Hydrophilidae

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Hydrophilidae
Temporal range: Late Jurassic–Recent
Grosser Kolbenwasserkafer Hydrous piceus 1.jpg
Hydrophilus piceus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Suborder: Polyphaga
Infraorder: Staphyliniformia
Superfamily: Hydrophiloidea
Family: Hydrophilidae
Latreille, 1802
Genera

169

Hydrophilidae, also known colloquially as water scavenger beetles, is a family of beetles. [1] [2] Aquatic hydrophilids are notable for their long maxillary palps, which are longer than their antennae. [3] Several of the former subfamilies of Hydrophilidae have recently been removed and elevated to family rank; Epimetopidae, Georissidae (= Georyssidae), Helophoridae, Hydrochidae, and Spercheidae. [1] [4] While the majority of hydrophilids are aquatic, around a third of described species are terrestrial, mostly belonging to the subfamily Sphaeridiinae. [5] [6]

Contents

With rare exceptions, the larvae are predatory while the adults may be herbivores or predators in addition to scavenging. [7] Many species are able to produce sounds. [8]

Species of Hydrophilus are reported as pests in fish hatcheries. [7] Other species are voracious consumers of mosquito larvae, and have potential as biological control agents. [7] [9]

This beetle family contains 2,835 species in 169 genera. [10]

Geography

Hydrophilid beetles are found worldwide. [5] In the Americas, they are usually found in areas of high humidity such as the tropics of Central and South America. They can also be found in rain pools and ponds in the forests of Guatemala and Argentina. [11] They also tend to exist in North America in areas with seasonal wetlands or lakes depending on the state. [12] Hydrophilus triangularis is found widely throughout the United States and is the biggest water beetle in the country. The family Hydrophilidae has been reported in every state in the United States. [13]

Evolutionary history

The oldest known fossils definitively assignable to the family are from the Late Jurassic Solnhofen of Germany and Talbragar Fish Bed in Australia. [14]

Habitat

A majority of the beetles in the family Hydrophilidae live in aquatic environments in both their larval and adult life stages. Some hydrophilid beetles will lay their eggs in ephemeral ponds and puddles where the larvae will live as they develop. [15] Other beetles such as Derralus angustus and Tropisternus setiger live in permanent ponds. Some beetles such as Tropisternus lateralis will only live in aquatic environments that lack fish because fish prey on their eggs, while others like D. angustus prefer aquatic habitats with a specific species of floating fern. [16] Generally, hydrophilids live in marshy, shallow, and heavily weeded aquatic environments. [17] There are some hydrophilid beetles that make their homes in fresh animal waste, decaying vegetation, or humus-rich soil. [18] They survive in a very wide variety of locations and because of that some types are more adapted to specific environments than others and will often only move to habitats of the same type. [19]

Physiology

Reproduction in hydrophilids takes place in bodies of water such as ponds. In the larval stage the beetle resides in a shallow area of the pond because they are dependent on the oxygen only available in the shallower areas. After the beetle exits the pupa stage they often take flight and move to a new area before they reproduce. [17] Wing growth depends on the environment that the beetle resides in. Beetles in lentic habitats have better wing development compared to lotic habitats because lentic habitats are less reliable and require the beetles to disperse quicker in order to survive. [20] Some beetles such as Berosus larvae can inhabit areas deeper in the water due to their thoracic gills while others like Berosus ingeminatus use cutaneous respiration allowing them to hold air for longer periods of time. The ability to consume oxygen at deeper water levels helps them avoid being preyed upon from surface predators. [17] Larvae use two main types of feeding mechanism, chewing and piercing-sucking, with the latter evolving at least three times independently in Hydrophilidae and once in Epimetopidae and allowing underwater extra-oral digestion, decreasing dependence on aerial environments. [21] Adult hydrophilid beetles can survive in deeper areas of water and stay under for longer due to their special abilities in acquiring oxygen. They can hold air bubbles under their elytra that connect to their spiracles for them to use the trapped oxygen. [22] They also have many fine hairs along their bodies that can trap oxygen. They can pull oxygen from the water into these areas on their body so that they can avoid returning to the surface for long periods of time. They only return to the surface when they need to replace the air they have acquired. [23]

