Parasitism of Cameraria ohridella (Lepidoptera, Gracillariidae) in natural and artificial horse-chestnut stands in the Balkans

1 The horse‐chestnut leafminer, Cameraria ohridella Deschka & Dimic, was discovered during a mass outbreak on planted horse‐chestnut trees, Aesculus hippocastanum, near Lake Ohrid in Macedonia. However, several peculiarities in the leafminer's biology and ecology, such as its inefficient control through natural enemies, indicate that it may be of exotic origin. 2 In this study, the parasitism of C. ohridella was examined in natural and artificial horse‐chestnut stands in South‐eastern Europe. The size and composition of the parasitoid complex of natural stands was comparable with that known from artificial stands. No specialist parasitoid species were detected in any of the samples. The only parasitoid species found exclusively in natural stands during this investigation were the braconid Colastes braconius and the eulophid Chrysocharis phryne. Both are known to be polyphagous on many leafminers in Europe and C. braconius has been often recorded from C. ohridella in artificial stands in previous studies. Parasitism levels varied within the same range (3.6–21.0%) throughout the entire region, in both types of habitats. 3 These results suggest that the parasitoid complex of C. ohridella in natural and planted horse‐chestnut stands in Europe is typically that of an exotic, recently introduced leafminer. Consequently, C. ohridella is probably not indigenous in the Balkans, but has been introduced to the area of its discovery from another continent.     

Laboratory rearing and biological parameters of the eulophid Pnigalio agraules,a parasitoid of Cameraria ohridella

Predators as well as parasitoids native to Europe accept the exotic horse chestnut leafminer, Cameraria ohridella Deschka and Dimi? 1986 (Lepidoptera: Gracillariidae), either as prey or as host. However, the influence of these antagonists on the populations of the pest insect is so far very low. Therefore, efforts to develop an integrated pest management system against C. ohridella should include methods which foster the natural enemy complex. In the present study we developed a laboratory rearing method and investigated several biological parameters of Pnigalio agraules (Walker 1839) (Hymenoptera: Eulophidae), a dominant species in the horse chestnut leafminer’s parasitoid complex in many European regions. This native parasitoid satisfies three basic requirements for successful use as a biocontrol agent with regard to C. ohridella. The parasitoid’s fecundity, longevity and the speed of juvenile development by far exceeds that of the leafminer, enabling the parasitoid population to increase faster than that of the host. Furthermore, our results show that the impact of this species on C. ohridella populations has been previously underestimated, because non‐reproductive killing (i.e. host‐feeding and host‐stinging) of the hosts, resulting in considerable larval mortality of the leafminer, has not been quantitatively assessed. However, naturally occurring parasitoid populations have negligible impact on C. ohridella populations. Further studies, including experimental releases of P. agraules, are necessary to understand the constraints limiting the parasitoid’s performance in the field and to assess the potential benefits of releases for the control of C. ohridella.     

Digestive enzymes activity in larvae of Cameraria ohridella (Lepidoptera: Gracillariidae)

This article presents the activity of carbohydratases and proteases in the midgut of Cameraria ohridella larvae--an oligophagous pest whose preferred feeding is horse chestnuts leaves. Optimal media pH of the assayed enzymes were similar to those of other Lepidopterans. Relatively high amylase activity, as well as maltase and sucrase activities, indicates that starch and sucrose are the main digested saccharides. Trehalase activity was similar to that described in other Lepidopterans. Activities of glycosidases were significantly lower than those of disaccharidases what suggests that neither cellulose nor glycosides are important for C. ohridella. Trypsin is the main endoprotease of this pest. Like in other leaf-eaters carboxypeptidase activity was higher than that of aminopeptidase. The activity of the majority of examined enzymes increased in the following successive pest generations, which could be explained by the decreased nutritional value of older leaves. Probably this phenomenon in hydrolases activity in Cameraria is a nonspecific mechanism present at this stage of co-evolution of the horse chestnut and its pest.     

A 3 year field survey of the natural enemies of the horse-chestnut leaf miner <Emphasis Type="Italic">Cameraria ohridella</Emphasis> in Lombardy,Italy

A survey to inventory the natural enemies of the horse-chestnut leafmining moth, Cameraria ohridella Deschka and Dimic (Lepidoptera, Gracillariidae) and to evaluate their relative abundance, was conducted from 2001 through 2003 in Lombardy, northern Italy. Two predators and 10 parasitoids were found during the 3-year survey. An increasing number of parasitoid species associated with the leafminer was detected (4 species in 2001, 5 in the 2002 and 10 in 2003). Eulophidae (Hymenoptera: Chalcidoidea) were most represented: Minotetrastichus frontalis Nees, Closterocerus trifasciatus Westwood, Pnigalio pectinicornis L., Pnigalio agraules Walker, Pediobius saulius Walker, Chrysocharis pentheus Walker, Cirrospilus talitzkii Bouek, Sympiesis sericeicornis Nees, Baryscapus nigroviolaceus (Nees). Only one Ichneumonidae (Hymenoptera: Ichneumonoidea) was found during 2003: Itoplectis alternans (Grav.).     

