Seasonal phenology and stage-specific parasitism of the apple ermine moth, Yponomeuta malinellus Zeller, in Korea

The apple ermine moth, Yponomeuta malinellus Zeller (Lepidoptera: Yponomeutidae), is a tent caterpillar that feeds on Malus spp. in Korea. Populations of the moth in native areas appeared to be regulated by the assemblage of parasitoids. Phenological associations between host stages and parasitoids, susceptible stage(s) of the host for each parasitoid, and stage‐specific parasitism were studied. The egg larval parasitoid Ageniaspis fuscicollis (Dalman) had highest parasitism of first instar larvae (24%), with 14% parasitism of other larval stages. Dolichogenidea delecta (Haliday) was recovered from all larval instars with the highest parasitism rate of second instar larvae (20.1%), followed by 19.9% parasitism of mid‐larval hosts. Herpestomus brunicornis Gravenhorst was reared from second instar larvae through to pupal collection, and had the highest parasitism rate (29.9%) at the pupal stage. The larval pupal parasitoid Zenillia dolosa (Meigen) was recovered from mid‐larval to pupal stages with the highest parasitism rate (5.5%) occurring in third to fourth instar larvae. The host stages for developing A. fuscicollis completely overlap with those of D. delecta, and with those of H. brunicornis to some degree. A statistically significant negative correlation exists between A. fuscicollis and these dominant parasitoids, indicating competitive interaction within the host.     

Larval parasitoids of the apple ermine moth, <Emphasis Type="Italic">Yponomeuta malinellus</Emphasis>in Korea,Japan, and China

Larval parasitoids of Yponomeuta malinellus Zell. (Lepidoptera: Yponomeutidae), the apple ermine moth (AEM), were sought in northeast Asia with the goal of identifying potential biological controls of the moth, which appeared to threaten the apple industry in Washington State, USA during the 1980s. Ten primary and four secondary parasitoids were found. Dolichogenidea delecta (Haliday) (Braconidae), Ageniaspis fuscicollis (Dalman) (Encyrtidae), Herpestomus brunnicornis Grav. (Icheumonidae), Bessa paralella (Meigen), and Zenillia dolosa (Meigen) (Tachinidae) were the most important parasitoids. The composition of parasitoid species was more diverse in Korea and Japan than in China; two species were found in China, compared to nine in Korea and seven in Japan. A. fuscicollis caused a greater mortality in all investigated countries; 22.7, 11.0, and 9.3% in China, Korea, and Japan, respectively. There was a high similarity in the composition of the parasitoid complex between Korean and Honshu populations but the rates of parasitization were different. The polyphagous B paralella caused significantly higher parasitism in Honshu (18.2%) than in Korea (0.5%). In contrast, H. brunnicornis caused significantly higher parasitism in Korea (8.7%) than in Japan (2.4%). A. fuscicollis and H. brunnicornis, judged to be the most appropriate parasitoids for biological control of AEM moth in the USA, were collected and sent to the USA for release. A. fuscicollis established and is contributing to the control of AEM.     

Parasitoid complex of citrus leafminer, Phyllocnistis citrellaon lemon in the Central Jordan Valley

From March 1997 to June 1999, samples of lemonleaves infested with citrus leafminer (CLM),Phyllocnistis citrella Stainton(Lepidoptera: Gracillariidae), were collectedfrom the Al-Masalha Citrus Orchard in theCentral Jordan Valley in order to rear CLMparasitoids and to study their populationtrends. Nine species of eulophid parasitoidswere reared from CLM larvae; these were Cirrospilus ingenuus Gahan, C. pictusNees, Pnigalio agraules, Pnigalio sp. B,Pnigalio sp. C, Citrostichusphyllocnistoides Narayanan, Ratzeburgiolaincompleata Boucek, Semielacherpetiolatus Girault, and Zagrammosoma sp.Cirrospilus ingenuus was the most commonparasitoid of CLM from March 1997 to June 1999and could be a good potential biologicalcontrol agent after rearing and releasing it incitrus orchards in the Jordan Valley. Onehyperparasitoid species was reared fromimmatures of parasitoids of CLM larvae, andidentified as Tyndrichus sp. An increasein the activity of parasitoids was observed inautumn 1999 when the temperature was between 15–20 °C and the relative humidity was between 55–65%.     

