ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Denisia Jahr/Year: 2002 Band/Volume: 0004 Autor(en)/Author(s): Gugliemino Adalgisa, Gugliemino Adalgisa Artikel/Article: Dryinidae (Hymenoptera Chrysidoidea): an interesting group among the natural enemies of the Auchenorrhyncha (Hemiptera) 549-556 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Dryinidae (Hymenoptera Chrysidoidea): an interesting group among the natural enemies of the Auchenorrhyncha (Hemiptera). A. GUGLIELMINO Abstract A survey on the family Dryinidae (Hymenoptera Chrysidoidea), an inte- resting group of natural enemies of the Auchenorrhyncha (Hemiptera), is empha- sized. In particular, informations on evo- lution, biology, natural enemies, economic importance, Auchenorrhyncha-Dryinidae relationships and biological control of this group on Auchenorrhyncha taxa are pro- vided. Denisia 04, zugleich Kataloge des OÖ. Landesmuseums, Neue Folge Nr. 176 (2002). 549-556 549 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Introduction tion. Females feed on sugar solutions or on their hosts consuming haemolymph and tis- The Dryinidae are a small cosmopolitan sue; in the last case, the prey may survive family of Hymenoptera Aculeata parasitizing (non-destructive host-feeding) or die Homoptera Auchenorrhyncha. It includes (destructive host-feeding) (JERVIS et al. 1996). about 1400 species belonging to 10 subfamilies Males do not feed or feed only on sugar (Anteoninae, Aphelopinae, Apodryininae, solutions. Bocchinae, Conganteoninae, Dryininae, Polyphagy appears to be the rule within Gonatopodinae, Laberitinae, Plesiodryininae, Dryinidae, as is the case in Strepsiptera and Transdryininae) and living in all terrestrial Pipunculidae which are other parasitoids of habitats from sea-level to the high mountains. Auchenorrhyncha (WALOFF & JERVIS 1987). Knowledge of the ecology and biology of Dry- Only Crovettia theliae (GAHAN), species belon- inid wasps is insufficient, the main gap regar- ging to the subfamily Aphelopinae and parasi- ding parasitoid - host interactions. Hosts are toid of Thelia bimaculata FABR1C1US (Membra- known only in 5 of the 10 above mentioned cidae), seems to be monophagous. subfamilies (Anteoninae, Aphelopinae, Dryinidae species parasitize adults and Bocchinae, Dryininae and Gonatopodinae) nymphs of Auchenorrhyncha. Usually the (GUCLIELMINO & OLMI 1997). Usually Dryi- parasitized host does not exhibit fundamental nids are both parasitoids and predators on changes, if it is parasitized in the adult stage. their hosts, being very effective for the biolo- On the contrary, the parasitization on nym- gical control of plant pests. Auchenorrhyn- phal stages can have different effects. If the cha, indeed, as well as causing direct damage host's metamorphosis is not stopped by the to plant through their feeding, oviposition, Dryinidae (as in the case of Crovettia and and production of honeydew and wax, may be Aphelopus species), the adult of the Auche- also vectors of several pathogen agents (virus, norrhyncha species exhibits fundamental mor- bacteria, spiroplasma, phytoplasma) and so phological, anatomical and physiological may cause considerable economic damage. changes (depigmentation, males similar in appearance to females, parasitic castration, Evolution and biology ecc). If the host's metamorphosis is stopped (as in Anteoninae, Bocchinae, Dryininae, In the evolution of Dryinidae, driving for- Gonatopodinae species), the nymph does not ces seem to be the host-capture and the host's reach the adult stage. According to PlLLAULT reaction (OLMI 1994)- They acted mainly on (1951), the host's metamorphosis is stopped the female sex, the only one having any by substances produced by dryinid larva. contact with the host, and involved changes of the prothorax (from very short and almost Dryinidae reproduction may be bisexual not visible in dorsal view to elongated and and/or parthenogenetic (thelytokous or arrhe- very mobile) and fore legs (from short and no notokous); the polyembryony is known only raptorial to elongated and raptorial with in Crovettia theliae (GAHAN) (Aphelopinae). robust and mobile chelae) (Figs 1-6). Regar- The females of many species of Dryinidae ding the 5 subfamilies known to have relati- are similar to ants in their general appearance. onships with Auchenorrhyncha, the Aphelo- This mimicry allows them to attack easily pinae females are the most plesiomorphic Dry- their hosts, because ants frequently feed on inids; on the contrary Dryininae and especial- the honeydew produced by Auchenorrhyncha ly Gonatopodinae females the most evolved. defending them from natural enemies. This The modest male selection explains also why phenomenon