Dytiscidae

The Dytiscidae – based on the Greek dytikos (δυτικός), "able to dive" – are the predaceous diving beetles, a family of water beetles. They occur in virtually any freshwater habitat around the world, but a few species live among leaf litter. The adults of most are between 1 and(-) long, though much variation is seen between species. The European Dytiscus latissimus and Brazilian Megadytes ducalis are the largest, reaching up to 4.5 cm and 4.75 cm respectively. In contrast, the smallest is likely the Australian Limbodessus atypicali of subterranean waters, which only is about 0.9 mm long. Most are dark brown, blackish, or dark olive in color with golden highlights in some subfamilies. The larvae are commonly known as water tigers due to their voracious appetite. They have short, but sharp mandibles 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.

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. Diving beetles live in water bodies in various landscapes, including agricultural and urban landscapes. Some species, such as Agabus uliginosus and Acilius canaliculatus, 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. Some species, such as Oreodytes sanmarkii, occur in exposed areas of waters, whereas many diving beetles species prefer habitats with aquatic plants, especially plants with complex structures, such as sedges and bulrush.

Larvae and development
When still in larval form, the beetles vary in size from about 1 to 5 cm (0.5 to 2.0 in). 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 tadpoles and glassworms, 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 edible. Remnants of C. explanatus were found in prehistoric human coprolites in a Nevada cave, likely sourced from the Humboldt Sink. In Mexico, C. explanatus is eaten roasted and salted to accompany tacos. In Japan, 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 C. bengalensis, C. guerini, C. limbatus, C. sugillatus, 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.

Diving beetle conservation
The greatest threat to diving beetles is the degradation and disappearance of their habitats due to anthropogenic activities. For example, urbanisation has led to the decreasing quantity and quality of dytiscid habitats, which consequentially has increased the distance between habitats. ; 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. 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. 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.

Dytiscid adults are eaten by many birds, mammals, reptiles, and other vertebrate predators, despite their arsenal of chemical defenses. 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 predators in small ponds, 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.

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. 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.

Parasites
Dytiscidae are parasitised by various mites. Those in genera Dytiscacarus and Eylais live beneath the elytra of their hosts, those in genus Acherontacarus attach to the mesosternal regions and those in genus Hydrachna attach to various locations. These mites are parasitic as larvae with the exception of Dytiscacarus, which are parasitic for their entire life cycle.

Systematics
The following taxonomic sequence gives the subfamilies, their associated genera.

Subfamily Agabinae Thomson, 1867
 * Agabinus Crotch, 1873
 * Agabus Leach, 1817
 * Agametrus Sharp, 1882
 * Andonectes Guéorguiev, 1971
 * Hydronebrius Jakovlev, 1897
 * Hydrotrupes Sharp, 1882
 * Ilybiosoma Crotch, 1873
 * Ilybius Erichson, 1832
 * Leuronectes Sharp, 1882
 * Platambus Thomson, 1859
 * Platynectes Régimbart, 1879

Subfamily Colymbetinae Erichson, 1837
 * Anisomeria Brinck, 1943
 * Senilites Brinck, 1948
 * Carabdytes Balke, Hendrich & Wewalka, 1992
 * Bunites Spangler, 1972
 * Colymbetes Clairville, 1806
 * Hoperius Fall, 1927
 * Meladema Laporte, 1835
 * Melanodytes Seidlitz, 1887
 * Neoscutopterus J.Balfour-Browne, 1943
 * Rhantus Dejean, 1833
 * Rugosus García, 2001

Subfamily Copelatinae Branden, 1885 Subfamily Coptotominae Branden, 1885 Subfamily Cybistrinae Subfamily Dytiscinae Leach, 1815
 * Agaporomorphus Zimmermann, 1921
 * Aglymbus Sharp, 1880
 * Copelatus Erichson, 1832
 * Exocelina Broun, 1886
 * Lacconectus Motschulsky, 1855
 * Liopterus Dejean, 1833
 * Madaglymbus Shaverdo & Balke, 2008
 * Rugosus García, 2001
 * Coptotomus Say, 1830
 * Austrodytes Watts, 1978
 * Cybister Curtis, 1827
 * Megadytes Sharp, 1882
 * Onychohydrus Schaum & White, 1847
 * Regimbartina Chatanay, 1911
 * Spencerhydrus Sharp, 1882
 * Sternhydrus Brinck, 1945


