The Dytiscidae – based on the Greek ''dytikos'' (δυτικός), "able to dive" – are the predaceous diving beetles, a
family
Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
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 long, though much variation is seen between species. The European ''
Dytiscus latissimus'' and Brazilian ''
Megadytes ducalis'' are the largest, reaching up to and respectively.
[ In contrast, the smallest is likely the Australian '' Limbodessus atypicali'' of subterranean waters, which only is about long.][ Most are dark brown, blackish, or dark ]olive
The olive, botanical name ''Olea europaea'', meaning 'European olive' in Latin, is a species of small tree or shrub in the family Oleaceae, found traditionally in the Mediterranean Basin. When in shrub form, it is known as ''Olea europaea'' ' ...
in color with golden highlights in some subfamilies. The larva
A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle.
...
e are commonly known as water tigers due to their voracious appetite. They have short, but sharp mandible
In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
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 biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
in numerous genera
Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
.
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
''Agabus uliginosus'' is a species of beetle native to the Palearctic, including Europe, where it is only found in Austria, Belarus, Belgium, Great Britain including Shetland, Orkney, Hebrides and Isle of Man, Croatia, the Czech Republic, ...
'' 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.
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
The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the c ...
, 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
A tadpole is the larval stage in the biological life cycle of an amphibian. Most tadpoles are fully aquatic, though some species of amphibians have tadpoles that are terrestrial. Tadpoles have some fish-like features that may not be found ...
s 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
A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in thei ...
. 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
''Cybister'' ('kybistētēr' = diver, tumbler), is a genus of beetle in family Dytiscidae. They are found in much of the world, including all continents except Antarctica. As of 2021 there are 96 species and 9 additional subspecies among four su ...
'', are edible. Remnants of '' C. explanatus'' were found in prehistoric human coprolite
A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name is ...
s in a Nevada cave, likely sourced from the Humboldt Sink
The Humboldt Sink is an intermittent dry lake bed, approximately 11 mi (18 km) long, and 4 mi (6 km) across, in northwestern Nevada in the United States. The body of water in the sink is known as Humboldt Lake. The sink and it ...
. In Mexico
Mexico (Spanish language, Spanish: México), officially the United Mexican States, is a List of sovereign states, country in the southern portion of North America. It is borders of Mexico, bordered to the north by the United States; to the so ...
, ''C. explanatus'' is eaten roasted and salted to accompany taco
A taco (, , ) is a traditional Mexican food consisting of a small hand-sized corn- or wheat-based tortilla topped with a filling. The tortilla is then folded around the filling and eaten by hand. A taco can be made with a variety of fillin ...
s. In Japan, '' C. japonicus'' has been used as food in certain regions such as Nagano prefecture. In the Guangdong Province
Guangdong (, ), alternatively romanized as Canton or Kwangtung, is a coastal province in South China on the north shore of the South China Sea. The capital of the province is Guangzhou. With a population of 126.01 million (as of 2020) ...
of China
China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
, 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
The great diving beetle (''Dytiscus marginalis'') is an aquatic diving beetle native to Europe and northern Asia, and in the UK is common in Wales, much of England and southern Scotland but less common on chalk and in the far north.
The grea ...
(''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
Taiwan, officially the Republic of China (ROC), is a country in East Asia, at the junction of the East and South China Seas in the northwestern Pacific Ocean, with the People's Republic of China (PRC) to the northwest, Japan to the no ...
, Thailand
Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is b ...
, and New Guinea
New Guinea (; Hiri Motu: ''Niu Gini''; id, Papua, or , historically ) is the world's second-largest island with an area of . Located in Oceania in the southwestern Pacific Ocean, the island is separated from Australia by the wide Torres ...
.
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.
Dytiscid adults are eaten by many bird
Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s, mammal
Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
s, reptile
Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates ( lizards and snakes) and rhynchocephalia ...
s, and other vertebrate predator
Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill t ...
s, 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 ]pond
A pond is an area filled with water, either natural or artificial, that is smaller than a lake. Defining them to be less than in area, less than deep, and with less than 30% emergent vegetation helps in distinguishing their ecology from tha ...
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 , and thus serve as umbrella species
Umbrella species are species selected for making conservation-related decisions, typically because protecting these species indirectly protects the many other species that make up the ecological community of its habitat (the umbrella effect). Speci ...
for the protection of natural aquatic habitats: '' Dytiscus latissimus'' and '' Graphoderus bilineatus''.
Cultural significance
The diving beetle plays a role in a Cherokee
The Cherokee (; chr, ᎠᏂᏴᏫᏯᎢ, translit=Aniyvwiyaʔi or Anigiduwagi, or chr, ᏣᎳᎩ, links=no, translit=Tsalagi) are one of the indigenous peoples of the Southeastern Woodlands of the United States. Prior to the 18th century, th ...