was observed mainly in apterous in several groups females and males are so dif- females of genus Gonatopus, but also in winged ferent (Figs 7, 8). females like the Australian Anteon myrmeco- Parallel with the evolution of these mor- philum (PERKINS), a myrmecophilous species phological characters, the Dryinidae females parasitoid of Ipo conferta (Cicadellidae). developed their predatory behaviour to obtain Knowledge on the biology of the Dryi- energy and nutrients necessary for egg produc- nidae is insufficient (see OLMI 1984 for a 550 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Figs 1-5. Evolution in Dryinidae females invol- ving changes mainly of thorax and fore legs. Fig. 1. Fig. 2. Crovettia masneri (OLMI 1984) Anteon ephippiger (DALMAN 1818) (Anteoninae) (from OLMI 1999). (= Biaphelopus masneri OLMI 1984) (Aphelopinae), (from OLMI Length 2,2 mm. 1984). Length 2.43 mm. Fig. 3. Fig. 4. Bocchus scaramozzinoi OLMI 1984 (Bocchinae) (from OLMI 1999). Dryinus collaris (LINNAEUS 1767) (Dryininae) (from OLMI 1999). Length 4.2 mm Length 5.9 mm. 551 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at review). In particular, few data are known on the postembryonic development. These insec- ts are hypermetamorphic; their postembryonic development consists of 4 or 5 larval instars, the last of which is the mature larva (PERKINS 1905; BUYCKX 1948; BARRETT et al. 1965; Fig. 5. HERNANDEZ & BELLOTI 1984; KITAMURA Gonatopus lunatus KLUG 1810 1985; GIRI & FREYTAG 1988; ABRIL RAMIREZ (Gonatopodinae) (from OLMI 1999). 1992; VlRLA 1992; GUCLIELMINO & VlRLA Length 3,3 mm. 1998). The immature larval instars are usually sacciform and endo-ectoparasitic. The anteri- or part of their body lies in the haemocoel of the host and the posterior part protrudes out- side between two overlapping sclerites of the host (Fig. 9). The immature larvae of the poly- embryonic genus Crovettia (Aphelopinae) are an exception being totally endoparasitic and developing inside the host haemocoel (KORN- HAUSER 1919; OLMI 1994). In each subfamily of the Dryinidae, the mature larva is hymenopteriform (sensu GAULD & BOLTON 1988); it is an eucephalous larva with developed mouth-parts. This larva kills the host by eating out all its internal organs; it then crawls out and pupates in a silk cocoon in the soil or on plants (Figs 9, 10) (see OLMI 1994 for a review). Few data are known on feeding habits of the immature larvae of the Dryinidae and con- cern only the subfamilies known having rela- tionships with Auchenorrhyncha. The following are few hypotheses quoted Fig. 6. Evolution of the chelae for each subfamily. in Dryinidae females In the Aphelopinae, the immature larval (from OLMI 1999) sho- wing an increase of the instars feed through a tissue which absorbs the number of bristles and substances from the host transforming them lamellae and a more into high energy nutrients (KEILIN &. THOMP- elongated distal part of SON 1915; FENTON 1918; BUYCKX 1948). How the 5th tarsal segment. A: Anteon jurineanum these nutrients are consumed by the Aphelo- LATREIUE 1809. Scale = pinae larvae is unknown. In addition, the 0.07 mm; B: Anteon ful- nature of this tissue is not clear. According viventre (HAUDAY 1828). Scale = 0.04 mm; KEILIN & THOMPSON (1915) and FENTON C: Dryinus canariensis (1918), this tissue is composed by hypodermal (CEBAUOS 1927). Scale = cells of the host body. These cells are stimula- 0.06 mm; ted to abnormal growth by the presence of the D: Gonatopus solidus parasite. The early stages of the parasite deve- (HAUPT 1938). Scale = 0.05 mm. lop inside of the host body and are enclosed a: arolium; b: basal part by this tissue forming a «cyst». When the of 5th segment; c: distal immature larvae assume a curved position and part of 5th segment; protrude outside of the host body, the cyst d: enlarged claw; e: bristles; f: lamellae. remains in intimate connection with the 552 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at attachement area of the larva and goes on one or more layers of transparent permeable absorbing and transforming the nutrients for cuticle. Besides, they are unrelated to the host the parasite. According to Bl'YCKX (1948), on gut. the contrary, this tissue is formed by cells of Immature larvae of Anteoninae and the trophamnion, a membrane surrounding Bocchinae seem to have a feeding habit inter- the embryo. After the eclosion, the tropham- mediate between Aphelopinae and Gonato- nion remains as a sac-like structure in front of podinae (PONOMARENKO 1975). The part of Fig. 8. the larval mouth, forming a barrier between the larva body lying into the host body is sur- Gonatopus clavipes (THUNBERG) (from the host and parasite. The cells of