon..29.2343L |hdl=10138/315037|hdl-access=free}}</ref> Some species, such as ''[[Agabus uliginosus]]''<ref name="Lundkvist" />  and ''[[Acilius canaliculatus]]'',<ref name="Liao1" /> are found to be relatively tolerant to recent urbanization. One of the most important limiting factors for diving beetle occurrence is the presence of fish, which predate on the beetles (mostly on larvae), compete for food, and change the structure of the habitat. The presence or absence of fish can also affect habitat use and habitat selection of dytiscids.<ref name="Liao6">{{cite journal|last1=Liao|first1=W.|last2=Zanca|first2=T.|last3=Niemelä|first3=J.|title=Predation risk modifies habitat use and habitat selection of diving beetles (Coleoptera: Dytiscidae) in an Urban Pondscape|journal=Global Ecology and Conservation|year=2024|volume=49|pages=e02801|doi=10.1016/j.gecco.2024.e02801|doi-access=free}}</ref><ref name="Brodin">{{cite journal|last1=Brodin|first1=T.|last2=Johansson|first2=F.|last3=Bergsten|first3=J.|title=Predator related oviposition site selection of aquatic beetles (Hydroporus spp.) and effects on offspring life-history|journal=Freshwater Biology|year=2006|volume=51|issue=7|pages=1277–1285|doi=10.1111/j.1365-2427.2006.01563.x|doi-access=free}}</ref> Some species, such as ''[[Oreodytes sanmarkii]]'', occur in exposed areas of waters,<ref>{{cite book |last1= Nilsson|first1=A. N. |last2= Holmen|first2=M. |date=1995 |title=The Aquatic Adephaga (Coleoptera) of the Fennoscandia and Denmark. II. Dytiscidae |location=Leiden, the Netherlands |publisher=Brill. |isbn=9004104569|issn=0106-8377}}</ref> whereas many diving beetles species prefer habitats with aquatic plants,<ref name="Law1" /><ref name="Liao6" /><ref>{{cite journal|last1=Liao|first1=W.|last2=Venn|first2=Stephen.|last3=Niemelä|first3=J.|title= Microhabitats with emergent plants counterbalance the negative effects of fish presence on diving beetle (Coleoptera: Dytiscidae) diversity in urban ponds |journal= Global Ecology and Conservation |year=2023|volume=41|pages=e02361|doi=10.1016/j.gecco.2022.e02361|doi-access=free|bibcode=2023GEcoC..4102361L |hdl=10138/352745|hdl-access=free}}</ref> especially plants with complex structures, such as sedges and bulrush.<ref name="Liao6" />

== Larvae and development ==
[[File:Dytiscus marginalis larva.jpg|thumb|left|Larva of the European diving beetle ''Dytiscus marginalis'']]
When still in larval form, the beetles vary in size from about {{convert|1|to|5|cm|in|abbr=on}}. The larval bodies are shaped like crescents, with the tail long and covered with thin hairs. Six legs protrude from along the [[thorax]], which also sports the same thin hairs. The head is flat and square, with a pair of long, large pincers. When hunting, they cling to grasses or pieces of wood along the bottom, and hold perfectly still until prey passes by, then they lunge, trapping their prey between their front legs and biting down with their pincers. The larvae are also known to partially consume prey and discard the carcass if another potential prey swims nearby. Their usual prey includes [[tadpole]]s and [[glassworm]]s, among other smaller water-dwelling creatures. As the larvae mature, they crawl from the water on the sturdy legs, and bury themselves in the mud for [[pupation]]. After about a week, or longer in some species, they emerge from the mud as adults. Adult diving beetles have been found to oviposit their eggs within frog spawn in highly ephemeral habitats, with their eggs hatching within 24 hours after the frogs and the larvae voraciously predating on the recently hatched tadpoles.