Reproduction

Some species of hydrophilid beetles in the genus Tropisternus have complex methods of signaling and communication including chirps, clicks, buzzing, and various body postures. These behaviors can assist in courtship. Courtship behaviors have been observed in Tropisternus ellipticus in which the females produced audible buzzing and shaking to rebuff potential suitors. In response to the rejection maneuvers of the female, male T. ellipticus will mimic the buzzing and shaking, often learning from repeated encounters of this kind to avoid females that produce these behaviors. Males may remember these encounters for up to 39 minutes when properly reinforced, which suggests that they have some capability for recording short-term memory. [24] Typical courtship in these beetles consists of the following steps:

1. The male approaches the female, buzzing and swimming around her.

2. If the female is receptive, she will either chirp in response and move toward the male or remain still and silent. No rejection behavior occurs at this time.

3. The male approaches further and mounts the female.

4. The male will touch his maxillary palps to that of the female while producing a buzzing sound.

5. If the female does not rebuff his advances, the male with move to the back and probe the abdomen of the female with his aedeagus.

6. If she is receptive, the female must lower her abdomen for the male to penetrate.

Males mount other beetles indiscriminately, and homosexual copulation has been known to occur. [24]

Anacaena lutescens , a species of hydrophilid beetle, has been observed as reproducing via parthenogenesis. Reproduction by parthenogenesis is relatively uncommon in other types of insects. [25]

Life stages

In Enochrus quadripunctatus, a species of hydrophilid beetle, it takes an average of 43 days for a newly hatched larva to reach its adult form. Juvenile mortality is high in this species, decreasing with each successive instar. [26]

Females will bury their silken egg case in the damp soil near a stream. Typically, the female will only create and lay one egg case per day, though they may create up to 20 egg cases in the weeks following a mating event and each case may produce from 1-32 individual larva. Females who have not mated will still create egg cases, but they will typically be empty or will not hatch at all. Not all egg cases laid by a mated female will hatch and produce viable offspring, but 42-70% will. Larval forms are carnivorous and may exhibit cannibalism among the individuals in a single egg case before hatching. The eggs begin to hatch an average of eight days after being laid. The egg hatches by way of the larvae eating their way out of the casing, usually by way of the lid. This process can take several days in some instances. Once the egg hatches, the first instar larvae will disperse, but will continue to feed on one another if the opportunity presents itself. They will remain in the first instar form for an average of 9.5 days before progressing to the second instar. Second instar individuals are much more robust, and voraciously feed on what meat they can find. They are strong swimmers at this stage and can be found at or beneath the surface of the stream. After an average of 11 days, they will transition to their third instar form. The third instar will last an average of 8.4 days, during which the larvae will continue feeding and become progressively slower as they reach their pupation stage. When they are ready to pupate, the larva will burrow into the sand where they will remain for 14 days before emerging as an adult. After vacating their burrow, the new adult will stand on the sand for up to 24 hours to allow its carapace to harden. Once fully hardened, the new fully grown adult beetle will travel to the water to hunt, where it will remain for the rest of its life. [26]

Although individuals of each species of hydrophilid (even those within the same genus) may vary in the duration of each of these life stages, mortality rates at each stage, and the number of offspring produced, few studies have been done to this effect on other species of hydrophilid. [26]

Diet

Larval hydrophilids are predatory by nature and different species have different food consumption habits. The larvae usually ingest small invertebrates and snails but have been known to also eat small fish and tadpoles. [27] Berosus ingeminatus prey mostly on Cricotopus sylvestris and other types of midges while Tropisternus setiger are tactile hunters and will eat whatever prey comes their way; they may even abandon a meal for a new one if the opportunity arises. [11] The predatory habits of this species are often left behind in the larval stage. Generally adult hydrophilids feed on various vegetation, alive or decaying, and will occasionally eat dead animal tissue. [17]

Predators

Hydrophilids have many predators from a range of different species. Fish, birds, predaceous insects, and turtles are the main predators in their aquatic environments. Humans have also been known to eat hydrophilid beetles. [28]

Behavior

Acoustic behavior

Member of the genus Tropisternus have been shown to perform acoustic sounds in their communicative behavior. Specific calls include stress calls, a male courtship call, a male copulating sound, and a female rejection buzz. [8] The sound production comes from the friction created by the beetle rubbing its spectrum (a well- defined ridge or lip) to its finely ridged surface called a pars stridens on the beetle's underside. [29]