 * Acilius Leach, 1817
 * Aethionectes Sharp, 1882
 * Austrodytes Watts, 1978
 * Dytiscus Linnaeus, 1758
 * Eretes Laporte, 1833
 * Graphoderus Dejean, 1833
 * Hydaticus Leach, 1817
 * Hyderodes Hope, 1838
 * Megadytes Sharp, 1882
 * Miodytiscus Wickham, 1911
 * Notaticus Zimmermann, 1928
 * Onychohydrus Schaum & White, 1847
 * Regimbartina Chatanay, 1911
 * Rhantaticus Sharp, 1880
 * Sandracottus Sharp, 1882
 * Spencerhydrus Sharp, 1882
 * Sternhydrus Brinck, 1945
 * Thermonectus Dejean, 1833
 * Tikoloshanes Omer-Cooper, 1956
 * †Ambarticus Yang et al. 2019 Burmese amber, Myanmar, Late Cretaceous (Cenomanian)

Subfamily Hydrodytinae K.B.Miller, 2001 Subfamily Hydroporinae Aubé, 1836
 * Hydrodytes K.B.Miller, 2001
 * Microhydrodytes K.B.Miller, 2002
 * Africodytes Biström, 1988
 * Agnoshydrus Biström, Nilsson & Wewalka, 1997
 * Allodessus Guignot, 1953
 * Allopachria Zimmermann, 1924
 * Amarodytes Régimbart, 1900
 * Amurodytes Fery & Petrov, 2013
 * Andex Sharp, 1882
 * Anginopachria Wewalka, Balke & Hendrich, 2001
 * Anodocheilus Babington, 1841
 * Antiporus Sharp, 1882
 * Barretthydrus Lea, 1927
 * Bidessodes Régimbart, 1895
 * Bidessonotus Régimbart, 1895
 * Bidessus Sharp, 1882
 * Boreonectes Angus, 2010
 * Borneodessus Balke, Hendrich, Mazzoldi & Biström, 2002
 * Brachyvatus Zimmermann, 1919
 * Brancuporus Hendrich, Toussaint & Balke, 2014
 * Canthyporus Zimmermann, 1919
 * Carabhydrus Watts, 1978
 * Celina Aubé, 1837
 * Chostonectes Sharp, 1880
 * Clypeodytes Régimbart, 1894
 * Coelhydrus Sharp, 1882
 * Comaldessus Spangler & Barr, 1995
 * Crinodessus K.B. Miller, 1997
 * Darwinhydrus Sharp, 1882
 * Deronectes Sharp, 1882
 * Derovatellus Sharp, 1882
 * Desmopachria Babington, 1841
 * Dimitshydrus Uéno, 1996
 * Ereboporus K.B. Miller, Gibson & Alarie, 2009
 * Etruscodytes Mazza, Cianferoni & Rocchi, 2013
 * Fontidessus K.B. Miller & Spangler, 2008
 * Geodessus Brancucci, 1979
 * Gibbidessus Watts, 1978
 * Glareadessus Wewalka & Biström, 1998
 * Graptodytes Seidlitz, 1887
 * Haideoporus Young & Longley, 1976
 * Hemibidessus Zimmermann, 1921
 * Heroceras Guignot, 1949
 * Herophydrus Sharp, 1880
 * Heterhydrus Fairmaire, 1869
 * Heterosternuta Strand, 1935
 * Hovahydrus Biström, 1982
 * Huxelhydrus Sharp, 1882
 * Hydrocolus Roughley & Larson in Larson, Alarie & Roughley, 2000
 * Hydrodessus J. Balfour-Browne, 1953
 * Hydroglyphus Motschulsky, 1853
 * Hydropeplus Sharp, 1882
 * Hydroporus Clairville, 1806
 * Hydrovatus Motschulsky, 1853
 * Hygrotus Stephens, 1828
 * Hyphoporus Sharp, 1880
 * Hyphovatus Wewalka & Biström, 1994
 * Hyphydrus Illiger, 1802
 * Hypodessus Guignot, 1939
 * Iberoporus Castro & Delgado, 2001
 * Incomptodessus K.B. Miller & García, 2011
 * Kakadudessus Hendrich & Balke, 2009
 * Kuschelydrus Ordish, 1976
 * Laccornellus Roughley & Wolfe, 1987
 * Laccornis Gozis, 1914
 * Leiodytes Guignot, 1936
 * Limbodessus Guignot, 1939
 * Liodessus Guignot, 1939
 * Lioporeus Guignot, 1950
 * Megaporus Brinck, 1943
 * Metaporus Guignot, 1945
 * Methles Sharp, 1882
 * Microdessus Young, 1967
 * Microdytes J. Balfour-Browne, 1946
 * Morimotoa Uéno, 1957
 * Nebrioporus Régimbart, 1906
 * Necterosoma W.J. Macleay, 1871
 * Neobidessodes Hendrich & Balke, 2009
 * Neobidessus Young, 1967
 * Neoclypeodytes Young, 1967
 * Neoporus Guignot, 1931
 * Oreodytes Seidlitz, 1887
 * Pachydrus Sharp, 1882
 * Pachynectes Régimbart, 1903
 * Papuadessus Balke, 2001
 * Paroster Sharp, 1882
 * Peschetius Guignot, 1942
 * Petrodessus K.B. Miller, 2012
 * Phreatodessus Ordish, 1976
 * Platydytes Biström, 1988
 * Porhydrus Guignot, 1945
 * Primospes Sharp, 1882
 * Pseuduvarus Biström, 1988
 * Psychopomporus Jean, Telles & K.B. Miller, 2012
 * Pteroporus Guignot, 1933
 * Queda Sharp, 1882
 * Rhithrodytes Bameul, 1989
 * Sanfilippodytes Franciscolo, 1979
 * Scarodytes Gozis, 1914
 * Schistomerus Palmer, 1957
 * Sekaliporus Watts, 1997
 * Sharphydrus Omer-Cooper, 1958
 * Siamoporus Spangler, 1996
 * Siettitia Abeille de Perrin, 1904
 * Sinodytes Spangler, 1996
 * Spanglerodessus K.B. Miller & García, 2011
 * Sternopriscus Sharp, 1880
 * Stictonectes Brinck, 1943
 * Stictotarsus Zimmermann, 1919
 * Stygoporus Larson & LaBonte, 1994
 * Suphrodytes Gozis, 1914
 * Tepuidessus Spangler, 1981
 * Terradessus Watts, 1982
 * Tiporus Watts, 1985
 * Trichonectes Guignot, 1941
 * Trogloguignotus Sanfilippo, 1958
 * Tyndallhydrus Sharp, 1882
 * Typhlodessus Brancucci, 1985
 * Uvarus Guignot, 1939
 * Vatellus Aubé, 1837
 * Yola Gozis, 1886
 * Yolina Guignot, 1936
 * † Calicovatellus K.B. Miller & Lubkin, 2001
 * † Procoelambus Théobald, 1937