Antibacterial and antifungal lysozyme-type activity in Cameraria ohridella pupae

Lysozyme-type antibacterial and antifungal activity in pupae of Cameraria ohridella was studied. Activity against Micrococcus luteus and Bacillus megaterium was detected in pupae extract. Also antifungal activity from C. ohridella pupae extract directed against Saccharomyces cerevisiae strain W 303 was shown. During immunoblotting two bands in pupae extract, with molecular mass of about 15 and 28 kDa were recognized by antibodies directed against HEWL. After acid electrophoresis followed by bioautography of the extract, two lytic zones showing lysozyme-type activity against M. luteus were observed. Two bacteria: Gram-positive Aerococcus viridans and Gram-negative Aeromonas salmonicida ssp. masoucida were isolated from pupae of C. ohridella. Their activity against M. luteus, B. megaterium, and S. cerevisiae W303 was detected. After immunoblotting with antibodies against HEWL, also two proteins from bacterial suspensions of A. viridans and A. salmonicida were detected, about 15 and 28 kDa.     

Recruitment of native parasitoids by an exotic leaf miner, Cameraria ohridella: host-parasitoid synchronization and influence of the environment

Abstract

• 1 The horse‐chestnut leaf miner, Cameraria ohridella, is a moth of unknown origin that has recently invaded Europe and severely defoliates the European horse‐chestnut, an important ornamental tree.
• 2 Several indigenous parasitoids have colonized this new host, but parasitism remains low. One of the hypotheses suggested to explain the low parasitism is that candidate parasitoids emerge too early in spring to attack the first host generation and, thus, need early‐occurring leaf miners as alternate hosts. This hypothesis was tested by observing the synchronization between the phenology of the moth and that of its main parasitoids, and by comparing parasitism rates and parasitoid richness in different environments with various levels of biological diversity.
• 3 In spring, the bulk of the parasitoids emerge at least 5 weeks before the occurrence of the first suitable larvae of C. ohridella whereas most parasitoid adults reared outdoors die within 5 weeks after emergence.
• 4 Parasitism rates and parasitoid richness do not increase with biological diversity, suggesting that most parasitoids attacking the first generation of C. ohridella do not come from alternate hosts. Parasitism does not increase later in the year in the subsequent generations, when host‐parasitoid synchronization becomes less critical.
• 5 We conclude that, although the spring emergence of parasitoids is not synchronized with the phenology of C. ohridella, the parasitoids attacking the first generation are probably old or late‐emerging adults of the overwintering generation. The lack of synchronization is probably not the only reason for the poor recruitment of native parasitoids by C. ohridella.

     

Cameraria ohridella: 10 years of sex pheromone and kairomone research

This review traces the identification and use of sex pheromone of the horse chestnut leafminer Cameraria ohridella Deschka et Dimi? (Lepidoptera, Gracillariidae, Lithocolletinae) to monitor population density and discuss its potential for direct control. Host volatiles endogenously emitted from Aesculus hippocastanum L. leaves are described in terms of their ability to control conspecific females.     

Spatial patterns and infestation processes in the horse chestnut leafminer Cameraria ohridella: a tale of two cities

Spatial patterns of the horse chestnut leafminer Cameraria ohridella Deschka & Dimic (Lepidoptera: Gracillariidae) population density was analysed in the cities of Bern and Brussels in order to explore its spatial population dynamics. The surveys were carried out in Bern in 1998 and 2000 and in Brussels in 2001 to assess population density in relation to local characteristics. In Brussels, population density was also measured using pheromone traps distributed over the city and collected twice per moth generation. A quantitative relationship was found between local population density (measured by pheromone traps and survey observations) and the amount of leaves left on the ground the previous fall. Several other factors were related to observed infestation levels: the occurrence of the pathogen Guignardia aesculi was inversely related to infestation by C. ohridella in Bern in 1998 and 2000, the number of horse chestnut trees within 800 m distance was positively related to infestation level in Bern in 2000, and the proportion of green areas within 100 m and the number of other horse chestnut trees within 2000 m were positively related to infestation levels in the 2001 Brussels survey. The pattern of infestation levels as a function of distance to potential population reservoirs suggested that C. ohridella re‐invades areas where overwintering leaves have been cleaned from refuge areas such as parks or urban forests. Our results indicate that the removal of leaves is a feasible first aid control measure to reduce moth population densities. However, leaf removal may not reduce moth densities when done improperly. In places where proper leaf removal is not feasible, other control measures are needed.     