Predatory behavior

Larval hydrophilids hunt a wide variety of prey such as copepods, mosquito larvae, snails, and conspecifics. [27] The larvae have a unique way of hunting by lifting their prey out of the water to consume it. The reason for this is not well known, but there is a suggestion that lifting the prey makes it more difficult for the prey to escape. [27] In addition to lifting their prey out of the water, hydrophilid larvae choose specific vegetation to wait in so that they may ambush their prey. [30] In addition, the larvae often change ambush sites according to prey density. The frequency is directly correlated to the amount of prey in the ambush areas and specific attack sites. [30]

Subfamilies

The classification to subfamilies and tribes was revised by Short and Fikáček in 2013: [5]

See also

Bibliography

Related Research Articles

<i>Spercheus</i> Genus of beetles

Spercheus is a genus of aquatic beetles which are placed in a family of their own, Spercheidae within the Hydrophiloidea. About 20 species are known from around the world except the Nearctic with the majority being from the Oriental and Afrotropical Realms.

<span class="mw-page-title-main">Telephone-pole beetle</span> Species of beetle

The telephone-pole beetle is a beetle native to the eastern United States, and the only living representative of the otherwise extinct family Micromalthidae. Larvae of the beetle live in decaying wood and can be pests to wooden structures, lending them their common name, the 'telephone-pole beetle.'

<span class="mw-page-title-main">Histeridae</span> Family of beetles

Histeridae is a family of beetles commonly known as clown beetles or hister beetles. This very diverse group of beetles contains 3,900 species found worldwide. They can be easily identified by their shortened elytra that leaves two of the seven tergites exposed, and their geniculate (elbowed) antennae with clubbed ends. These predatory feeders are most active at night and will fake death if they feel threatened. This family of beetles will occupy almost any kind of niche throughout the world. Hister beetles have proved useful during forensic investigations to help in time of death estimation. Also, certain species are used in the control of livestock pests that infest dung and to control houseflies. Because they are predacious and will even eat other hister beetles, they must be isolated when collected.

<span class="mw-page-title-main">Hydrophiloidea</span> Superfamily of beetles

Hydrophiloidea, known as water scavenger beetles, is a superfamily of beetles. Until recently it included only a single family, the Hydrophilidae, but several of the subfamilies have been removed and raised to family rank. Hydrophiliidae remains by far the largest member of the group, with nearly 3,000 described species. The other families have no more than 400 species. The Histeroidea are closely related and sometimes considered part of a sensu lato Hydrophiloidea. The majority of the clade is aquatic, which is thought to be the ancestral ecology of the group, with some lineages like Sphaeridiinae becoming secondarily terrestrial. Modern representatives of the group first appeared during the Late Jurassic.

<i>Thermonectus marmoratus</i> Species of beetle

Thermonectus marmoratus is a relatively colorful North American species of diving beetle known by the common names sunburst diving beetle and yellow-spotted diving beetle. The behavior of this diving beetle has been compared to a scuba diver, since it carries with it a bubble of air as it dives down into the water. Its aquatic larval stage was the first ever recorded use of bifocals in the animal world. The beetle uses in its principal eyes two retinas and two distinct focal planes that are substantially separated, in the manner of bifocals to switch their vision from up-close to distance, for easy and efficient capture of their prey.

<i>Hydrophilus</i> (beetle) Genus of beetles

Hydrophilus is a genus of beetles in the family Hydrophilidae, the water scavenger beetles. There are 53 species in three subgenera in the genus: Hydrophilus, Dibolocelus, and Temnopterus.

<i>Hydrophilus palpalis</i> Species of beetle

Hydrophilus palpalis is a species of water scavenger beetles belonging to the Hydrophilinae subfamily.

<i>Tropisternus</i> Genus of beetles

Tropisternus is a genus of hydrophilid beetles with 63 species in five subgenera in North and South America.

<i>Tropisternus lateralis</i> Species of beetle

Tropisternus lateralis is a species of hydrophilid beetle that ranges across much of the Americas.

<span class="mw-page-title-main">Hydrobiusini</span> Tribe of beetles

Hydrobiusini is a tribe in the subfamily Hydrophilinae of aquatic beetles, and containing 47 species in 9 genera.

