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.     

Interactions between host attributes and wasp size: a laboratory evaluation of Trichogramma platneri as an augmentative biological control agent for two avocado pests

We compared egg parasitization of two sporadic pests of avocado, Persea americana Miller, by Trichogramma platneri Nagarkatti reared on Sitotroga cerealella (Oliver) (=smaller parasitoids) with those reared on Trichoplusia ni (Hübner) (= larger parasitoids) in the laboratory. Large T. platneri females invested less time than smaller parasitoids in obtaining offspring from eggs of either Amorbia cuneana Walsingham or Sabulodes aegrotata (Gueneé). Furthermore, large T. platneri invested substantially more time (ca. 3-fold) in obtaining offspring from S. aegrotata than A. cuneana eggs. These investment times positively correlated with percentage egg-load retained by a female after it had ceased exploiting a host species. This occurred even though the offspring produced by a parasitoid from either host species manifested the same size and fecundity (= quality). Host species utilization in the laboratory mirrored that observed in the avocado groves of southern California following augmentative release of commercially produced T. platneri (Oatman & Platner, 1985). Explanations for these differences in host utilization are discussed along with their implications for augmentative biological control.     

Egg Size, Intrinsic Competition, and Lethal Interference in the Parasitoids Encarsia pergandiella and Encarsia formosa

Recent population dynamic theory predicts that disruption of biological control may occur when one parasitoid species' superiority in intrinsic competition is associated with a lower ability to find and exploit hosts (i.e., ability in extrinsic competition). One might expect such a trade-off, for instance, if parasitoids with larger (and fewer) eggs are more likely to prevail in intrinsic competition than species with smaller (and more numerous) eggs. We tested the idea that relative egg size could be used to predict the outcome of intrinsic competition in two closely related endoparasitoids, Encarsia pergandiella Howard and Encarsia formosa Gahan. Contrary to expectation, the parasitoid species with smaller eggs, E. pergandiella, prevailed in intrinsic competition, regardless of the order that hosts were exposed to the two species. In a literature survey, we found four studies of competing pairs of endoparasitoid species for which: (a) egg size estimates were available and (b) one species was consistently superior in intrinsic competition. In three of the four studies, the small-egged species prevailed in intrinsic competition, as we also found. Although E. formosa lost in intrinsic competition, this species negatively affected E. pergandiella's progeny production by host feeding on and killing hosts containing E. pergandiella eggs. E. formosa females also host fed on conspecific-parasitized hosts. As a mechanism of both intra- and interspecific interference competition, host feeding on parasitized hosts contradicts assumptions about the nature of interference competition in existing population dynamics models.     

Chemical signals mediating interactions betweenGaleruca tanaceti L. (Coleoptera, Chrysomelidae) and its egg parasitoidOomyzus galerucivorus (Hedqvits) (Hymenoptera, Eulophidae)

Chemical signals mediating interactions betweenGaleruca tanaceti and its egg parasitoidOomyzus galerucivorus (Hymenoptera, Eulophidae) were studied. Neither odor of gravid females ofG. tanaceti nor volatiles of their feces were attractive to the parasitoid. However, the presence of the beetles’ feces on a substrate arrested the parasitoid and elicited frequent antennal drumming. Thus, feces may contain a kairomone important for host finding. Odors of damaged and undamaged host plants had no effect on the parasitoids.O. galerucivorus did not detect its host eggs at close range but encountered them by chance. Neither the structure nor the dark color of the egg surface play a key role in host recognition, but chemicals of the extrachorion which could be isolated by dichloromethane. Fractionation of the dichloromethane extract by TLC revealed a single active fraction which induced host recognition behavior. Since the eggs ofG. tanaceti contain anthraquinones and anthrones which are active as feeding deterrents against predators, we hypothesized that reproductive success ofO. galerucivorus is due to sequestration of these protective compounds. However, GC-MS analyses revealed that there was no transfer of them from the host egg into the adult parasitoid.     