==Edibility==
Adult Dytiscidae, particularly of the genus ''[[Cybister]]'', are [[Entomophagy|edible]]. Remnants of ''[[Cybister explanatus|C. explanatus]]'' were found in prehistoric human [[coprolite]]s in a Nevada cave, likely sourced from the [[Humboldt Sink]].<ref name=Miller>{{cite book|last1=Miller|first1=Kelly|last2=Bergsten|first2=Johannes|title=Diving Beetles of the World: Systematics and Biology of the Dytiscidae|date=3 October 2016|publisher=Johns Hopkins University Press|location=Baltimore|page=20}}</ref> In [[Mexico]], ''C. explanatus'' is eaten roasted and salted to accompany [[taco]]s. In Japan, ''[[Cybister japonicus|C. japonicus]]'' has been used as food in certain regions such as Nagano prefecture. In the [[Guangdong Province]] of [[China]], the latter species, as well as ''[[Cybister bengalensis|C. bengalensis]]'', ''[[Cybister guerini|C. guerini]]'', ''[[Cybister limbatus|C. limbatus]]'', ''[[Cybister sugillatus|C. sugillatus]]'', ''[[Cybister tripunctatus|C. tripunctatus]]'', and probably also the well-known [[great diving beetle]] (''D. marginalis'') are bred for human consumption, though as they are cumbersome to raise due to their carnivorous habit and have a fairly bland (though apparently not offensive) taste and little meat, this is decreasing. Dytiscidae are reportedly also eaten in [[Taiwan]], [[Thailand]], and [[New Guinea]].<ref>De Foliart (2002), Jäch (2003), CSIRO (2004)</ref>
[[File:Dytiscidae.png|thumb|Dytiscidae sp.]]

==Diving beetle conservation==
The greatest threat to diving beetles is the degradation and disappearance of their habitats due to anthropogenic activities.<ref name=Foster2014/> For example, urbanisation has led to the decreasing quantity and quality of dytiscid habitats,<ref name="Liao1"/> which consequentially has increased the distance between habitats.;<ref name="Liao2">{{cite journal|last1=Liao|first1=W.|last2=Venn|first2=S.|last3=Niemelä|first3=J.|title=Diving beetle (Coleoptera: Dytiscidae) community dissimilarity reveals how low landscape connectivity restricts the ecological value of urban ponds|journal=Landscape Ecology|year=2022|volume=37|issue=4 |pages=1049–1058|doi=10.1007/s10980-022-01413-z|doi-access=free|bibcode=2022LaEco..37.1049L }}</ref> thus, dytiscids may be exposed to high predation risks during dispersal. Urbanisation has complex effects on the inter- and intraspecific variation in dytiscid traits. Some flight-related traits of ''[[Acilius canaliculatus]]'' and ''[[Hydaticus seminiger]]'', such as body length and hindwing traits, were found to change along the urban gradient at different scales, whereas the traits of ''[[Ilybius ater]]'' exhibited no change.<ref name="Liao5">{{cite journal|last1=Liao|first1=W.|last2=Lin|first2=H.|title=Urbanisation drives inter- and intraspecific variation in flight-related morphological traits of aquatic insects at different landscape scales|journal=Insect Conservation and Diversity|year=2024|volume=17|issue=2 |pages=287–303|doi=10.1111/icad.12703|doi-access=free}}</ref> Brownification, which refers to the change in surface water colour towards yellow–brown hues caused by recent climate change and land-use change, can also drive changes in dytiscid communities.<ref name="Liao4">{{cite journal|last1=Liao|first1=W.|title=Water Colour Shapes Diving Beetle (Coleoptera: Dytiscidae) Assemblages in Urban Ponds|journal=Insects|year=2024|volume=15|issue=4 |pages=308|doi=10.3390/insects15050308|doi-access=free|pmid=38786864 |pmc=11122460}}</ref> As some species, such as ''[[Dytiscus marginalis]]'', are tolerant to brown water, whereas some species, ''[[Hyphydrus ovatus]]'', tend to occur in clear water, brownification may threaten dytiscid species that are intolerant to highly coloured waters.<ref name="Liao4"/>

Dytiscid adults are eaten by many [[bird]]s, [[mammal]]s, [[reptile]]s, and other vertebrate [[predator]]s, despite their arsenal of chemical defenses.<ref name=Dettner2014/> But by far the most important predator of diving beetles are fish, which limit the occurrence of most diving beetle species to fishless ponds, or to margins of aquatic habitats. Although the larvae of a few dytiscid species may become [[apex predator]]s in small [[pond]]s, their presence is also often incompatible with fish. Therefore, the main focus of water beetle conservation is the protection of natural, fish-less habitats.
In the European Union, two species of diving beetles are protected by the [[Berne Convention on the Conservation of European Wildlife and Natural Habitats]], and thus serve as [[umbrella species]] for the protection of natural aquatic habitats: ''[[Dytiscus latissimus]]'' and ''[[Graphoderus bilineatus]]''.

