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Wedge-shaped beetles; Temporal range: Albian–Recent PreꞒ Ꞓ O S D C P T J K Pg N
	Ripiphorus diadasiae male.; Note the characteristically small elytra and flabellate antennae
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Coleoptera
Suborder:	Polyphaga
Infraorder:	Cucujiformia
Superfamily:	Tenebrionoidea
Family:	Ripiphoridae; Laporte, 1840 proposed
Subfamilies
	Hemirhipidiinae; Micholaeminae; Pelecotominae; Ptilophorinae; Ripidiinae; Ripiphorinae;

Ripiphoridae (formerly spelled Rhipiphoridae) is a cosmopolitan family of some 450 described species of beetles sometimes called "wedge-shaped beetles". Ripiphoridae are unusual among beetle families in that many species are hypermetamorphic parasitoids, an attribute that they share with the Meloidae. Members of the family differ in their choice of hosts, but most attack various species of bees or wasps, while some others attack cockroaches or beetles. Many species of Ripiphoridae have abbreviated elytra, and flabellate or pectinate antennae.

Biology

The subfamily Ripiphorinae parasitise bees and wasps (Hymenoptera), while Ripidiinae parasitises cockroaches (Blattodea) and Pelecotominae parasitises larvae of wood-boring beetles (Coleoptera).^[3]

Species that attack bees typically lay their eggs on flowers. There the eggs hatch almost immediately into small planidial larvae and lie in wait for a visiting host. The planidium mounts the bee and rides it back to the hive. There it dismounts and seeks a cell occupied by a host larva. The planidium then enters the body of the host. It changes its skin and shape, then remains more or less dormant until the host larva pupates. It then emerges from the bee pupa and begins to feed. It eats the entire pupa, then pupates in its turn and completes its metamorphosis before emerging from the hive to mate and lay eggs.^[4]^[5]

In species that parasitise cockroaches, males are winged while the females are wingless and larviform. Both sexes of adults cannot feed due to reduced mouthparts. Adult females attract males using pheromones to mate, then lay eggs on the spot. The eggs hatch into larvae which attack cockroach nymphs. Upon maturity, the ripiphorid larva emerges from its host's last abdominal segments and pupates nearby.^[6]

In species that attack wood-boring beetles, adults occur on dead trees or on dead parts of living trees. They mate and then the females lay eggs into wood using a long, stiff, needle-shaped ovipositor. Larvae hatch and actively search for host beetle larvae, able to survive for at least 10 days without finding a host. When a host is found, the ripiphorid larva pierces into it with the help of a narcotising substance it injects. It feeds within the host, then emerges and continues feeding as an ectoparasitoid. When development is complete, the larva acts as a wood-borer itself, creating an emergence gallery and pupating at the apical end of this gallery.^[7]

Evolution

Fossil species in the genera Paleoripiphorus, Macrosiagon, Cretaceoripidius, Flabellotoma, Burmitoma, Plesiotoma, and Amberocula have been described from mid- to lower-Cretaceous amber from sites in France, Germany and Myanmar.^[6]^[8]^[9]^[10]

Genera

Alloclinops Broun, 1921^g
Ancholaemus Gerstaecker, 1855^g
Blattivorus Chobaut, 1891^g
Clinopalpus Batelka, 2009^g
Clinops Gerstaecker, 1855^g
Elytroxystrotus Manfrini de Brewer, 1963
Euctenia Gerstaecker, 1855
Falsorhipidius Pic, 1947
Geoscopus Gerstaecker, 1855^g
Hemirhipidius Heller, 1920
Heteromeroxylon Pic, 1939
Ivierhipidius Barclay, 2015
Macrosiagon Hentz, 1830^{i c g b}
Metoecus Dejean, 1834^g
Micholaemus Viana, 1971^g
Micropelecotoides Pic, 1910
Neonephrites Riek, 1955
Neopauroripidius Falin & Engel, 2014^g
Neorhipidius Riek, 1955
Neorrhipidius Viana, 1958^g
Nephrites Shuckard, 1838
Paranephrites Riek, 1955
Pelecotoma Fischer, 1809^{i c g b}
Pirhidius Besuchet, 1957^b
Pseudorhipidius Chobaut, 1894
Pterydrias Reitter, 1895
Ptilophorus Dejean, 1834^{i c g b}
Quasipirhidius Zaragoza, 1992
Quasirhipidius Zaragoza, 1992^g
Rhipidioides Riek, 1955
Rhipidocyrtus Falin & Engel, 2014^g
Rhipistena Sharp, 1878^g
Riekella Selander, 1957
Ripidius Thunberg, 1806^{i c g}
Ripiphorus Bosc, 1791^{i c g b}
Scotoscopus Reitter, 1884^g
Sharpides Kirkaldy, 1910
Sitarida White, 1846
Trigonodera Dejean, 1834^{i c g b}
Zapotecotoma Engel, Falin, & Batelka, 2019^{i c g b}