Role of host-plant stimuli in sexual behaviour of small ermine moths (Yponomeuta)

ABSTRACT. 1. Studies involving experiments with olfactometers and field experiments have shown that the release of sex-pheromone by females of Yponomeuta spp. is stimulated by the presence of host plants.
2. While in Y.evonymellus L. less females emitted pheromone in the absence of host plants, in Y.padellus L. the initiation of pheromone release was delayed.
3. Females of Y.evonymellus, Y.cagnagellus Hüb., Y.padellus and Y.plumbellus D. & S. all preferred a place with host-plant odour to one with non-host odours for sex-pheromone release.
4. Male preference for host-plant odour was demonstrated in the laboratory only for Y.cagnagellus.
5. Trap catches in the field indicated a reinforcement of female attractiveness by host-plant stimuli.     

Microsatellite markers for testing host plant‐related population differentiation in the moth genus Yponomeuta

Nine microsatellite primers were developed for Yponomeuta padellus (Lepidoptera: Yponomeutidae) and tested for their applicability in analysing genetic population structure. Eight of the nine loci were highly polymorphic with on average 11.4 alleles. Cross‐species amplification of the nine primer pairs was tested in five other moth species. Primer pairs amplified in Y. cagnagellus, Y. malinellus, Y. evonymellus, and Y. rorellus but not in Y. sedellus and Plutella xylostella.     

Population structure in the small ermine moth Yponomeuta padelus: an estimate of male dispersal

Abstract.

• 1 An extensive investigation was carried out into the structure of a large, effectively continuous population of Yponomeuta padellus Latreille, feeding on hawthorn, Crataegus monogyna, and blackthorn, Prunus spinosa.
• 2 Mark—recapture studies were used to obtain direct estimates of the male dispersal rate, σ, in two successive years.
• 3 Dispersal was found to be extremely localized with respect to the total area of the study population. Estimates of u ranged from a minimum of 2.47 m to a maximum of 12.5 m.
• 4 Limited dispersal could be an important factor governing the genetic differentiation of Y.padellus populations, and its implications for the supposed existence of host races in this species are discussed.

     

Parasitoid complex of the pistachio twig borer moth,Kermania pistaciella,in Iran

The parasitoid complex of the pistachio twig borer moth, Kermania pistaciella Amsel (Lepidoptera: Tineidae), a native pest of pistachio trees, was investigated at 27 pistachio plantation sites in Kerman province, the major pistachio growing area of Iran. The present study was conducted to document the naturally established parasitoid complex and to assess the need for improving the biological control of this species. In total, 22,390 moth cocoons were collected from 186 samples collected from commercial orchards during 2006–2008 and kept singly in controlled conditions to rear immature insects. An average of 2.8% of moth cocoons had been attacked by predators at time of sampling. Of the collected cocoons, on average 46.7% completed development and emerged as adult moths, no insects emerged from 8%, suggesting that the moth or wasp died before maturing, and parasitoids emerged from the remaining 42.5%. The overall percentage of host cocoons from which wasps emerged ranged from 25.6 to 59%. Fifteen hymenopterous parasitoid species were recovered from cocoons, of which three species were primary parasitoids, two were obligatory hyperparasitoids and the remaining 10 species were facultative hyperparasitoids. The primary parasitoid, Chelonus kermakiae (Tobias) (Hymenoptera: Braconidae), was the most abundant comprising almost 85% of the total emerging parasitoids. In addition, a further four species of larval parasitoid developed within the PTBM's larval tunnels in pistachio fruit cluster-stem tissue. Conservation of these parasitoids in the pistachio growing areas is recommended since a high level of parasitized moths’ cocoons was found at the majority of experimental sites.     

Behavioural and electrophysiological activity of unsaturated analogues of the pheromone tetradecyl acetate in the small ermine moth Yponomeuta rorellus

ABSTRACT In field-trapping experiments the unsaturated analogues ( E )-6-, ( E )-12-, and ( Z )-12-tetradecenyl acetate were as attractive to male Yponomeuta rorellus Latr. as the native pheromone component tetradecyl acetate. All four analogues attracted more males than virgin females did, whereas ( Z )-6-, ( E )-11-, ( Z )-10- and ( Z )-11-tetradecenyl acetate were essentially non-attractive. Addition of 1–30% of ( Z )-11-tetradecenyl acetate to the pheromone tetradecyl acetate reduced the attraction to less than 2%. Flight tunnel experiments with Y. rorellus confirmed the activity of the ( E )-6- and ( E )-12-tetradecenyl acetates and demonstrated the activity of ( E )-7-tetradecenyl acetate as well. These analogues elicited orientation behaviour, upwind flight and landing at the odour source as frequently as the native pheromone did. Single sensillum recordings from male Y. rorellus showed two types of cells in most sensilla. A large spike amplitude cell was stimulated by tetradecyl acetate and the unsaturated analogues ( E )-11-, ( E )-6- and ( E )-12-tetradecenyl acetate, and to a lower extent by the ( Z )-6-, ( Z )-11- and ( Z )-12-isomers. A cell with medium spike amplitude was stimulated by ( Z )-9-tetradecenyl and ( Z )-11-hexadecenyl acetate. Some sensilla contained a third cell firing with a small spike amplitude which was activated by ( Z )-11-tetradecenol. Thus the tetradecyl acetate receptor was stimulated not only by the behaviourally active analogues, but also by behavioural antagonists. The interaction of ( E )-11-tetra-decenyl acetate and tetradecyl acetate with the same antennal receptor cell was also demonstrated in Y.cagnagellus . Electrophysiological discrimination between behavioural attractants and antagonists and the role of behavioural antagonists in the interspecific relations between Y.rorellus and sympatric closely related species are discussed.     

