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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
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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
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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.
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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 the rounded by a tissue which is not as wide as in OLMI 1999): male. Length =2.5 mm.
Fig. 7. Gonatopus clavipes (THUNBERG) (from OLMI 1999): female. Length = 3.3 mm. trophamnion select the substances from the Fig. 9. host haemolymph and transform them in Immature larva of Dryinidae on nutrients for the parasite. the host. Regarding Dryininae and Gonatopodinae interesting data on the feeding habits were provided by PONOMARENKO (1975) and espe- cially by CARCUPINO et al. (1998). Immature larvae of Dryininae and Gonatopodinae have a pair of vesicles in front of their head. They are generally considered to be involved in lar- val feeding. According to CARCUPINO et al. (1998), the vesicles are evaginations of the body wall , unrelated to the digestive apparatus of the larva. They are formed by large epithelial cells showing signs of intense protein synthesis and completely enveloped in
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the Aphelopinae. It is composed by a layer of Natural enemies of hypodermal cells of the host lined by a mem- Dryinidae brane on the larva side. The larvae have a pair The main natural enemies both para- of small conical processes which emerge from sitoids and predators of Dryinidae (OLMl the oral cavity and are in contact with the 1999) are listed (Tab. 1, 2). The activity of above tissue. According to PONOMARENKO these may greatly reduce the efficiency of this (1975), these processes are probably homo- group. Fig. 10. logous to the oral vesicles of Dryininae and Mature larva of Dryinidae just after emergence from the host Gonatopodinae. Auchenorrhyncha-Dryinidae relationships
Up to now about 800 relationships to dif- ferent systematic levels between the two groups are known (GUGLIELMINO &. OLMI Dryinidae species has relationships with more Auchenorrhyncha species belonging to the same or several systematic levels; only few cases of monospecific taxa are known (for instance the Aphelopinae Crovettia theliae which seems to parasitize only the Membraci- dae Thelia maculata ¥.), but this probably could he due to scarcity of research. Regarding the number of species involved, these relation- ^ips include just 320 Auchenorrhyncha and 230 Dryinidae species. This number is insigni- ficant when compared with the number of known Auchenorrhyncha (over 40.000 spe- cies) and Dryinidae (about 1.400 species). From a biogeographical point of view, the most studied regions are the Palaearctic and Nearctic. Moreover, it is important to point
Tab. 2. Predators known to attack Dryinidae
DRYINIDAE PREDATORS stage attacked
Adults Ants
Larvae Crabonidae Sphecidae
Cocoons Insects Birds Rodents
Fig. 11. Tab. 1. Cocoon of Dryinidae Predators known to attack Dryinidae
Aphelopinae Anteoninae Bocchinae Dryininae Gonatopodinae
Aphelinidae + Ceraphronidae + Chalcididae + Diapriidae + Encyrtidae + + Ichneumonidae + Pteromalidae +
554 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at out that at least one case of Auchenorrhyncha give the pests less chances of damaging plants parasitization by Anteoninae, Aphelopinae, directly or transmitting pathogen agents. Dryininae and Gonatopodinae is known in Unfortunately, little is known on the role each zoogeographical region; on the contrary, of dryinids and other parasitoids in limiting the records of parasitization by Bocchinae are Auchenorrhyncha populations. Indeed many known only in the Palaearctic and Nearctic factors, like the parasitoid interference, the regions. In spite of insufficient knowledge of heterogeneous parasitism rates, the presence the Auchenorrhyncha-Dryinidae relations- of refugia, the presence of invulnerable deve- hips, some apparent constant links at several lopmental stages, can influence Auchenorr- systematic levels emerge from these data. hyncha population control (GUGLIELMINO While Auchenorrhyncha-Dryinidae relations- 2000). According to WALOFF & JERVIS (1987), hips are certainly the result of a co-evolution it is very difficult to evaluate the factors which between the two groups, the scarce knowledge can determine the outcome of a biological of the interactions (from both qualitative and control attempt in terms of parasitoid-hopper quantitative points of view), the biology and population dynamics. the phylogeny of both taxa does not presently Presently, Dryinidae are used in some bio- allow us to theorize on how the relationships logical control programmes. In Italy, France evolved. For the same reason, up to now we and Switzerland, Neodryinus thyphlocybae are not able to understand the sensory and (ASHMEAD) is used in the control of Metcalfa biological bases of Auchenorrhyncha-Dryi- pruinosa (SAY) (in Italy, a second dryinid, nidae links. Any hypothesis on a phylogenetic Thaumatodryinns danieli OLMI, is being reared congruence should be premature. for the same purpose); in New Zealand, Neo- dryinus nelsoni PERKINS to control Scolypopa australis (WALKER); in Nigeria, at the I.I.T.A. Biological control (Ibadan) control projects using Dryinidae against Cicadulina spp., dangerous to maize, Dryinidae have a high potential as biolo- are being studied. gical control agents. The predatory and para- sitic efficiency of this family has been studied Acknowledgements in many species. In fact, females of many taxa, mainly of Gonatopodinae, kill significant I would like to thank Prof. M. Olmi of numbers of hosts by feeding as well as by para- Viterbo, E.J. Brill (Leiden, The Netherlands) sitism. At least within 4 subfamilies (Gonato- and Calderini Edizioni (Bologna) for their podinae, Dryininae, Bocchinae and Anteo- kind permission to reproduce figs 1-8, first ninae) predation is almost as important as published in OLMI 1999. I wish to thank again parasitization (OLMI 1994). Gonatopodinae Prof. M. Olmi for his valuable suggestions and are very active in predation (WALOFF 1974; Mr. M. Vollaro (Viterbo) for his nice photo- CHUA & DICK 1982; KITAMURA 1982; GIRI & graphs. FREYTAG 1988; ALMA & ARZONE 1994). This study was supported by a grant of the Anteoninae show a lower activity of predation University of Tuscia: 60%. than Gonatopodinae (WALOFF 1974). Dryi- ninae and Bocchinae, both very scarcely inve- Zusammenfassung stigated, surely have predation activity, while Aphelopinae have none (OLMI 1994). As pre- Die Familie Dryinidae (Zikadenwespen; viously mentioned, the host-feeding can be Hymenoptera Chrysidoidea), eine interessan- «non-destructive» or «destructive», if the te Gruppe natürlicher Feinde der Zikaden, wounds inflicted by the dryinid female permit wird überblicksartig vorgestellt. Informatio- the host to survive or not. From an applied nen zu Evolution, Lebensweise, natürlichen point of view, the destructive host feeders are Feinden und wirtschaftlicher Bedeutung wer- valuable because they kill their prey more den dargestellt und Beziehungen zwischen rapidly than not feeder parasitoids, i.e. they Zikaden und Dryiniden erläutert.
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