Impact of the leaf miner Cameraria ohridella (Lepidoptera: Gracillariidae) and bleeding canker disease on horse‐chestnut: direct effects and interaction

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Oviposition patterns and larval damage by the invasive horse‐chestnut leaf miner Cameraria ohridella on different species of Aesculus

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Parasitism of the apple maggot,Rhagoletis pomonella,infesting hawthorns in Washington

Five species of larval parasitoids were reared fromRhagoletis pomonella (Walsh) infested fruit of hawthorn,Crataegus, collected from several locations in southwest Washington over a four year period. A braconid,Biosteres melleus (Gahan), parasitized larvae infesting fruits of a native hawthorn species,Crataegus douglasii Lindl. Another braconid,Opius downesi Gahan, emerged exclusively fromR. pomonella pupae reared from fruits of an introduced species of hawthorn,Crataegus monogyna Jacq. A pteromalid,Pteromalus sp., and two eulophids,Tetrastichus spp., attackedR. pomonella larvae infesting fruits of both hawthorn species. No parasitoids emerged from a total of 4385 pupae reared from apple. Percent parasitism ofR. pomonella was higher inC. monogyna compared toC. douglasii fruits. The highest average levels of parasitism ofR. pomonella inC. monogyna andC. douglasii fruits were 90% and 23% respectively. The kinds of parasitoids, their relative abundances and timing of parasitization on the two hawthorns was related to differences in fruit ripening patterns and its effect on the development ofR. pomonella on these two hosts. Parasitization ofR. pomonella byTetrastichus spp. is a new host record. The detection of these species andPteromalus sp. in southwest Washington are the first records of ectoparasitoids attacking this tephritid.     

Parasitism byEphedrus cerasicola [Hym.: Aphidiidae] developing in different stages ofMyzus persicae [Hom.: Aphididae]

The parasitoidEphedrus cerasicola Stary oviposited in the 4 nymphal instars and in newly moulted apterous adults ofMyzus persicae (Sulzer). Development and reproduction of unparasitized and parasitized aphids at 21°C were compared. Unparasitized aphids developed to adults in 6.5 days and started to reproduce after 7 days. Longevity varied between 7 and 42 days. Net reproductive rate (R₀) was 40.7. In contrast to older nymphs, aphids parasitized in the 1 st instar almost never reached the adult stage before mummification. Aphids parasitized in 2nd, 3rd and 4th instar and as newly moulted adults produced respectively 0.07 %, 2 %, 23 % and 32 % of offspring produced by unparasitized aphids. Corresponding reproductive periods were 1, 1.4, 3 and 4 days. Host age at parasitization had a slight effect on the parasitoid's developmental rate and had no effect on egg or pupal survival, or on the sex ratio of the emerging parasitoids.     

Sperm structure and ultrastructure of the Melittobia hawaiiensis, Perkins and M. australica, Girault (chalcidoidea: eulophidae)

Spermatozoa morphology has, for some years, been used to help answer some phylogenetic questions for Hymenoptera. This is the second study describing spermatozoa morphology of an Eulophidae species in which important characteristics were observed. Melittobia spermatozoa are spiralled and measure approximately 270mum in length. The head contains a small acrosome, apparently formed only by an acrosomal vesicle, which, together with the initial nuclear region, is surrounded by an extracellular sheath, from which innumerable filaments irradiate. The nucleus is helicoidal and completely filled with compact chromatin. A centriolar adjunct is observed at the nucleus-flagellum transition; it associates laterally with the nucleus and exhibits two small expansions, which reach around the centriole. In the flagellum there are two mitochondrial derivatives, which in cross-sections are asymmetric. In the derivative with the larger diameter, two distinct regions are observed, a small one, near the axoneme, with a clear "fissure" inside, and a larger region where the cristae occur. Both derivatives initiate at the nuclear base, but the larger diameter derivative finishes first, before the flagellum extremity. At the end of the axoneme, the accessory microtubules are the first to finish.     

Role of photoperiod and temperature in the induction of overwintering pupal diapause in the spotted tentiform leafminer,Phyllonorycter blancardella

The role of photoperiod and temperature in the induction of overwintering diapause inPhyllonorycter blancardella (F.) (Lepidoptera: Gracillariidae) was examined in the laboratory and field using leafminers from commercial apple orchards in Ontario, Canada.P. blancardella exhibited a long-day response to photoperiod: long daylengths resulted in uninterrupted development whereas short daylengths induced diapause. The estimated critical photoperiod for diapause induction was L14.25∶D9.75. The larvae of leafminers destined to enter diapause took ca. 3× longer to complete development than the larvae of non-diapausing leafminers. The development prolonging effect of photoperiod decreased with decreasing daylength. Temperature modified the diapause inducing effect of photoperiod. At L14.25∶D9.75, diapause incidence was similar at 15 and 20°C but was lower at 25°C. Photoperiod also altered the normal relationship between development rate and temperature. At L14.25∶D9.75, the duration of larval development of diapausing leafminers was similar at 15, 20 and 25°C. Temperature alone is unlikely to have a role in the induction of diapause because leafminers exposed to natural late summer and fall temperature regimes and L16∶D8 did not enter diapause.     

