Host location in Oomyzus gallerucae (Hymenoptera: Eulophidae), an egg parasitoid of the elm leaf beetle Xanthogaleruca luteola (Coleoptera: Chrysomelidae)

Eggs of the elm leaf beetle Xanthogaleruca luteola are often heavily attacked by the chalcidoid wasp Oomyzus gallerucae. We studied the chemical signals mediating interactions between the egg parasitoid, its host, and the plant Ulmus campestris. Olfactometer bioassays with O. gallerucae showed that volatiles of the host-plant complex attract the parasitoid. In order to determine the source of attractive volatiles within this host-plant-complex, we tested separately the effect of odours of eggs, gravid elm leaf beetle females, faeces of the beetles and elm twigs (with undamaged leaves and leaves damaged either mechanically or by feeding of the beetles). Odours of faeces of the elm leaf beetle were attractive, whereas neither volatiles from eggs nor from gravid females acted as attractants. Volatiles from undamaged or damaged plants did not elicit a positive reaction in O. gallerucae, whereas volatiles from feeding-damaged plants onto which host eggs had been deposited were attractive. This latter result suggests that it is not feeding but deposition of host eggs onto elm leaves that induces the production of plant volatiles attractive to the egg parasitoid. Investigations of the search patterns of O. gallerucae within the habitat by laboratory bioassays revealed that the egg parasitoid encounters host eggs by chance. Contact kairomones from faeces were demonstrated to be important in microhabitat acceptance, while contact kairomones isolated from the host eggs are relevant for host recognition. Received: 12 February 1997 / Accepted: 29 April 1997     

RearingTetrastichus gallerucae [hymenoptera: Eulophidae] for biological control of the ELM leaf beetleXanthogaleruca luteola

Tetrastichus gallerucae (Fonscolombe) was collected in southern France in 1985 and is being reared at the Division of Biological Control in Albany, Ca for use against the elm leaf beetle (ELB),Xanthogaleruca luteola (Müller) in northern California. This egg parasitoid can be reared easily by keeping the beetle eggs on a moist substrate to prevent desiccation and by inhibiting the growth of mold with air currents. A laboratory colony of ELB adults provides a constant supply of fresh eggs for the parasitoids. In 1985 and 1986 over 80,000T. gallerucae were released at 17 sites.      

Patch-exploitation patterns in an egg parasite of elm leaf beetle

Eggs of elm beetle,Pyrrhalta luteola (Muller), normally occur in masses which can be viewed as discrete host patches in space and time. Analysis of>800 egg masses from 3 field sites in northern California revealed that the number of eggs/mass varied from 2 to>40 and that the imported egg parasiteTetrastichus gallerucae (Fonsc.) exploited a relatively large proported of the egg masses at certain times. The spatial relationship between parasitization and number of eggs/egg mass was assessed in 2 ways-i.e., for exploited masses only and for all masses combined (exploited+nonexploited). Percent parasitization was density independent in most cases for both data sets. It is suggested that both methods of analysis can provide patterns which are relevant to biological control of insect pests.      

Response of the elm leaf beetle to host plants induced by oviposition and feeding: the infestation rate matters

We investigated by olfactometry and feeding‐ and oviposition‐choice‐tests how the highly specialised elm leaf beetle, Xanthogaleruca luteola Müller (Coleoptera: Chrysomelidae), responds to conspecifically induced defences in the field elm Ulmus minor Miller (Ulmaceae). While egg deposition of the beetle induced elms to release volatiles attractive to the egg parasitoid Oomyzus gallerucae Fonscolombe (Hymenoptera: Eulophidae), feeding alone did not. In the present study, females of the elm leaf beetle showed preferences for the odours of twigs induced by low egg deposition and feeding over odours from uninfested twigs. In contrast, heavy infestation rendered elm odours less attractive to the beetles. Feeding and oviposition bioassays revealed an oviposition preference for leaves from uninfested twigs when compared to locally infested leaves. However, beetles preferred to feed upon systemically induced leaves compared to uninfested ones. The different preferences of the elm leaf beetle during host plant approach might be explained by a strategy that accounts for both gaining access to high quality nutrition and avoiding competition or parasitism.     

Functional response of Tetrastichus gallerucae (Hym: Eulophidae) to different densities of the elm leaf beetle eggs,Xanthogaleruca luteola (Col: Chrysomelidae)

The elm leaf beetle is often considered as much of an indoor nuisance as a garden pest since large numbers of adults migrate into homes seeking a protected place to overwinter. Outdoors, both adults and larvae feed on the emerging leaves of virtually all species of elm trees, leaving skeleton zed foliage in their wake. Repeated severe infestations can markedly weaken a host tree, making it susceptible to other insects and diseases. The elm leaf beetle does not transmit the well-known Dutch elm disease, which is carried by another insect than elm bark beetle. The functional response of Tetrastichus gallerucae bees, that is the most important natural enemy of egg elm beetle Xanthogallerucae luteola, has been detected in Kerman province. The experiment was carried out in the growth room under conditions of 25 ± 1°C, 60 ± 5 RH and 16L:8D. Every female bee was exposed for 24 hours at densities of 2, 4, 6, 8, 10, 12, 16, 20, 30 and 40 eggs of elm leaf beetle. Every density had 10 repetitions. The results showed that T. gallerucae has the second type of functional response. The searching efficiency and the handling time of T. gallerucae was 2.63 and 0.114 on the eggs of elm leaf beetle and 2.193 and 0.112 on the eggs of hostess with diluted honey.     