Olfactory receptors on the maxillary palps of small ermine moth larvae: evolutionary history of benzaldehyde sensitivity

In lepidopterous larvae the maxillary palps contain a large portion of the sensory equipment of the insect. Yet, knowledge about the sensitivity of these cells is limited. In this paper a morphological, behavioral, and electrophysiological investigation of the maxillary palps of Yponomeuta cagnagellus (Lepidoptera: Yponomeutidae) is presented. In addition to thermoreceptors, CO₂ receptors, and gustatory receptors, evidence is reported for the existence of two groups of receptor cells sensitive to plant volatiles. Cells that are mainly sensitive to (E)-2-hexenal and hexanal or to (Z)-3-hexen-1-ol and 1-hexanol were found. Interestingly, a high sensitivity for benzaldehyde was also found. This compound is not known to be present in Euonymus europaeus, the host plant of the monophagous Yponomeuta cagnagellus, but it is a prominent compound in Rosaceae, the presumed hosts of the ancestors of Y. cagnagellus. To elucidate the evolutionary history of this sensitivity, and its possible role in host shifts, feeding responses of three Yponomeuta species to benzaldehyde were investigated. The results confirm the hypothesis that the sensitivity to benzaldehyde evolved during the ancestral shift from Celastraceae to Rosaceae and can be considered an evolutionary relict, retained in the recently backshifted Celastraceae-specialist Y. cagnagellus.     

Oviposition in an eruptive moth species, Yponomeuta evonymellus, is insensitive to the population density experienced during the larval period

We studied if the population density experienced during larval development affects the reproductive schedule of Yponomeuta evonymellus L. (Lepidoptera, Yponomeutidae), a moth species characterized by outbreak population dynamics. More specifically, we predicted that reproduction would be delayed to facilitate emigration from sites with suboptimally high densities of conspecifics. We manipulated larval densities in the laboratory, as well as in those pupae collected from the extremes of natural densities. As the response, we also recorded the timing and sizes of egg clutches laid. The results did not support our initial predictions: the timing of oviposition was not dependent on larval growth conditions. This apparent lack of adaptation might be related to the loss of ‘memory’ during metamorphosis in holometabolous insects. However, oviposition schedules were also only minimally sensitive to elevated adult density. An inability to respond to high larval densities may contribute to the outbreak dynamics in this species.     

Parasitoid diversity reduces the variability in pest control services across time on farms

Recent declines in biodiversity have increased interest in the link between biodiversity and the provision and sustainability of ecosystem services across space and time. We mapped the complex network of interactions between herbivores and parasitoids to examine the relationship between parasitoid species richness, functional group diversity and the provision of natural pest control services. Quantitative food webs were constructed for 10 organic and 10 conventional farms. Parasitoid species richness varied from 26 to 58 species and we found a significant positive relationship between parasitoid species richness and temporal stability in parasitism rates. Higher species richness was associated with lower variation in parasitism rate. A functional group analysis showed significantly greater parasitoid species complementarity on organic farms, with on average more species in each functional group. We simulated parasitoid removal to predict whether organic farms experienced greater robustness of parasitism in the face of local extinctions. This analysis showed no consistent differences between the organic and conventional farm pairs in terms of loss of pest control service. Finally, it was found that the different habitats that make up each farm do not contribute equally to parasitoid species diversity, and that hedgerows produced more parasitoid species, significantly more so on organic farms.     