The introduction and recovery in the United States ofAnaphes diana [Hym.: Mymaridae], an egg parasite ofSitona weevils [Col.: Curculionidae]

Anaphes diana (Girault) (=Patasson lameerei Debauche), a mymarid egg parasite ofSitona spp., was introduced from Europe beginning in 1976 and is now tentatively established in the United States. Techniques are described for the separation of eggs ofSitona spp. from soil, using a series of fine-mesh sieves, water, and a saturated salt solution. Data from 9 years of sampling in an alfalfa field at Newark, Del. (>19,300 host eggs extracted), showed that the mean peak density of viable overwintering eggs ofSitona hispidulus (F.) was 14.6 per 100 cm³ of 1 cm deep surface soil. At the study site,Sitona egg densities consistently increased during the fall as a result of oviposition, peaked during January and February and decreased during the spring as a result of egg hatch. Although the incidence of parasitism byA. diana remained surprisingly low (0.29%), the fact that the species was recovered during 3 years and up to 7 years after the last release, indicates that it has colonized at the Delaware release site.      

Do grains whisper for help? Evidence for herbivore‐induced synomones in wheat grains

Thus far, the release of herbivore‐induced synomones (HIS) has almost exclusively been demonstrated in somatic plant tissue. Here we present evidence for the production of HIS from reproductive tissue, i.e., seeds. The study system consisted of wheat grains (Triticum aestivum L., Poaceae) infested by larvae of the granary weevil Sitophilus granarius L. (Coleoptera, Curculionidae), which in turn are attacked by the parasitic wasp Lariophagus distinguendus Förster (Hymenoptera, Pteromalidae). The use of potential chemical signals from the infested grain for host recognition of L. distinguendus was studied with weevil‐infested grains that had developed under a range of humidity conditions. Wasps always performed longer antennal drumming on infested than healthy grains, demonstrating that they were able to recognize infested grains under all humidity conditions. In grains developed at a high humidity (75% r.h.), host recognition is enabled by chemical signals arising from the host faeces which densely cover infested grains. However, in grains from a low humidity (45% r.h.) almost no faeces are present, indicating that other cues are used by the wasp. Control experiments revealed that these other cues are neither chemical signals from faeces contaminations, volatiles from faeces nor host larvae inside the grain, movement of host larvae, or moisture content of infested grains. Therefore, wasp recognition of infested grains in low humidity conditions is probably based on chemical signals from the grain itself that are induced by the feeding of granary weevil larva. Further experiments revealed that infested grains increase their viability when granary weevil larvae are killed by the idiobiont L. distinguendus. This justifies the categorisation of potential signals from the grain as HIS. To our knowledge, this is the only system where potential HIS in plant seeds have been studied. We discuss whether potential HIS are actively produced by infested grains or are a by‐product of grain metabolism.     

Responses of a generalist and a specialist parasitoid (Hymenoptera: Eucoilidae) to Drosophilid larval kairomones

Foraging parasitoids are thought to need more specific information than generalists on the presence, identity, availability, and suitability of their insect host species. In the present paper, we compare responses to host kairomones by two phylogenetically related parasitoid species that attack Drosophilidae and that differ in the width of their host range. As predicted, the behavioral response of the parasitoids to host kairomones reflected their difference in host range. The response of the specialist parasitoid Leptopilina boulardiwas restricted to contact kairomones from its natural hosts and one closely related species. In contrast, the generalist parasitoid Leptopilina heterotomaresponded to contact kairomones of a variety of Drosophilidae species.     

The Response Specificity of <Emphasis Type="Italic">Trichogramma</Emphasis> Egg Parasitoids towards Infochemicals during Host Location

Parasitoids are confronted with many different infochemicals of their hosts and food plants during host selection. Here, we investigated the effect of kairomones from the adult host Pieris brassicae and of cues present on Brussels sprout plants infested by P. brassicae eggs on the behavioral response of the egg parasitoid Trichogramma evanescens. Additionally, we tested whether the parasitoid’s acceptance of P. brassicae eggs changes with different host ages. The wasps did not discriminate between olfactory cues from mated and virgin females or between mated females and males of P. brassicae. T. evanescens randomly climbed on the butterflies, showing a phoretic behavior without any preference for a certain sex. The parasitoid was arrested on leaf parts next to 1-day-old host egg masses. This arrestment might be due to cues deposited during oviposition. The wasps parasitized host eggs up to 3 days old equally well. Our results were compared with former studies on responses by T .brassicae showing that T. evanescens makes less use of infochemicals from P. brassicae than T. brassicae.     