<span class="mw-page-title-main">Acidocerinae</span> Subfamily of beetles

Acidocerinae is a subfamily in the family Hydrophilidae of aquatic beetles, and it contains over 500 species in 23 genera.

<i>Cymbiodyta bifidus</i> Species of beetle

Cymbiodyta bifidus is a species of hydrophilid beetle. The species was formerly the sole member of the genus Helocombus.

<span class="mw-page-title-main">Sphaeridiinae</span> Subfamily of beetles

Sphaeridiinae is a subfamily of water scavenger beetles. Some species live in fresh water as both larvae and adults.

<i>Hydrobiomorpha</i> Genus of beetles

Hydrobiomorpha is a genus of water scavenger beetles in the family Hydrophilidae. There are 56 extant described species in Hydrobiomorpha, along with several fossil species.

<i>Rivacindela hudsoni</i> Species of beetle

Rivacindela hudsoni is an Australian species of the family Cicindelinae or "tiger beetle" and is the fastest running insect. The genus Rivacindela is contentiously treated as a subgenus of the broader Cicindela and are typically found in salty habitats such as dry salt lakes and salt streams and cannot fly far. The species was discovered in South Australia and described in 1997, with an adult form of approximately 20–21mm in length and a running speed of 2.49m/s, or 120 body lengths per second.

Hydrophilus ensifer is a species of water scavenger beetle in the family Hydrophilidae found in the Americas. As of 2020, there are two valid subspecies of H. ensifer, H. e. ensifer and H. e. duvali, however the differences among the subspecies are not well understood.

Cercyon (Clinocercyon) lineolatus, is a species of water scavenger beetle found in several Oriental and Oceanian countries.

Coelostoma (Holocoelostoma) stultum, is a species of water scavenger beetle widely distributed in Palearctic and Oriental realms from West Pacific towards Indian Ocean, such as China, Taiwan, Andaman Islands, Myanmar, India, Indonesia, Japan, Korea, Malaysia, Mascarene Islands, Nicobar Islands, Oman, Philippines, Saudi Arabia, South Korea, Sri Lanka, Thailand, United Arab Emirates, Oman, and Vietnam.

<i>Coelostoma vitalisi</i> Species of beetle

Coelostoma (Coelostoma) vitalisi, is a species of water scavenger beetle found in China, Taiwan, Japan, India, Indonesia, Japan, Malaysia, Nepal, Singapore, Sri Lanka, Cambodia, Thailand and Vietnam.

Regimbartia attenuata, commonly known as Japanese water scavenger beetle, is a species of water scavenger beetle widely distributed in the Old World, from northern Australia and Japan westward to the countries of Arabian Peninsula, including Oman and Yemen. It is the only species of the genus occurring in the Arabian Peninsula.

