Rhipiphoridae
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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 In evolutionary ecology, a parasitoid is an organism that lives in close association with its host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionary strategies within parasi ...
, an attribute that they share with the
Meloidae Blister beetles are beetles of the family Meloidae, so called for their defensive secretion of a blistering agent, cantharidin. About 7,500 species are known worldwide. Many are conspicuous and some are aposematically colored, announcing their ...
. Members of the family differ in their choice of hosts, but most attack various species of bees or
wasps A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. ...
, while some others attack
cockroaches Cockroaches (or roaches) are a paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known a ...
or beetles. Many species of Ripiphoridae have abbreviated
elytra An elytron (; ; , ) is a modified, hardened forewing of beetles (Coleoptera), though a few of the true bugs (Hemiptera) such as the family Schizopteridae are extremely similar; in true bugs, the forewings are called hemelytra (sometimes alterna ...
, and flabellate or pectinate antennae.


Biology

The subfamily Ripiphorinae parasitise bees and wasps ( Hymenoptera), while Ripidiinae parasitises
cockroaches Cockroaches (or roaches) are a paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known a ...
(
Blattodea Blattodea is an order of insects that contains cockroaches and termites. Formerly, termites were considered a separate order, Isoptera, but genetic and molecular evidence suggests they evolved from within the cockroach lineage, cladistically ...
) and Pelecotominae parasitises larvae of wood-boring beetles ( Coleoptera). 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. In species that parasitise cockroaches, males are winged while the females are wingless and
larviform Larviform female is a biological phenomenon occurring in some insect species, where the females in the adult stage of metamorphosis resemble the larvae to various degrees, while the male appears more morphologically adult (as imagoes). The resemblan ...
. Both sexes of adults cannot feed due to reduced mouthparts. Adult females attract males using
pheromones A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
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. 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.


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.


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, c = Catalogue of Life, g = GBIF, b = Bugguide.net


Extinct genera

* Subfamily Pelecotominae ** †'' Burmitoma'' Batelka et al., 2018
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, Cenomanian ** †'' Flabellotoma'' Batelka et al., 2016 Burmese amber, Myanmar, Cenomanian ** †'' Plesiotoma'' Batelka et al., 2018 Burmese amber, Myanmar, Cenomanian ** †'' Samlandotoma'' Alekseev, 2019
Baltic amber The Baltic region is home to the largest known deposit of amber, called Baltic amber or succinite. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that these forests created more than ...
, 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


External links

* * {{Taxonbar, from=Q590486 Tenebrionoidea Beetle families