==Cultural significance==
The diving beetle plays a role in a [[Cherokee]] creation story. According to the narrative, upon finding nowhere to rest in the "liquid chaos" the beetle brought up soft mud from the bottom. This mud then spread out to form all of the land on Earth.<ref name=Miller/>

==Ethnobiology==
Adult Dytiscidae, as well as [[Gyrinidae]], are collected by young girls in [[East Africa]]. It is believed that inducing the beetles to bite the nipples will stimulate breast growth.<ref name=Miller/> The effect of that habit has not been tested, but it is notable that the defense glands of diving beetles contain many types of bioactive steroids.<ref name=Dettner2014>{{cite book| author1=Konrad Dettner | year=2014 | chapter=Chemical Ecology and Biochemistry of Dytiscidae | editor=D.A. Yee | title=Ecology, Systematics, and the Natural History of Predaceous Diving Beetles (Coleoptera: Dytiscidae) | pages=235–306 | publisher=Springer | isbn=978-94-017-9109-0 }}</ref>

==Parasites==
Dytiscidae are parasitised by various [[mites]]. Those in genera ''[[Dytiscacarus]]'' and ''[[Eylais]]'' live beneath the [[elytra]] of their hosts,<ref name=":0">{{Cite journal |last1=Mortazavi |first1=Abdolazim |last2=Hajiqanbar |first2=Hamidreza |last3=Lindquist |first3=Evert E |date=2018-10-20 |title=A new family of mites (Acari: Prostigmata: Raphignathina), highly specialized subelytral parasites of dytiscid water beetles (Coleoptera: Dytiscidae: Dytiscinae) |url=https://doi.org/10.1093/zoolinnean/zlx113 |journal=Zoological Journal of the Linnean Society |volume=184 |issue=3 |pages=695–749 |doi=10.1093/zoolinnean/zlx113 |issn=0024-4082}}</ref><ref>{{Cite journal |last=Aiken |first=R. B. |date=1985-02-01 |title=Attachment sites, phenology, and growth of larvae of Eylais sp. (Acari) on Dytiscus alaskanus J. Balfour-Browne (Coleoptera: Dytiscidae) |url=http://www.nrcresearchpress.com/doi/10.1139/z85-041 |journal=Canadian Journal of Zoology |language=en |volume=63 |issue=2 |pages=267–271 |doi=10.1139/z85-041 |issn=0008-4301}}</ref> those in genus ''Acherontacarus'' attach to the mesosternal regions<ref>{{Cite journal |last1=Aykut |first1=Medeni |last2=Esen |first2=Yunus |last3=Taşar |first3=Gani Erhan |date=2016-07-03 |title=New host-parasite association of Acherontacarus rutilans (Acari, Hydrachnidia, Acherontacaridae) on Scarodytes halensis (Coleoptera: Dytiscidae) |url=http://www.tandfonline.com/doi/full/10.1080/01647954.2016.1174304 |journal=International Journal of Acarology |language=en |volume=42 |issue=5 |pages=242–246 |doi=10.1080/01647954.2016.1174304 |bibcode=2016IJAca..42..242A |s2cid=88103217 |issn=0164-7954}}</ref> and those in genus ''[[Hydrachna]]'' attach to various locations.<ref>{{Cite journal |last1=Arjomandi |first1=Elham |last2=Zawal |first2=Andrzej |last3=Hajiqanbar |first3=Hamidreza |last4=Filip |first4=Ewa |last5=Szenejko |first5=Magdalena |date=2019-07-22 |title=New record of a parasitising species of Hydrachna (Acari, Hydrachnidia) on water beetles Eretes griseus (Fabricius, 1781) (Coleoptera, Dytiscidae, Dytiscinae, Eretini) |url=https://zookeys.pensoft.net/article/34532/ |journal=ZooKeys |issue=865 |pages=31–38 |doi=10.3897/zookeys.865.34532 |issn=1313-2970 |pmc=6663934 |pmid=31379442|doi-access=free |bibcode=2019ZooK..865...31A }}</ref> These mites are parasitic as larvae with the exception of ''Dytiscacarus'', which are parasitic for their entire life cycle.<ref name=":0" />

== Systematics ==
The following taxonomic sequence gives the [[subfamilies]], their associated [[genera]].<ref name=gbif/><ref name=Nilsson/><ref name=Bouchard2011/><ref name=itis/>

