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Parent bug
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==Parental care== [[File:Elasmucha grisea 20050608 916 part.jpg|thumb|250px|left|''Elasmucha grisea'' with eggs]] The common name of "parent bug" comes from the relatively rare insect behaviour of prolonged caring for eggs and juveniles, exhibited by females of this species. Predators, such as bugs, beetles, earwigs and ants, can eliminate all the offspring of the parent bug if there is no maternal care. The repertoire of female [[defensive behaviour]]s includes wing fanning, body jerking, tilting towards the enemy and, finally, releasing of 'nasty' odours from the scent glands,<ref name="mappes1"/><ref>Jordan KHC (1958) Die Biologie von ''Elasmucha grisea'' L. (Heteroptera: Acanthosomatidae). Beitr Entomol 8:385-397</ref><ref>Melber A, Hölsher L, Schmidt GH (1980) Further studies on the social behaviour and the ecological significance in ''Elasmucha grisea'' L. (Hem.-Het.: Acanthostomatidae) Zool Anz Jena 205:27-38</ref> After oviposition, the parent bug female stands over the egg batch and shields it throughout egg development. Predation appears to limit the clutch size in ''E. grisea''. Experiments have shown that large females lay larger egg clutches than small females. However, when the clutch size was manipulated, small females protecting large clutches lost significantly more eggs than large females guarding small clutches or females in the control groups (guarding clutches of optimal size).<ref>{{cite journal | last1 = Mappes | first1 = J | last2 = Kaitala | first2 = A | year = 1994 | title = Experiments with ''Elasmucha grisea'' L. (Heteroptera: Acanthosomatidae). Does a female parent bug lay as many eggs as she can defend? | journal = Behav Ecol | volume = 5 | issue = 3| pages = 314–317 | doi=10.1093/beheco/5.3.314}}</ref> After hatching, larvae of the parent bug remain in a tight aggregation, feeding on their empty egg shells.<ref name="hanelova"/> When any larva tries to abandon the aggregation, the female tilts her body, stretches her antennae to reach the larva and pushes the larva back to the aggregation.<ref name="roth"/> During the second and third [[instar]] they move, for food, towards catkins then back to the leaf with the female in close attendance.<ref name="hanelova"/> The female keeps a lookout for the larvae constantly and manages them with touches of her antennae. Finally, larvae form smaller groups and disperse at the end of the third instar, at which point the female leaves them. [[File: Eggs of Ectophasia crassipennis.JPG|thumb|200px|right|''E. grisea'' with the white eggs of ''[[Subclytia rotundiventris]]'' on pronotum]] It has been noted that, in ''E. grisea'', [[moulting]]s during the early instar stages can be asynchronous.<ref name="roth"/> While some larvae are still at the first instar stage, others have already moulted to the stage of second instar larvae and abandon the brood leaf for food. Under such circumstances, the female is no longer able to provide effective protection for all her larvae. The offspring of different females make contact with each other and form mixed groups. There is no kin-recognition in this species. Both single and joint guarding females provide parental care for their own larvae or other females' offspring. Larvae are likely to benefit from 'kindergartens', when their mothers disappear or die.<ref name="roth"/> Moreover, joint-guarding females defend the egg clutches much more successfully than single females.<ref>{{cite journal | last1 = Mappes | first1 = J | last2 = Kaitala | first2 = A | last3 = Alato | first3 = RV | year = 1995 | title = Joint brood guarding in parent bugs – an experiment on defence against predation | journal = Behav Ecol Sociobiol | volume = 36 | issue = 5| pages = 343–347 | doi=10.1007/bf00167795}}</ref> The [[Tachinidae|tachinid]] fly, ''[[Subclytia rotundiventris]]'', is a specialist [[endoparasite]] of the parent bug females.<ref name="mappes2">{{cite journal | last1 = Mappes | first1 = J | year = 2010 | title = Parasites and female ability to defend offspring in the parent bug ''Elsamucha grisea'' L | doi = 10.1111/j.1439-0310.1994.tb01030.x | journal = Ethology | volume = 97 | issue = 1–2| pages = 76–80 }}</ref> The parasite inserts a single egg through the upper [[prothorax]] of an ''E. grisea'' female and, after hatching, the larva feeds on its host. At the beginning the parasite feeds only on the non-vital parts of the bug, but finally it kills it and [[pupa]]tes outside the host. The larva ‘permits’ the parent bug to continue caring for the juveniles until their third stage. In experiments, the wing-fanning (regarded as the most effective defensive behaviour of the parent bug) did not differ significantly between parasited and non-parasited females until the nymphs were at the second instar stage. However, with older nymphs, females were much less effective in their defensive behaviour and often died before the end of maternal care. Considering that the older nymphs are more mobile and may escape from predators, some offspring of the parasited females of the parent bug probably survive and may also be potential hosts for the new generation of the parasite.<ref name="mappes2"/>
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