References

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  2. ↑ "Hydrophilidae – UNH Center for Freshwater Biology". UNH. Retrieved 2015-06-30.
  3. ↑ "Sphaeridiinae Definition. Crossword Dictionary". worddomination.com. Retrieved 2015-06-30.
  4. ↑ Hansen, Michael (1999). "Fifteen new genera of Hydrophilidae (Coleoptera), with remarks on the generic classification of the family". Insect Systematics & Evolution. 30 (2): 121–172. doi:10.1163/187631200X00228.
  5. 1 2 3 4 Short, Andrew Edward Z.; Fikáček, Martin (2013). "Molecular phylogeny, evolution and classification of the Hydrophilidae (Coleoptera)". Systematic Entomology. 38 (4): 723–752. doi:10.1111/syen.12024. ISSN   1365-3113. S2CID   83050858.
  6. ↑ Y.N. Minoshima, M. Seidel, J.R. Wood, R.A.B. Leschen, N.L. Gunter, M. Fikáček Morphology and biology of the flower-visiting water scavenger beetle genus Rygmodus (Coleoptera: Hydrophilidae) Entomological Science, 21 (2018), pp. 363-384, 10.1111/ens.12316
  7. 1 2 3 "Water Scavenger Beetles". Missouri Department of Conservation. Archived from the original on 2015-07-02. Retrieved 2015-06-30.
  8. 1 2 Ryker, Lee C. (1976). "Acoustic Behavior of Tropisternus ellipticus, T. columbianus, and T. lateralis limbalis in Western Oregon (Coleoptera: Hydrophilidae)". The Coleopterists Bulletin. 30 (2): 147–156. ISSN   0010-065X. JSTOR   3999809.
  9. ↑ Shaalan, Essam; Canyon, D. V. (2009). "Aquatic insect predators and mosquito control". Tropical Biomedicine. 26 (3): 223–61. PMID   20237438 . Retrieved 2015-06-30– via academia.edu.
  10. ↑ Short & Fikáček, 2011: 85, table 1 (see bibliography)
  11. 1 2 Pereyra, Daniela; Archangelsky, Miguel (2007-04-24). "The effect of prey density on the developmental time of larvae of an aquatic beetle: Tropisternus setiger (Insecta, Coleoptera: Hydrophilidae)". Hydrobiologia. 586 (1): 367–372. doi:10.1007/s10750-007-0710-x. hdl: 11336/101797 . ISSN   0018-8158. S2CID   37162746.
  12. ↑ Batzer, Darold P.; Resh, Vincent H. (March 1991). "Trophic Interactions among a Beetle Predator, a Chironomid Grazer, and Periphyton in a Seasonal Wetland". Oikos. 60 (2): 251. doi:10.2307/3544872. ISSN   0030-1299. JSTOR   3544872.
  13. ↑ "BugGuide: Family Hydrophilidae – Water Scavenger Beetles Data". bugguide.net. Retrieved 2024-03-20.
  14. ↑ Fikáček, Martin; Prokin, Alexander; Yan, Evgeny; Yue, Yanli; Wang, Bo; Ren, Dong; Beattie, Robert (April 2014). "Modern hydrophilid clades present and widespread in the Late Jurassic and Early Cretaceous (Coleoptera: Hydrophiloidea: Hydrophilidae): Modern hydrophilid clades in the Mesozoic". Zoological Journal of the Linnean Society. 170 (4): 710–734. doi:10.1111/zoj.12114. S2CID   82580893.
  15. ↑ Landin, Jan (July 1980). "Habitats, life histories, migration and dispersal by flight of two water-beetles Helophorus brevipalpis and H. strigifrons (Hydrophilidae)". Ecography. 3 (3): 190–201. doi: 10.1111/j.1600-0587.1980.tb00725.x . ISSN   0906-7590. PMC   7163735 . S2CID   84482581.
  16. ↑ Resetarits, William J. (September 2001). "Colonization under threat of predation: avoidance of fish by an aquatic beetle, Tropisternus lateralis (Coleoptera: Hydrophilidae)". Oecologia. 129 (1): 155–160. Bibcode:2001Oecol.129..155R. doi:10.1007/s004420100704. ISSN   0029-8549. PMID   28547063. S2CID   22620190.
  17. 1 2 3 4 Usinger, Robert L. (1956). Aquatic insects of California, with keys to North American genera and California species. Berkeley: University of California Press. doi:10.5962/bhl.title.61952.
  18. ↑ "Hydrophilidae", Encyclopedia of Entomology, Kluwer Academic Publishers, 2004, p. 1140, doi:10.1007/0-306-48380-7_2111, ISBN   0-7923-8670-1
  19. ↑ Song, Ke-Qing; Xue, Huai-Jun; Beutel, Rolf G.; Bai, Ming; Bian, Dong-Ju; Liu, Jie; Ruan, Yong-Ying; Li, Wen-Zhu; Jia, Feng-Long; Yang, Xing-Ke (2014-10-01). "Habitat-dependent diversification and parallel molecular evolution: Water scavenger beetles as a case study". Current Zoology. 60 (5): 561–570. doi: 10.1093/czoolo/60.5.561 . ISSN   2396-9814.