Subfamily [[Agabinae]] <small>Thomson, 1867</small>{{Div col|colwidth=23em}}
* ''[[Agabinus]]'' <small>Crotch, 1873</small>
* ''[[Agabus (beetle)|Agabus]]'' <small>Leach, 1817</small>
* ''[[Agametrus]]'' <small>Sharp, 1882</small>
* ''[[Andonectes]]'' <small>Guéorguiev, 1971</small>
* ''[[Hydronebrius]]'' <small>Jakovlev, 1897</small>
* ''[[Hydrotrupes]]'' <small>Sharp, 1882</small>
* ''[[Ilybiosoma]]'' <small>Crotch, 1873</small>
* ''[[Ilybius]]'' <small>Erichson, 1832</small>
* ''[[Leuronectes]]'' <small>Sharp, 1882</small>
* ''[[Platambus]]'' <small>Thomson, 1859</small>
* ''[[Platynectes]]'' <small>Régimbart, 1879</small>
{{div col end}}
Subfamily [[Colymbetinae]] <small>Erichson, 1837</small>{{Div col|colwidth=23em}}
* ''[[Anisomeria (beetle)|Anisomeria]]'' <small>Brinck, 1943</small>
* ''[[Senilites]]'' <small>Brinck, 1948</small>
* ''[[Carabdytes]]'' <small>Balke, Hendrich & Wewalka, 1992</small>
* ''[[Bunites]]'' <small>Spangler, 1972</small>
* ''[[Colymbetes]]'' <small>Clairville, 1806</small>
* ''[[Hoperius]]'' <small>Fall, 1927</small>
* ''[[Meladema]]'' <small>Laporte, 1835</small>
* ''[[Melanodytes]]'' <small>Seidlitz, 1887</small>
* ''[[Neoscutopterus]]'' <small>J.Balfour-Browne, 1943</small>
* ''[[Rhantus]]'' <small>Dejean, 1833</small>
* ''[[Rugosus]]'' <small>García, 2001</small>
{{div col end}}
Subfamily [[Copelatinae]] <small>Branden, 1885</small>
* ''[[Agaporomorphus]]'' <small>Zimmermann, 1921</small>
* ''[[Aglymbus]]'' <small>Sharp, 1880</small>
* ''[[Copelatus]]'' <small>Erichson, 1832</small>
* ''[[Exocelina]]'' <small>Broun, 1886</small>
* ''[[Lacconectus]]'' <small>Motschulsky, 1855</small>
* ''[[Liopterus]]'' <small>Dejean, 1833</small>
* ''[[Madaglymbus]]'' <small>Shaverdo & Balke, 2008</small>
* ''[[Rugosus]]'' <small>García, 2001</small>
Subfamily [[Coptotominae]] <small>Branden, 1885</small>
* ''[[Coptotomus]]'' <small>Say, 1830</small>
Subfamily [[Cybistrinae]]{{Div col|colwidth=23em}}
* ''[[Austrodytes]]'' <small>Watts, 1978</small>
* ''[[Bifurcitus]]'' <small>Brinck, 1945</small>
* ''[[Cybister]]'' <small>Curtis, 1827</small>
* ''[[Nilssondytes]]'' <small>Miller, Michat, & Ferreira Jr., 2024</small>
* ''[[Megadytes]]'' <small>Sharp, 1882</small>
* ''[[Metaxydytes]]'' <small>Miller, Michat, & Ferreira Jr., 2024</small>
* ''[[Paramegadytes]]'' <small>Trémouilles & Bachmann, 1980</small>
* ''[[Onychohydrus]]'' <small>Schaum & White, 1847</small>
* ''[[Regimbartina]]'' <small>Chatanay, 1911</small>
* ''[[Spencerhydrus]]'' <small>Sharp, 1882</small>
* ''[[Sternhydrus]]'' <small>Brinck, 1945</small>
* ''[[Trifurcitus]]'' <small>Brinck, 1945</small>
{{div col end}}
Subfamily [[Dytiscinae]] <small>Leach, 1815</small>

{{Div col|colwidth=23em}}
* ''[[Acilius (genus)|Acilius]]'' <small>Leach, 1817</small>
* ''[[Aethionectes]]'' <small>Sharp, 1882</small>
* ''[[Austrodytes]]'' <small>Watts, 1978</small>
* ''[[Dytiscus]]'' <small>Linnaeus, 1758</small>
* ''[[Eretes]]'' <small>Laporte, 1833</small>
* ''[[Graphoderus]]'' <small>Dejean, 1833</small>
* ''[[Hydaticus]]'' <small>Leach, 1817</small>
* ''[[Hyderodes]]'' <small>Hope, 1838</small>
* ''[[Megadytes]]'' <small>Sharp, 1882</small>
* ''[[Miodytiscus]]'' <small>Wickham, 1911</small>
* ''[[Notaticus]]'' <small>Zimmermann, 1928</small>
* ''[[Onychohydrus]]'' <small>Schaum & White, 1847</small>
* ''[[Regimbartina]]'' <small>Chatanay, 1911</small>
* ''[[Rhantaticus]]'' <small>Sharp, 1880</small>
* ''[[Sandracottus (beetle)|Sandracottus]]'' <small>Sharp, 1882</small>
* ''[[Spencerhydrus]]'' <small>Sharp, 1882</small>
* ''[[Sternhydrus]]'' <small>Brinck, 1945</small>
* ''[[Thermonectus]]'' <small>Dejean, 1833</small>
* ''[[Tikoloshanes]]'' <small>Omer-Cooper, 1956</small>
* †''[[Ambarticus]]''  <small>Yang et al. 2019</small> [[Burmese amber]], Myanmar, Late Cretaceous ([[Cenomanian]]){{div col end}}