Data sources: i = ITIS,^[11] c = Catalogue of Life,^[12] g = GBIF,^[13] b = Bugguide.net^[3]

Extinct genera

Subfamily Pelecotominae
- †Burmitoma Batelka et al., 2018 Burmese amber, Myanmar, Cenomanian
- †Flabellotoma Batelka et al., 2016 Burmese amber, Myanmar, Cenomanian
- †Plesiotoma Batelka et al., 2018 Burmese amber, Myanmar, Cenomanian
- †Samlandotoma Alekseev, 2019 Baltic amber, Russia, Eocene
- †Spinotoma Hsiao & Huang, 2017 Burmese amber, Myanmar, Cenomanian
Subfamily Ripidiinae
- †Olemehliella Batelka, 2017 Baltic amber, Russia, Eocene
- †Pauroripidius Kaupp & Nagel, 2001 Baltic amber, Russia, Eocene
- Tribe Ripidiini
  - †Amberocula Batelka et al., 2018 Burmese amber, Myanmar, Cenomanian
  - †Cretaceoripidius Falin & Engel, 2010 Burmese amber, Myanmar, Cenomanian
  - †Paleoripiphorus Perrichot et al., 2004 Charentese amber, France, Cenomanian Burmese amber, Myanmar, Cenomanian
  - †Protoripidius Cai et al., 2018 Burmese amber, Myanmar, Cenomanian

References

^ Bousquet, Yves; Bouchard, Patrice (2018). "Case 3746 — Ripiphoridae Laporte, 1840 and Ripiphorus Bosc, 1791 (Insecta, Coleoptera): Proposed conservation of usage by designating Ripiphorus subdipterus Fabricius, 1792 as the type species of Ripiphorus and proposed ruling that Laporte (1840) used the type genus Ripiphorus in the sense defined B". The Bulletin of Zoological Nomenclature. 75: 36. doi:10.21805/bzn.v75.a010.
^ Not resolved
^ ^a ^b "Ripiphoridae Family Information". BugGuide.net. Retrieved 2018-04-22.
^ Falin, Z.H. (2002). "102. Ripiphoridae. Gemminger and Harold 1870 (1853)". In Arnett, R.H. Jr.; Thomas, M.C.; Skelley, P.E.; Frank, J.H. (eds.). American beetles. Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea. Boca Raton, Florida: CRC Press LLC. pp. 431–444. doi:10.1201/9781420041231.ch6 (inactive 2024-11-12). ISBN 978-0-8493-0954-0.{{cite book}}: CS1 maint: DOI inactive as of November 2024 (link)
^ Lawrence, J.F.; Falin, Z.H.; Ślipiński, A. (2010). "Ripiphoridae Gemminger and Harold, 1870 (Gerstaecker, 1855)". In Leschen, R.A.B.; Beutel, R.G.; Lawrence, J.F. (eds.). Coleoptera, beetles. Volume 2: Morphology and systematics (Elateroidea, Bostrichiformia, Cucujiformia partim). New York: Walter de Gruyter. pp. 538–548. doi:10.1515/9783110911213.538. ISBN 978-3110190755.
^ ^a ^b Batelka, Jan; Engel, Michael S.; Prokop, Jakub (2021). "The complete life cycle of a Cretaceous beetle parasitoid". Current Biology. 31 (3): R118 – R119. doi:10.1016/j.cub.2020.12.007. PMID 33561406. S2CID 231848712.
^ Švácha, P. (1994). "Bionomics, behaviour and immature stages of Pelecotoma fennica (Paykull) (Coleoptera: Rhipiphoridae)". Journal of Natural History. 28 (3): 585–618. doi:10.1080/00222939400770271. ISSN 0022-2933.
^ Perrichot V.; Nel A.; Neraudeau D. (2004). "Two new wedge-shaped beetles in Albo-Cenomanian ambers of France (Coleoptera: Ripiphoridae: Ripiphorinae)" (PDF). European Journal of Entomology. 101 (4): 577–581. doi:10.14411/eje.2004.081.
^ Batelka, J; François-Marie Collomb & André Nel (2006). "Macrosiagon deuvei n. sp. (Coleoptera: Ripiphoridae) from the French Eocene amber" (PDF). Ann. Soc. Entomol. Fr. 42 (1): 75–78. doi:10.1080/00379271.2006.10697451.
^ Batelka, J; MS Engel & J Prokop (2018). "A remarkable diversity of parasitoid beetles (Ripiphoridae) in Cretaceous amber, with a summary of the Mesozoic record of Tenebrionoidea". Cretaceous Research. 90 (1): 296–310. doi:10.1016/j.cretres.2018.04.019. S2CID 134951153.
^ "Ripiphoridae Report". Integrated Taxonomic Information System. Retrieved 2018-04-22.
^ "Browse Ripiphoridae". Catalogue of Life. Retrieved 2018-04-22.
^ "Ripiphoridae". GBIF. Retrieved 2018-04-22.