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
East Africa, Eastern Africa, or East of Africa, is the eastern subregion of the African continent. In the United Nations Statistics Division scheme of geographic regions, 10-11-(16*) territories make up Eastern Africa:
Due to the historica ...
. 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
Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear e ...
. Those in genera '' Dytiscacarus'' and ''Eylais
''Eylais'' is a genus of mites belonging to the family Eylaidae.
The genus has cosmopolitan distribution.
Species:
* '' Eylais abitibiensis'' Marshall, 1929
* '' Eylais amplipons'' Viets
Biology
''Eylais'' mite larvae are ectoparasites of d ...
'' 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
Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
.[
Subfamily ]Agabinae
Agabinae is a subfamily of predaceous diving beetles in the family Dytiscidae. There are about 11 genera and more than 460 described species in Agabinae.
Genera
These 11 genera belong to the subfamily Agabinae:
* '' Agabinus'' Crotch, 1873
* ' ...
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
Colymbetinae is a subfamily of predaceous diving beetles in the family Dytiscidae. There are about 11 genera and at least 130 described species in Colymbetinae.
Genera
These 11 genera belong to the subfamily Colymbetinae:
* '' Anisomeria'' Bri ...
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
* '' 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
Subfamily Coptotominae
''Coptotomus'' is a genus of predaceous diving beetles in the family Dytiscidae, the only genus of the subfamily Coptotominae. There are about six described species in ''Coptotomus'', found in North America and the Neotropics.
There is one exti ...
Branden, 1885
* '' Coptotomus'' Say, 1830
Subfamily Cybistrinae
* '' Austrodytes'' Watts, 1978
* ''Cybister
''Cybister'' ('kybistētēr' = diver, tumbler), is a genus of beetle in family Dytiscidae. They are found in much of the world, including all continents except Antarctica. As of 2021 there are 96 species and 9 additional subspecies among four su ...
'' Curtis, 1827
* '' Megadytes'' Sharp, 1882
* '' Onychohydrus'' Schaum & White, 1847
* '' Regimbartina'' Chatanay, 1911
* '' Spencerhydrus'' Sharp, 1882
* '' Sternhydrus'' Brinck, 1945
Subfamily Dytiscinae Leach, 1815
* '' 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
Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. The ...
, Myanmar, Late Cretaceous (Cenomanian
The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in ...
)
Subfamily Hydrodytinae K.B.Miller, 2001
* '' Hydrodytes'' K.B.Miller, 2001
* '' Microhydrodytes'' K.B.Miller, 2002
Subfamily Hydroporinae Aubé, 1836
* '' 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
* '' 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
* † '' Kuschelydrus'' Ordish, 1976
* † '' 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
* '' Lancetes'' Sharp, 1882
Subfamily Matinae Branden, 1885
* '' Allomatus'' Mouchamps, 1964
* '' Batrachomatus'' Clark, 1863
* '' Matus'' Aubé, 1836
Subfamily † Liadytiscinae Prokin & Ren, 2010
* † '' Liadroporus'' Prokin & Ren, 2010 Yixian Formation, China, Early Cretaceous (Aptian
The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ag ...
)
* † '' 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
Subfamily Incertae sedis
* † '' Cretodytes'' Ponomarenko, 1977 Doronino Formation, Russia, Early Cretaceous (Barremian
The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma (million years ago) and 121.4 ± 1.0 Ma). It is a subdivision of the Early Cretaceous Epoch (or Lower Cretaceous Series). It is preceded ...
), Kzyl-Zhar, Kazakhstan, Late Cretaceous (Turonian
The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded b ...
)
* † '' Palaeodytes'' Ponomarenko, 1987 Karabastau Formation, Kazakhstan, Late Jurassic ( Oxfordian), Durlston Formation
The Durlston Formation is a geologic formation in England. Particularly in the Isle of Purbeck. It preserves fossils dating back to the Berriasian stage
Stage or stages may refer to:
Acting
* Stage (theatre), a space for the performance of ...
, United Kingdom, Early Cretaceous (Berriasian), Zaza Formation, Russia, Aptian
* † '' Sinoporus'' Prokin & Ren, 2010 Yixian Formation, China, Aptian
References
* (2004): Water for a Healthy Country
Family Dytiscidae
Version of 2004-JUL-02. Retrieved 2008-AUG-04
* (2002)
''In: The Human Use of Insects as a Food Resource: A Bibliographic Account in Progress''.
* (2003): Fried water beetles Cantonese style. ''American Entomologist'' 49(1): 34-37
PDF fulltext
* (2000): Predaceous Diving Beetles (Coleoptera: Dytiscidae) of the Nearctic Region, with emphasis on the fauna of Canada and Alaska. NRC Research Press, Ottawa. .
External links
*
*
{{DEFAULTSORT:Dytiscidae
Beetle families
Aquatic insects