Subfamily [[Hydrodytinae]] <small>K.B.Miller, 2001</small>
* ''[[Hydrodytes]]'' <small>K.B.Miller, 2001</small>
* ''[[Microhydrodytes]]'' <small>K.B.Miller, 2002</small>
Subfamily [[Hydroporinae]] <small>Aubé, 1836</small>{{Div col|colwidth=23em}}
* ''[[Africodytes]]'' <small>Biström, 1988</small>
* ''[[Agnoshydrus]]'' <small>Biström, Nilsson & Wewalka, 1997</small>
* ''[[Allodessus]]'' <small>Guignot, 1953</small>
* ''[[Allopachria]]'' <small>Zimmermann, 1924</small>
* ''[[Amarodytes]]'' <small>Régimbart, 1900</small>
* ''[[Amurodytes]]'' <small>Fery & Petrov, 2013</small>
* ''[[Andex]]'' <small>Sharp, 1882</small>
* ''[[Anginopachria]]'' <small>Wewalka, Balke & Hendrich, 2001</small>
* ''[[Anodocheilus]]'' <small>Babington, 1841</small>
* ''[[Antiporus]]'' <small>Sharp, 1882</small>
* ''[[Barretthydrus]]'' <small>Lea, 1927</small>
* ''[[Bidessodes]]'' <small>Régimbart, 1895</small>
* ''[[Bidessonotus]]'' <small>Régimbart, 1895</small>
* ''[[Bidessus]]'' <small>Sharp, 1882</small>
* ''[[Boreonectes]]'' <small>Angus, 2010</small>
* ''[[Borneodessus]]'' <small>Balke, Hendrich, Mazzoldi & Biström, 2002</small>
* ''[[Brachyvatus]]'' <small>Zimmermann, 1919</small>
* ''[[Brancuporus]]'' <small>Hendrich, Toussaint & Balke, 2014</small>
* ''[[Canthyporus]]'' <small>Zimmermann, 1919</small>
* ''[[Carabhydrus]]'' <small>Watts, 1978</small>
* ''[[Celina (beetle)|Celina]]'' <small>Aubé, 1837</small>
* ''[[Chostonectes]]'' <small>Sharp, 1880</small>
* ''[[Clypeodytes]]'' <small>Régimbart, 1894</small>
* ''[[Coelhydrus]]'' <small>Sharp, 1882</small>
* ''[[Comaldessus]]'' <small>Spangler & Barr, 1995</small>
* ''[[Crinodessus]]'' <small>K.B. Miller, 1997</small>
* ''[[Darwinhydrus]]'' <small>Sharp, 1882</small>
* ''[[Deronectes]]'' <small>Sharp, 1882</small>
* ''[[Derovatellus]]'' <small>Sharp, 1882</small>
* ''[[Desmopachria]]'' <small>Babington, 1841</small>
* ''[[Dimitshydrus]]'' <small>Uéno, 1996</small>
* ''[[Ereboporus]]'' <small>K.B. Miller, Gibson & Alarie, 2009</small>
* ''[[Etruscodytes]]'' <small>Mazza, Cianferoni & Rocchi, 2013</small>
* ''[[Fontidessus]]'' <small>K.B. Miller & Spangler, 2008</small>
* ''[[Geodessus]]'' <small>Brancucci, 1979</small>
* ''[[Gibbidessus]]'' <small>Watts, 1978</small>
* ''[[Glareadessus]]'' <small>Wewalka & Biström, 1998</small>
* ''[[Graptodytes]]'' <small>Seidlitz, 1887</small>
* ''[[Haideoporus]]'' <small>Young & Longley, 1976</small>
* ''[[Hemibidessus]]'' <small>Zimmermann, 1921</small>
* ''[[Heroceras]]'' <small>Guignot, 1949</small>
* ''[[Herophydrus]]'' <small>Sharp, 1880</small>
* ''[[Heterhydrus]]'' <small>Fairmaire, 1869</small>
* ''[[Heterosternuta]]'' <small>Strand, 1935</small>
* ''[[Hovahydrus]]'' <small>Biström, 1982</small>
* ''[[Huxelhydrus]]'' <small>Sharp, 1882</small>
* ''[[Hydrocolus]]'' <small>Roughley & Larson in Larson, Alarie & Roughley, 2000</small>
* ''[[Hydrodessus]]'' <small>J. Balfour-Browne, 1953</small>
* ''[[Hydroglyphus]]'' <small>Motschulsky, 1853</small>
* ''[[Hydropeplus]]'' <small>Sharp, 1882</small>
* ''[[Hydroporus]]'' <small>Clairville, 1806</small>
* ''[[Hydrovatus]]'' <small>Motschulsky, 1853</small>
* ''[[Hygrotus]]'' <small>Stephens, 1828</small>