  20. ↑ Arribas, Paula; Velasco, Josefa; Abellán, Pedro; Sánchez-Fernández, David; Andújar, Carmelo; Calosi, Piero; Millán, Andrés; Ribera, Ignacio; Bilton, David T. (2011-12-15). "Dispersal ability rather than ecological tolerance drives differences in range size between lentic and lotic water beetles (Coleoptera: Hydrophilidae)". Journal of Biogeography. 39 (5): 984–994. doi:10.1111/j.1365-2699.2011.02641.x. ISSN   0305-0270. S2CID   3902117.
  21. ↑ Rodriguez, Georgina; Fikáček, Martin; Minoshima, Yȗsuke N.; Archangelsky, Miguel; Torres, Patricia L. M. (2021). "Going underwater: multiple origins and functional morphology of piercing-sucking feeding and tracheal system adaptations in water scavenger beetle larvae (Coleoptera: Hydrophiloidea)". Zoological Journal of the Linnean Society. 193 (1): 1–30. doi: 10.1093/zoolinnean/zlaa132 . hdl: 11336/139362 .
  22. ↑ Short, A. E. Z.; McIntosh, C. E. (2014-06-01). "Review of the Giant Water Scavenger Beetle Genus Hydrophilus Geoffroy (Coleoptera: Hydrophilidae) of the United States and Canada". The Coleopterists Bulletin. 68 (2): 187. doi:10.1649/0010-065x-68.2.187. ISSN   0010-065X. S2CID   84497637.
  23. ↑ Robinson, Ayanna; Cherry, Sabrina; Elliott, Michelle; Davis, Marsha; Adams, Grace (2016-06-15). "Leveraging university-community partnerships in rural Georgia: A community health needs assessment template for hospitals". Journal of the Georgia Public Health Association. 5 (4): 365–372. doi: 10.21633/jgpha.5.409 . ISSN   2471-9773.
  24. 1 2 Ryker, Lee C. (1994). "Male Avoidance of Female Rejection: Learning in Tropisternus Solier Water Beetles (Coleoptera: Hydrophilidae)". The Coleopterists Bulletin. 48 (3): 207–212. ISSN   0010-065X. JSTOR   4009098.
  25. ↑ Angus, Robert B.; Jia, Fenglong (2020-01-13). "Triploidy in Chinese parthenogenetic Helophorus orientalis Motschulsky, 1860, further data on parthenogenetic H. brevipalpis Bedel, 1881 and a brief discussion of parthenogenesis in Hydrophiloidea (Coleoptera)". Comparative Cytogenetics. 14 (1): 1–10. doi: 10.3897/CompCytogen.v14i1.47656 . ISSN   1993-078X. PMC   6971125 . PMID   31988701.
  26. 1 2 3 "Life History, Behavior and Morphology of the Immature Stages of Enochrus Quadripunctatus Herbst in the Laboratory (Coleoptera: Hydrophilidae) 11- Morphology". Journal of Sciences, Islamic Republic of Iran. 6 (4). 1995-12-01. ISSN   1016-1104.
  27. 1 2 3 Plague, Gordon R. (1996). "Examination of the Feeding Behavior of Larval Tropisternus (Coleoptera: Hydrophilidae)". Journal of the Kansas Entomological Society. 69 (1): 104–107. ISSN   0022-8567. JSTOR   25085655.
  28. ↑ Bameul, F (1993). "Omicrini from Sulawesi Utara, Indonesia (Coleoptera : Hydrophilidae : Sphaeridiinae)". Invertebrate Systematics. 7 (4): 751. doi:10.1071/it9930751. ISSN   1445-5226.
  29. ↑ Ryker, Lee (1972). "Acoustic behavior of four sympatric species of water scavenger beetles (Coleoptera, Hydrophilidae, Tropisternus)" (PDF). Occasional Papers of the Museum of Zoology.
  30. 1 2 Formanowicz, Daniel R.; Brodie, Edmund D. (1988). "Prey Density and Ambush Site Changes in Tropisternus lateralis Larvae (Coleoptera: Hydrophilidae)". Journal of the Kansas Entomological Society. 61 (4): 420–427. ISSN   0022-8567. JSTOR   25085034.
  31. ↑ Girón, Jennifer C.; Short, Andrew Edward Z. (2021-06-18). "The Acidocerinae (Coleoptera, Hydrophilidae): taxonomy, classification, and catalog of species". ZooKeys (1045): 1–236. doi: 10.3897/zookeys.1045.63810 . ISSN   1313-2970. PMC   8233300 . PMID   34228772.
  32. ↑ Seidel, M.; Arriaga-Varela, E.; Fikáček, M. (2016). "Establishment of Cylominae Zaitzev, 1908 as a valid name for the subfamily Rygmodinae Orchymont, 1916 with an updated list of genera (Coleoptera: Hydrophilidae)". Acta Entomologica Musei Nationalis Pragae . 56: 159–165.
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