肉苁蓉人工控制离体寄生关键技术研究

以梭梭无菌苗和肉苁蓉萌发种子为材料,对肉苁蓉离体寄生培养基质、离体寄生方式和离体寄生环境条件等进行了研究,以明确肉苁蓉的寄生特性,为肉苁蓉人工繁殖技术体系的建立与优化奠定基础。结果表明:玻璃纤维滤纸是肉苁蓉离体寄生适宜的培养基质;在黑暗、相对湿度80%、温度18℃～30℃的环境中,萌发的肉苁蓉种子与寄主梭梭寄生培养8d后开始建立寄生关系,寄生率达82%;而光照条件下,萌发的肉苁蓉种子与梭梭不能建立寄生关系。     

The preimaginal stages of Pnigalio gyamiensis Myartseva & Kurashev, 1990 (Hymenoptera, Eulophidae), a parasitoid associated with Chrysoesthia sexguttella (Thunberg) (Lepidoptera, Gelechiidae)

The larval instars of Pnigalio gyamiensis Myartseva and Kurashev are described in detail for the first time. This species is a larval-pupal ectoparasitoid of Chrysoesthia sexguttella (Thunberg) (Lepidoptera, Gelechiidae), which forms leaf mines in the plant Chenopodium album L. (Caryophyllales: Amaranthaceae). The female of Pnigalio gyamiensis lays a single egg on the skin of the host larva or nearby it, without any significant preference for a particular variant. The presence of long hairs on its body provides the newly-hatched first larval instar with high mobility. Some peculiarities in this parasitoid-host relationship are described.     

Interacting effects of increased tannin levels on leaf-mining insects

Tannin-rich extracts were painted on leaves of Q. emoryi (Fagaceae) that contained first instar larvae of a leaf miner, Cameraria sp. (Lepidoptera: Gracillariidae, Agrifoliella group), to test the role of tannins in increasing parasitism, while controlling for changes in protein content and physical damage. Miners from leaves that were treated with tannins had higher rates of parasitism by hymenopterans, but decreased mortality from other causes. The negative and positive effects of tannins balanced since overall larval survivorship was equivalent for leaf miners on tannin-enriched and control leaves. The tannin-rich extract also affected larval development because pupal weight was decreased for leaf miners in treated leaves. Tannins can have both adverse and beneficial effects on these phytophagous insects.

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Résumé Des extraits riches en tanins ont été étalés sur des feuilles de Quercus emoryi contenant des chenilles de ler stade de Cameraria sp., pour contrôler l'influence du tanin dans l'accroissement du parasitisme sans modifier la teneur en protéines ni provoquer de dommages physiques. Les mineuses de feuilles traitées avaient des taux plus élevés de parasitisme par les hyménoptères, mais une diminution des autres causes de mortalité. Les effets négatifs et positifs des tanins se sont équilibrés, puisque la survie des mineuses était la même pour les feuilles traitées que pour les témoins. Les extraits riches en tanin ont agi sur le développement larvaire puisque le poids des chrysalides des mineuses de feuilles traitées était plus faible. Les tanins peuvent ainsi avoir des effets nuisibles et bénéfiques sur ces insectes phytophages.

     

Host Preferences of Trichogramma pretiosum and the Influence of Prior Ovipositional Experience on the Parasitism of Plutella xylostella and Pseudoplusia includens Eggs

Trichogramma wasps are generalist egg parasitoids used in biological control efforts. In a multi host situation they may preferentially parasitize a non-target host species to the detriment of the control program. Plutella xylostella Linnaeus (Lepidoptera: Plutellidae) is a very serious pest of cabbage, but is only one in a number of species in the ‘cabbageworm’ complex. We investigated the host preferences of Trichogramma pretiosum Riley (Hymenoptera: Chalcidoidea) when offered the eggs of Plutella xylostella and the eggs of Pseudoplusia includens Hübner (Lepidoptera: Noctuidae), another species in the ‘cabbageworm’ complex. Trichogramma pretiosum reared on the eggs of the factitious host Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) parasitized both Plutella xylostella and Pseudoplusia includens eggs under laboratory conditions. For both choice and no-choice experiments, T. pretiosum parasitized significantly more P. xylostella eggs than P. includens eggs. Prior ovipositional experience with one or other of the two host species had no effect on the subsequent parasitism levels of the two host species. The preference for P. xylostella eggs was also not affected by this prior ovipositional experience.