Temporal variation in the genetic structure of host-associated populations of the small ermine moth Yponomeuta padellus (Lepidoptera, Yponomeutidae)

Temporal changes in allele frequencies were studied in host-associated populations of the small ermine moth Yponomeuta padellus. At one site, populations from three host plants (Sorbus aucuparia, Amelanchier larnarckii , and Crataegus spp.) were sampled annually during a four-year-period and analysed with 20 polymorphic allozyme markers. At two other sites, allele frequencies at 5- 6 enzyme loci of Y. padellus populations from two different host plants were also tested for consistency over a 13-year-pcriod. Significant allele frequency changes occurred in the short-term analysis, whereas allele frequencies remained relatively stable through time in the long-term analyses. Furthermore, allele frequencies of Y. padellus populations from Crataegus spp. were relatively stable compared to the other host populations. The role of the agents responsible for the observed patterns is discussed.     

Introduction and establishment of parasitoids for the biological control of the apple ermine moth, Yponomeuta malinellus (Lepidoptera: Yponomeutidae), in the Pacific Northwest

Four parasitoids were imported from five countries in Eurasia and released in northwestern Washington, US, to control the apple ermine moth (AEM), Yponomeuta malinellus Zeller, which colonized the Northwest around 1981. From 1988 to 1991, 95,474 individuals of Ageniaspis fuscicollis (Dalman) from France, China, Korea, and Russia were released in Washington. Parasitism of AEM increased 4- to 5-fold over that produced by preexisting natural enemies between 1989 and 1994 at 22 monitored sites. Subsequently, the wasp dispersed up to 20 km from release sites. A. fuscicollis also parasitized the cherry ermine moth, Yponomeuta padellus (L.), which was discovered in the Pacific Northwest in 1993. A total of 1813 individuals of Herpestomus brunnicornis (Gravenhorst) from France, Korea, and Japan were released in 1989–1991, and 26 wasps were recovered in 1994–1995. From 1989 to 1991, 2647 Diadegma armillata (Gravenhorst) individuals from France were released. D. armillata was recovered at one site in 1991 two months following release, but no other recoveries have been made. A total of 8274 Eurystheae scutellaris(Robineau-Desvoidy) individuals were released in 1989 to 1991. However, this tachinid has not been recovered. A consistent decline of AEM populations occurred in 1989–1995, including at sites where A. fuscicollis was not recovered, suggesting other factors also contributed to this pest’s decline. Now well established in western Washington, A. fuscicollis may help suppress future outbreaks of Y. malinellus and its congener, Y. padellus.     

Parasitoid diversity and impact on populations of the diamondback moth Plutella xylostella (L.) on Brassica crops in central México

Three experimental plots were established and maintained during one year at two sites in the state of Querétaro, México, in order to identify species of parasitoids attacking the diamondback moth (DBM) Plutella xylostella (L.) (Lepidoptera: Plutellidae), and to gather information concerning their relative importance and patterns of population fluctuation. At both sites, the plots were planted with broccoli, cabbage and cauliflower during three cropping seasons. Parasitoid species identified were: Diadegma insulare Cresson, Diadromus (= Thyraeella) collaris Gravenhorst (Hymenoptera: Ichneumonidae); Habrobracon sp. (Hymenoptera: Braconidae); Oomyzus (= Tetrastichus) sokolowoskii Kurdjumov (Hymenoptera: Eulophidae); and Spilochalcis (= Conura) sp. (Hymenoptera: Chalcididae). This last species is a hyperparasitoid of D. insulare. The most abundant and frequently occurring species was D. insulare, it occurred in both localities during all three cropping seasons. The highest levels of parasitism caused by D. insulare on DBM were registered in the spring-summer season of 1996 at `La Soledad' farm with averages of 42.7, 45.0 and 44.5% on cabbage, broccoli, and cauliflower, respectively. Because D. insulare was detected attacking the pest at very low population densities during the initial stages of the crop cycle, it is assumed that the parasitoid has a high searching capacity. Correlation (r) between DBM and D. insulare population numbers was positive and significant. The other species identified occurred sporadically and had little impact on pest populations. The identification of D. collaris represents the first record of this species in North America.     

Intra- and interspecific sex-pheromone communication in the genus Yponomeuta

ABSTRACT. Male behaviour in pheromone attraction was observed in a wind tunnel for eight species of the genus Yponomeuta Latreille. Interspecific responses and the effect of some combinations of female pheromones were also investigated. The results show that the species Yponomeuta cagnagellus (Hübner), Y. malinellus Zeller, Y. rorellus (Hübner) and Y. plumbellus (Den. & Schiff.) are isolated by the specificity of their female pheromones. The other four species, Y. evonymellus (L.), Y. padellus (L.), Y. irrorellus (Hübner) and Y. vigintipunctatus (Retz.), exhibited interspecific pheromone attraction.     