Subfamily Laccophilinae Gistel, 1856
 * Africophilus Guignot, 1948
 * Agabetes Crotch, 1873
 * Australphilus Watts, 1978
 * Japanolaccophilus Satô, 1972
 * Laccodytes Régimbart, 1895
 * Laccophilus Leach, 1815
 * Laccoporus J. Balfour-Browne, 1939
 * Laccosternus Brancucci, 1983
 * Napodytes Steiner, 1981
 * Neptosternus Sharp, 1882
 * Philaccolilus Guignot, 1937
 * Philaccolus Guignot, 1937
 * Philodytes J. Balfour-Browne, 1939

Subfamily Lancetinae Branden, 1885 Subfamily Matinae Branden, 1885 Subfamily †Liadytiscinae Prokin & Ren, 2010 Subfamily Incertae sedis
 * Lancetes Sharp, 1882
 * Allomatus Mouchamps, 1964
 * Batrachomatus Clark, 1863
 * Matus Aubé, 1836
 * † Liadroporus Prokin & Ren, 2010 Yixian Formation, China, Early Cretaceous (Aptian)
 * † Liadytiscus Prokin & Ren, 2010 Yixian Formation, China, Aptian
 * † Mesoderus Prokin & Ren, 2010 Yixian Formation, China, Aptian
 * † Liadyxianus Prokin, Petrov, B. Wang & Ponomarenko, 2013 Yixian Formation, China, Aptian
 * † Mesodytes Prokin, Petrov, Wang & Ponomarenko, 2013 Yixian Formation, China, Aptian
 * † Cretodytes Ponomarenko, 1977 Doronino Formation, Russia, Early Cretaceous (Barremian), Kzyl-Zhar, Kazakhstan, Late Cretaceous (Turonian)
 * † Palaeodytes Ponomarenko, 1987 Karabastau Formation, Kazakhstan, Late Jurassic (Oxfordian), Zaza Formation, Russia, Aptian
 * † "Palaeodytes" incompletus Ponomarenko, Coram & Jarzembowski, 2005 Durlston Formation, United Kingdom, Early Cretaceous (Berriasian) (undescribed genus)
 * † Sinoporus Prokin & Ren, 2010 Yixian Formation, China, Aptian