Resistance to Dutch Elm Disease Reduces Presence of Xylem Endophytic Fungi in Elms (Ulmus spp.)

Efforts to introduce pathogen resistance into landscape tree species by breeding may have unintended consequences for fungal diversity. To address this issue, we compared the frequency and diversity of endophytic fungi and defensive phenolic metabolites in elm (Ulmus spp.) trees with genotypes known to differ in resistance to Dutch elm disease. Our results indicate that resistant U. minor and U. pumila genotypes exhibit a lower frequency and diversity of fungal endophytes in the xylem than susceptible U. minor genotypes. However, resistant and susceptible genotypes showed a similar frequency and diversity of endophytes in the leaves and bark. The resistant and susceptible genotypes could be discriminated on the basis of the phenolic profile of the xylem, but not on basis of phenolics in the leaves or bark. As the Dutch elm disease pathogen develops within xylem tissues, the defensive chemistry of resistant elm genotypes thus appears to be one of the factors that may limit colonization by both the pathogen and endophytes. We discuss a potential trade-off between the benefits of breeding resistance into tree species, versus concomitant losses of fungal endophytes and the ecosystem services they provide.     

水椰八角铁甲和椰心叶甲对椰子和银海枣的寄主选择性

【目的】水椰八角铁甲和椰心叶甲均为棕榈科植物的重要入侵害虫,两者的外部形态、取食部位和危害特征相似。研究它们的寄主选择性有助于了解这2种害虫的扩散和成灾机制。【方法】在室内用椰子和银海枣2种寄主植物分别饲养水椰八角铁甲和椰心叶甲,研究在不同寄主植物上水椰八角铁甲和椰心叶甲的存活率、产卵率、发育历期等以及这2种害虫对不同寄主植物的选择性。【结果】水椰八角铁甲在2种寄主上的存活率差异显著,除了卵期和蛹期之外,幼虫期各虫态在银海枣上的存活率明显比在椰子上的存活率高;椰心叶甲在椰子上的存活率高于银海枣,各虫态平均存活率分别为95%和86%。取食银海枣的水椰八角铁甲达标准卵量概率为0.23,取食椰子不产卵,无法完成整个世代;取食椰子的椰心叶甲达标准卵量概率为0.86,取食银海枣不产卵,也无法完成整个世代;水椰八角铁甲取食银海枣完成世代的实验种群趋势指数为12.55,椰心叶甲取食椰子完成世代的实验种群趋势指数为66.55。【结论】水椰八角铁甲和椰心叶甲分别对银海枣和椰子这2种寄主植物具有明显的选择性。在海南椰子树的数量远远超过银海枣,该实验结果在一定程度上解释了椰心叶甲在海南岛广泛分布而水椰八角铁甲只是零星发生的原因。     

Egg-limited functional response of Uscana lariophaga, egg parasitoid of bruchid beetle pests in stored cowpea

The functional response of the egg parasitoid Uscana lariophaga Steffan (Hymenoptera: Trichogrammatidae) to eggs of its host Callosobruchus maculatus Fab. (Coleoptera: Bruchidae) was investigated in storage containers filled with a cowpea (Vigna unguiculata Walp.) seed mass. Foraging time was limited to 4 or 24 h. These indirect experiments were supplemented by direct observations of the parasitoid's handling time, egg-laying capacity and initial egg load. The foraging process of U. lariophaga can be divided into two distinct stages: the process leading to detection of host clusters and, after arrival within a host cluster, the response of the parasitoid to the host density within the cluster. The chance that a cluster is found by U. lariophaga appeared independent of the number of host eggs per cluster, but was influenced by the available foraging time. Within a host cluster, U. lariophaga demonstrated a Holling II type functional response. Parasitoids were strongly arrested in host clusters, leading to high levels of parasitism. Direct observations proved that handling time was not a limiting factor, but that U. lariophaga's initial egg load and egg maturation rate limited the plateau level of her functional response. As such, direct observations were essential for a correct interpretation of the mechanisms underlying the shape of the functional response.