* ''[[Hyphoporus]]'' <small>Sharp, 1880</small>
* ''[[Hyphovatus]]'' <small>Wewalka & Biström, 1994</small>
* ''[[Hyphydrus]]'' <small>Illiger, 1802</small>
* ''[[Hypodessus]]'' <small>Guignot, 1939</small>
* ''[[Iberoporus]]'' <small>Castro & Delgado, 2001</small>
* ''[[Incomptodessus]]'' <small>K.B. Miller & García, 2011</small>
* ''[[Kakadudessus]]'' <small>Hendrich & Balke, 2009</small>
* ''[[Kuschelydrus]]'' <small>Ordish, 1976</small>
* ''[[Laccornellus]]'' <small>Roughley & Wolfe, 1987</small>
* ''[[Laccornis]]'' <small>Gozis, 1914</small>
* ''[[Leiodytes]]'' <small>Guignot, 1936</small>
* ''[[Limbodessus]]'' <small>Guignot, 1939</small>
* ''[[Liodessus]]'' <small>Guignot, 1939</small>
* ''[[Lioporeus]]'' <small>Guignot, 1950</small>
* ''[[Megaporus]]'' <small>Brinck, 1943</small>
* ''[[Metaporus]]'' <small>Guignot, 1945</small>
* ''[[Methles]]'' <small>Sharp, 1882</small>
* ''[[Microdessus]]'' <small>Young, 1967</small>
* ''[[Microdytes]]'' <small>J. Balfour-Browne, 1946</small>
* ''[[Morimotoa]]'' <small>Uéno, 1957</small>
* ''[[Nebrioporus]]'' <small>Régimbart, 1906</small>
* ''[[Necterosoma]]'' <small>W.J. Macleay, 1871</small>
* ''[[Neobidessodes]]'' <small>Hendrich & Balke, 2009</small>
* ''[[Neobidessus]]'' <small>Young, 1967</small>
* ''[[Neoclypeodytes]]'' <small>Young, 1967</small>
* ''[[Neoporus]]'' <small>Guignot, 1931</small>
* ''[[Oreodytes]]'' <small>Seidlitz, 1887</small>
* ''[[Pachydrus]]'' <small>Sharp, 1882</small>
* ''[[Pachynectes]]'' <small>Régimbart, 1903</small>
* ''[[Papuadessus]]'' <small>Balke, 2001</small>
* ''[[Paroster]]'' <small>Sharp, 1882</small>
* ''[[Peschetius]]'' <small>Guignot, 1942</small>
* ''[[Petrodessus]]'' <small>K.B. Miller, 2012</small>
* ''[[Phreatodessus]]'' <small>Ordish, 1976</small>
* ''[[Platydytes]]'' <small>Biström, 1988</small>
* ''[[Porhydrus]]'' <small>Guignot, 1945</small>
* ''[[Primospes]]'' <small>Sharp, 1882</small>
* ''[[Pseuduvarus]]'' <small>Biström, 1988</small>
* ''[[Psychopomporus]]'' <small>Jean, Telles & K.B. Miller, 2012</small>
* ''[[Pteroporus]]'' <small>Guignot, 1933</small>
* ''[[Queda (beetle)|Queda]]'' <small>Sharp, 1882</small>
* ''[[Rhithrodytes]]'' <small>Bameul, 1989</small>
* ''[[Sanfilippodytes]]'' <small>Franciscolo, 1979</small>
* ''[[Scarodytes]]'' <small>Gozis, 1914</small>
* ''[[Schistomerus]]'' <small>Palmer, 1957</small>
* ''[[Sekaliporus]]'' <small>Watts, 1997</small>
* ''[[Sharphydrus]]'' <small>Omer-Cooper, 1958</small>
* ''[[Siamoporus]]'' <small>Spangler, 1996</small>
* ''[[Siettitia]]'' <small>Abeille de Perrin, 1904</small>
* ''[[Sinodytes]]'' <small>Spangler, 1996</small>
* ''[[Spanglerodessus]]'' <small>K.B. Miller & García, 2011</small>
* ''[[Sternopriscus]]'' <small>Sharp, 1880</small>
* ''[[Stictonectes]]'' <small>Brinck, 1943</small>
* ''[[Stictotarsus]]'' <small>Zimmermann, 1919</small>
* ''[[Stygoporus]]'' <small>Larson & LaBonte, 1994</small>
* ''[[Suphrodytes]]'' <small>Gozis, 1914</small>
* ''[[Tepuidessus]]'' <small>Spangler, 1981</small>
* ''[[Terradessus]]'' <small>Watts, 1982</small>
* ''[[Tiporus]]'' <small>Watts, 1985</small>
* ''[[Trichonectes]]'' <small>Guignot, 1941</small>
* ''[[Trogloguignotus]]'' <small>Sanfilippo, 1958</small>