Study on some biological characteristics of Tetrastichus gallerucae (Hym. Eulophidae), important parasitoid of Elm leaf beetle eggs in Kerman,Iran

In this study, a number of biologic characteristics of Tetrastichus gallerucae, the most important natural enemy of Xanthogaleruca luteola (Col.: Chrysomellidae), were studied. Experiments were carried out in growth room with a temperature of 25°C±1°C, relative humidity of 60%±5% and light period of 16L:8D. The results demonstrated the following outcomes: without food and host egg, the longevity of male and female bees was respectively, 1.23 and 1.35 days. On the host egg, their longevities were 3.85 and 6.02 days, respectively. On rarefied honey their longevities were 14.04 and 24.74 days, respectively and on the host eggs together with rarefied honey, their longevities were 13.4 and 34.72 days, respectively. The average growth period times in the egg of elm leaf beetle of male and female bees were 12.97 and 12.63 days, respectively. The percentage of female bees decreases as the longevity of the insects increases. These bees lay eggs in the 1–9-day-old eggs of elm leaf beetle but 1-day-old eggs of the host are preferred by T. gallerucae.     

Smelling the tree and the forest: elm background odours affect egg parasitoid orientation to herbivore induced terpenoids

Parasitic insects use herbivore induced plant volatiles as signals for host location. However, their responses to these volatiles in the background of natural habitat odours need further evaluation for developing successful biological control strategies. Field elms (Ulmus minor Miller (Ulmaceae)) release a blend of volatiles in response to oviposition of the elm leaf beetle, Xanthogaleruca luteola Müller (Coleoptera: Chrysomelidae), a major urban and forest pest in the USA and Australia. This induced blend attracts the beneficial egg parasitoid Oomyzus gallerucae Fonscolombe (Hymenoptera: Eulophidae). Our olfactory assays showed that an odorous background of non-attractive host plant volatiles from feeding damaged elms or (Z)-3-hexenyl acetate masks the attractive effect of the host-induced (E)-β-caryophyllene to O. gallerucae. Quantitative GC–MS analyses revealed decreased concentrations of (Z)-3-hexenyl acetate accompanied by highly increased concentrations of sesquiterpenes in oviposition and feeding treated elms compared to undamaged elms. This finding hints to how the parasitoid might distinguish between different odorous backgrounds. It is corroborated by the outcome of our field study in natural elm stands, where the egg parasitoid parasitized more host egg masses due to an artificially induced blend of elm terpenoids.     

Host handling behavior of the egg parasitoid Telenomus triptus to the egg mass of the stink bug Piezodorus hybneri

Host handling behavior of Telenomus triptus Nixon (Hymenoptera: Scelionidae) to an egg mass of Piezodorus hybneri Gmelin (Heteroptera: Pentatomidae) was studied in the laboratory. Five distinct behavioral events could be distinguished in the host handling behavior: drumming, ovipositor-insertion, marking, walking, and resting. Female wasps showed two types of behavioral sequence in an ovipositional bout. One consisted of drumming, ovipositor-insertion, and marking, and the other drumming, ovipositor-insertion, walking, and resting. Females did not seem to lay an egg when ovipositor-insertion was not followed by marking. This was observed frequently in the early oviposition bout, on average 2.9 times per female. Females finally succeeded in parasitizing all the eggs in a host egg mass in most cases. The durations of drumming, ovipositor-insertion, and marking on an egg mass were nearly constant, while the total time spent by a female on and beside a host egg mass varied considerably as a result of variable durations of walking and resting. Self-superparasitism occurred when almost all the eggs in a host egg mass were parasitized. Females laid the first male egg within the first four eggs; this could be an adaptation to small egg masses or single egg.     

Flavonoid chemistry and taxonomy in Ulmus

There is evidence in Ulmus of impoverishment of flavonoid constituents with evolutionary advancement and dispersal, but this is less marked in Ulmus than in Geranium and Dillenia. lnfraspecific variability is present in U. minor and U. macrocarpa. The phylogeny of Ulmus, systematic relationships within the Ulmaceae and the systematic position of the Urticales are discussed.     

Host recognition cues of the granary weevil parasitoid Lariophagus distinguendus

Host recognition was examined in Lariophagus distinguendus (Förster), a parasitoid of larvae of the granary weevil Sitophilus granarius (L.) that live endophytically in wheat grains. On encountering a grain infested with S. granarius, females of L. distinguendus behave in a set sequence. First they show antennal drumming on the grain, second they tap with the tip of the abdomen on the grain surface, third they drill into the grain and then insert their ovipositor. Bioassays revealed that drumming and drilling was stimulated by non-volatile chemicals present on the grain-host complex. Host faeces and herbivore damaged grain material stimulated the most activity, followed by artificially damaged grain, and healthy grain. This is the first report on non-volatile chemicals released from herbivore-damaged seeds as signals for foraging parasitoids. Volatile chemicals from the faeces alone were not active. Experiments on the use of physical cues revealed that the presence of a three-dimensional structure increased the response towards chemicals from the faeces. The shape (ovoid or rectangular solid) and colour (brown or white) of the structure had no impact. Thus, physical cues alone were insufficient to stimulate host recognition behaviour, but acted by increasing the response towards the chemical stimuli.