Oviposition in Yponomeuta cagnagellus: the importance of contact cues for host plant acceptance

Small ermine moths (Yponomeutidae: Lepidoptera) are specialist herbivores. Species within the genus Yponomeuta are each specialized on a limited number of plant species, mainly within genera belonging to the Celastraceae. European Yponomeuta species have developed new specialized host affiliations, mainly on rosaceous hosts. Since these host shifts are reputed to be of consequence for speciation, the role of the ovipositing female is of particular interest. Study of the pre-oviposition behaviour of gravid Y. cagnagellus (Hb.) moths on host (Euonymus europaeus), non-host (Crateagus monogyna) and artificial oviposition substrates, provided information on the nature of the cues used for host plant acceptance and the insect’s perception of these cues. Host selection by adult females occurs with contact chemoreceptors probably located on the antennae or tarsi. MeOH-soluble, non-volatile phytochemical compounds washed from the host plant’s surface and applied on an artificial twig are sufficient to stimulate a complete sequence of behavioural elements leading to oviposition. Volatiles do not have a large effect on the pre-oviposition behaviour.     

A review of the hymenopterous natural enemies of cockroaches with emphasis on biological control

The biology and control potential of hymenopterous natural enemies of cockroaches in the familiesAmpulicidae, Evaniidae, Pteromalidae, Eulophidae, Eupelmidae, andEncyrtidae were reviewed. Emphasis was placed on parasitoids of major domiciliary cockroach species, especially those that have had experimental usage in integrated pest management programs. The most promising parasitoids wereAprostocetus hagenowii (Ratzeburg) againstPeriplaneta species, andComperia merceti (Compere) againstSupella longipalpa (F.). Future research should focus on practical augmentation techniques that will be compatible with other control methodologies in the urban environment.      

Parasitization of potato tuber moth eggs (Lep., Gelechiidae) from irradiated adults by Trichogramma (Hym., Trichogrammatidae) and control of moth population with combined releases of sterile insect and egg parasitoid

Abstract:  Eggs of the potato tuber moth (PTM) Phthorimaea operculella (Zeller) obtained from 150 or 300 Gy-irradiated moths, either inbred or out-crossed to untreated counterparts, were offered as host material to three generalist Trichogramma species; Trichogramma cacoeciae , Trichogramma evanescens , and Trichogramma principium , to determine the ability of these egg parasitoids to develop in F₁ sterile eggs. All eggs laid by the different parental crosses at 150 or 300 Gy, were acceptable and suitable for Trichogramma development. However, in no choice and paired comparison tests, eggs from 300 Gy-irradiated parents were significantly less preferred compared with eggs laid by non-irradiated moths. No intra-specific differences for PTM sterile eggs were recorded among the tested Trichogramma species. The mean number of wasps emerging from each egg types was not affected by an interaction between parental crosses and applied doses. Overall, this study demonstrates that release of irradiated and non-irradiated moths at a 5 : 1 over-flooding ratio and Trichogramma over potatoes in small plexiglass boxes, is complementary and more effective in reducing PTM F₁-emerged progeny than either technique used alone. The results provide baseline data for developing an integrated control approach by using inherited sterility technique in conjunction with Trichogramma for PTM population suppression.     

Analysis of mitochondrial COI sequences of the Diachasmimorpha longicaudata (Hymenoptera: Braconidae) species complex in Thailand

Diachasmimorpha longicaudata is an Opiinae parasitoid used to control tephritid fruit flies, which cause tremendous economic losses of fruits worldwide. In Thailand, D. longicaudata is classified as three sibling species, DLA, DLB and DLBB, based on the morphological and biological species concepts but their genetic variation has not been studied. Therefore, we investigated the genetic differentiation of the mitochondrial COI gene to clarify the ambiguous taxonomy of this species complex. The 603‐bp COI region was sequenced from laboratory‐bred colonies and field‐collected specimens from seven locations representing five geographical regions in Thailand. DLA was associated with the host Bactrocera correcta while DLB and DLBB were associated with Bactrocera dorsalis. The interspecific nucleotide differences of COI sequences among the three groups ranged from 6.70% to 7.62% (Kimura 2‐parameter distance), which adequately separates species complexes within the order Hymenoptera and supports the current sibling species classification. The neighbor joining, maximum likelihood and consensus Bayesian phylogenetic trees constructed from COI sequences revealed that the three sibling species of laboratory and field‐collected D. longicaudata are monophyletic with 100% support. The high genetic variation and molecular phylogeny of the COI sequences were shown to discriminate between the D. longicaudata species examined in this study.