* ''[[Tyndallhydrus]]'' <small>Sharp, 1882</small>
* ''[[Typhlodessus]]'' <small>Brancucci, 1985</small>
* ''[[Uvarus]]'' <small>Guignot, 1939</small>
* ''[[Vatellus]]'' <small>Aubé, 1837</small>
* ''[[yola (beetle)|Yola]]'' <small>Gozis, 1886</small>
* ''[[Yolina]]'' <small>Guignot, 1936</small>
* † ''[[Calicovatellus]]'' <small>K.B. Miller & Lubkin, 2001</small>
* † ''[[Procoelambus]]'' <small>Théobald, 1937</small>
{{div col end}}
Subfamily [[Laccophilinae]] <small>Gistel, 1856</small>{{Div col|colwidth=23em}}
* ''[[Africophilus]]'' <small>Guignot, 1948</small>
* ''[[Agabetes]]'' <small>Crotch, 1873</small>
* ''[[Australphilus]]'' <small>Watts, 1978</small>
* ''[[Japanolaccophilus]]'' <small>Satô, 1972</small>
* ''[[Laccodytes]]'' <small>Régimbart, 1895</small>
* ''[[Laccophilus]]'' <small>Leach, 1815</small>
* ''[[Laccoporus]]'' <small>J. Balfour-Browne, 1939</small>
* ''[[Laccosternus]]'' <small>Brancucci, 1983</small>
* ''[[Napodytes]]'' <small>Steiner, 1981</small>
* ''[[Neptosternus]]'' <small>Sharp, 1882</small>
* ''[[Philaccolilus]]'' <small>Guignot, 1937</small>
* ''[[Philaccolus]]'' <small>Guignot, 1937</small>
* ''[[Philodytes]]'' <small>J. Balfour-Browne, 1939</small>
{{div col end}}
Subfamily [[Lancetinae]] <small>Branden, 1885</small>
* ''[[Lancetes]]'' <small>Sharp, 1882</small>
Subfamily [[Matinae]] <small>Branden, 1885</small>
* ''[[Allomatus]]'' <small>Mouchamps, 1964</small>
* ''[[Batrachomatus]]'' <small>Clark, 1863</small>
* ''[[Matus (beetle)|Matus]]'' <small>Aubé, 1836</small>
Subfamily †[[Liadytiscinae]] <small>Prokin & Ren, 2010</small>
* † ''[[Liadroporus]]'' <small>Prokin & Ren, 2010</small> [[Yixian Formation]], China, Early Cretaceous ([[Aptian]])
* † ''[[Liadytiscus]]'' <small>Prokin & Ren, 2010</small> Yixian Formation, China, Aptian
* † ''[[Mesoderus]]'' <small>Prokin & Ren, 2010</small> Yixian Formation, China, Aptian
* † ''[[Liadyxianus]]'' <small>Prokin, Petrov, B. Wang & Ponomarenko, 2013</small> Yixian Formation, China, Aptian
* † ''[[Mesodytes]]'' <small>Prokin, Petrov, Wang & Ponomarenko, 2013</small> Yixian Formation, China, Aptian
Subfamily Incertae sedis
* † ''[[Cretodytes]]'' <small>Ponomarenko, 1977</small> [[Doronino Formation]], Russia, Early Cretaceous ([[Barremian]]), Kzyl-Zhar, Kazakhstan, Late Cretaceous ([[Turonian]])
* † ''[[Palaeodytes]]'' <small>Ponomarenko, 1987</small> [[Karabastau Formation]], Kazakhstan, Late Jurassic ([[Oxfordian (stage)|Oxfordian]]), [[Zaza Formation]], Russia, Aptian
* † ''"Palaeodytes" incompletus'' <small>Ponomarenko, Coram & Jarzembowski, 2005</small> [[Durlston Formation]], United Kingdom, Early Cretaceous ([[Berriasian]]) (undescribed genus)<ref>{{cite journal|last1=Prokin|first1=A.A.|last2=Petrov|first2=P.N.|last3=Wang|first3=B.|last4=Ponomarenko|first4=A.G.|year=2013|title=New fossil taxa and notes on the Mesozoic evolution of Liadytidae and Dytiscidae (Coleoptera)|journal=[[Zootaxa]]|volume=3666|issue=2|pages=137–159|doi=10.11646/zootaxa.3666.2.2|url=https://www.zin.ru/animalia/coleoptera/pdf/Prokin_Petrov_Wang_Ponomarenko_2013_Mesozoic_Liadytidae_Dytiscidae_z03666p159p594.pdf}}</ref>
* † ''[[Sinoporus]]'' <small>Prokin & Ren, 2010</small> Yixian Formation, China, Aptian

== References ==
{{Reflist| refs=
<ref name=itis>
{{Cite web| title=Dytiscidae Report
| url=https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=111963
| website=Integrated Taxonomic Information System
| access-date=2019-06-17
}}</ref>

<ref name=gbif>
{{Cite web| title=Dytiscidae
| url=https://www.gbif.org/species/7824
| website=GBIF
| access-date=2019-06-17
}}</ref>

<ref name=Nilsson>{{Cite report
| title = A World Catalogue of the Family Dytiscidae, or the Diving Beetles (Coleoptera, Adephaga)
| last1 = Nilsson
| first1 = A.N.
| url = http://www2.emg.umu.se/projects/biginst/andersn/World%20catalogue%20of%20Dytiscidae%202015.pdf
| access-date = 2019-06-18
| archive-url = https://web.archive.org/web/20190726055144/http://www2.emg.umu.se/projects/biginst/andersn/World%20catalogue%20of%20Dytiscidae%202015.pdf
| archive-date = 2019-07-26
| url-status = dead
}}</ref>

<ref name=Bouchard2011>
{{Cite journal
| title = Family-group names in Coleoptera (Insecta)
| date = 2011
| last1 = Bouchard | first1 = Patrice
| last2 = Bousquet | first2 = Yves
| last3 = Davies | first3 = Anthony E.
| last4 = Alonso-Zarazaga | first4 = Miguel A.
| last5 = Lawrence | first5 = John F.
| last6 = Lyal | first6 = Chris H. C.
| last7 = Newton | first7 = Alfred F.
| last8 = Reid | first8 = Chris A. M.
| last9 = Schmitt | first9 = Michael
| last10 = Ślipiński | first10 = S. Adam
| last11 = Smith | first11 = Andrew B. T.
| display-authors = 4
| journal = ZooKeys
| issue = 88| pages = 1–972
| issn = 1313-2989
| url = https://zookeys.pensoft.net/articles.php?id=4001
| doi = 10.3897/zookeys.88.807
| pmc=3088472
| pmid=21594053
| doi-access = free
| bibcode = 2011ZooK...88....1B
}}</ref>
}}
{{Refbegin}}
* {{aut|[[Commonwealth Scientific and Industrial Research Organisation]] (CSIRO)}} (2004): Water for a Healthy Country - [http://www.anbg.gov.au/cpbr/WfHC/Dytiscidae/index.html Family Dytiscidae]. Version of 2004-JUL-02. Retrieved 2008-AUG-04
* {{aut|De Foliart, Gene R.}} (2002): [https://web.archive.org/web/20080705034552/http://www.food-insects.com/book7_31/Chapter%2026%20Eastern%20Asia.htm Chapter 26 - Eastern Asia: China, Japan, and other countries]. ''In: The Human Use of Insects as a Food Resource: A Bibliographic Account in Progress''.
* {{aut|Jäch, Manfred A.}} (2003): Fried water beetles Cantonese style. ''American Entomologist'' '''49'''(1): 34–37. [https://web.archive.org/web/20081029184708/http://wcv.nhm-wien.ac.at/coleoptera/publications/183.pdf PDF fulltext]
* {{aut|Larson, D.J., Alarie, Y., and Roughley, R.E.}} (2000): Predaceous Diving Beetles (Coleoptera: Dytiscidae) of the Nearctic Region, with emphasis on the fauna of Canada and Alaska. NRC Research Press, Ottawa. {{ISBN|978-0-660-17967-4}}.
{{Refend}}

== External links ==
{{Refbegin}}
* {{Commons-inline| category|Dytiscidae}}
* {{Wikispecies-inline|Dytiscidae}}
{{Refend}}

{{Coleoptera|2}}
{{Taxonbar|from=Q327149}}
{{Authority control}}

{{DEFAULTSORT:Dytiscidae}}
[[Category:Dytiscidae| ]]
[[Category:Beetle families]]
[[Category:Aquatic insects]]