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.     

Morphology of female sex pheromone gland in the horse chestnut leafminer Cameraria ohridella (Lep., Gracillariidae)

Abstract: The internal and external morphology of the female sex pheromone gland in Cameraria ohridella Deschka & Dimic, an European pest on Aesculus hippocastanum L., has been investigated by histological and electron microscopic techniques. The gland consists of a single layer of modified epidermal cells in the dorsal part of the intersegmental membrane between the eighth and ninth abdominal segments and laterally extends to the posterior apophyses. The epithelium contains large columnar- and cone-shaped cells with basally situated nuclei. The cuticle, which covers the glandular region, has a wrapped appearance and is divided into a hyalin and thickened endocuticle and a thin outer epicuticle: it considerably expands when the gland is protruded and provides a sufficiently large surface for evaporation of the pheromone. The cuticle does not show any orifices of pore channels. In the retracted position, the gland is folded within the body cavity of the seventh and eighth abdominal segments but is exposed to the environment by extension of the abdominal tip along with female calling. In virgin females, pheromone glands are well developed at least within the first days after eclosion; if copulation occurs, glandular epithelia degenerate soon. According to the current classification, the glandular type of C. ohridella most easily is consistent with eversible dorsal scent folds that are widely distributed amongst diverse taxa of Lepidoptera. However, this is the first report on the morphology of pheromone glands in the Gracillariidae.     

Towards understanding Cameraria ohridella (Lepidoptera: Gracillariidae) development: effects of microhabitat variability in naturally growing horse-chestnut tree canopy

International Journal of Biometeorology - Dwelling intensity of horse-chestnut miner (Cameraria ohridella) larvae in various leaves insolation and temperature was measured to determine whether this...     

Occurrence and phenology of parasitic Chalcidoidea on the horse chestnut leafminer,Cameraria ohridella Deschka & Dimic (Lep., Gracillariidae)

Abstract: The parasitoid spectrum of the horse chestnut leafminer, Cameraria ohridella, was examined for its adaptation to a newly introduced host. A total of 15 parasitic species belonging to the suprafamilies Ichneumonoidea and Chalcidoidea was reared, all of them are polyphagous and common on other leafminers in Europe. The abundance of the moth and its natural enemies were studied during one entire vegetation period. The phenology of some of its major parasitic species was investigated and compared with the occurrence of the leafminer with special regard to its possible use in biological pest control.     

Possibilities to control the horse chestnut leaf miner (Cameraria ohridella) in urban environments

The continuous epidemic infestation of horse chestnuts by C. ohridella is a serious aesthetic and economic problem in urban environments. The aim of the presented studies was therefore to develop control measures which meet the special needs of urban plant protection. Removal of infested leaves in autumn and appropriate disposal of the leaflitter is the only practicable measure to reduce infestation levels so far. Another two methods, which are currently in an experimental stage, had an impact on the leafminer populations. A gelformulated combination of the C. ohridella-pheromone with a contact insecticide as attract-and-kill-system proved to be efficient at low population levels. Easily applicable and safe banks of systemic insecticides turned out to be effective in killing preimaginal stages of the leafminer on young trees. The complex of parasitoids attacking C. ohridella is comparable to that of other leafminers. Nevertheless, parasitism rates are very low at present and far away from having any controlling effect on the pest insect.     

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

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Genetic variation of horse chestnut and red horse chestnut and trees susceptibility to Erysiphe flexuosa and Cameraria ohridella

Trees of Aesculus sp. are often present in European landscapes. Common horse chestnut and red horse chestnut are seriously damaged by the fungus Erysiphe flexuosa as well as the pest Cameraria ohridella. Both pathogen and pest have spread throughout Europe. A genetic background of trees resistance still remains unknown. The present study was undertaken to estimate genetic variation of A. hippocastanum and A.×carnea grown in urban greens and their susceptibility to powdery mildew and horse chestnut leafminer. According to obtained results both species were infected by E. flexuosa in a similar degree but there were significant differences in susceptibility of particular trees (ANOVA). C. ohridella damaged only A. hippocastanum. The correlation between age of trees and degree of infestation by pathogen and pest was not observed. The significant genetic variability between two Aesculus species was revealed by the analysis of molecular variance (AMOVA) where both the intra- and the inter-species variation were found to be significant. It was shown that 73.0% of the genetic variance was contributed by the differentiation between A. hippocastanum and A.×carnea, whereas 27.0% was partitioned within species. The medium level of genetic diversity of Aesculus spp. was determined using SRAP and RAPD analyses. The mean value of genetic similarity was equal to 0.55 for common horse chestnut and 0.98 for red horse chestnut. Among 40 polymorphic SRAP and RAPD markers 17 were associated with degree of leaf damage caused by C. ohridella.     

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.     

Predator complex of the horse chestnut leafminer Cameraria ohridella: identification and impact assessment

Abstract:  The control of Cameraria ohridella Deschka and Dimic (Lepidoptera, Gracillariidae) by natural enemies in Europe is poor. Thus, in the past 15 years epidemic population densities of the moth regularly caused a continuous pre-mature defoliation of horse chestnut trees, Aesculus hippocastanum L.. Whereas several studies investigated the parasitoid complex of the leafminer and revealed its inefficiency, only little is known about the predators of C. ohridella . The aim of this study was to identify the predator complex in European horse chestnut stands by visual observations and exposure experiments and to assess the impact of different predator guilds on C. ohridella populations by an exclusion experiment. Of all potential predators observed, only blue tits, Parus caeruleus L., great tits, Parus major L., marsh tits, Parus palustris L. and the southern oak bushcricket Meconema meridionale (Costa) were found to prey on the pre-imaginal leafminer stages. Bushcrickets have to our knowledge never before been recorded preying on leafminers; moreover M. meridionale showed a measurable negative impact on the C. ohridella population. However, birds seem to be of greater importance as predators of C. ohridella than arthropods. Predation rates of birds ranged from 2 to 4% of leafminer populations, which is of the same order of magnitude as published parasitism rates. We conclude that the augmentation of tit densities, for example via nesting boxes in chestnut trees, should be considered as a component of an integrated control strategy against the horse chestnut leafminer.     

Mitochondrial and microsatellite DNA markers reveal a Balkan origin for the highly invasive horse-chestnut leaf miner Cameraria ohridella (Lepidoptera, Gracillariidae)

Biological invasions usually start with a small number of founder individuals. These founders are likely to represent a small fraction of the total genetic diversity found in the source population. Our study set out to trace genetically the geographical origin of the horse-chestnut leafminer, Cameraria ohridella , an invasive microlepidopteran whose area of origin is still unkown. Since its discovery in Macedonia 25 years ago, this insect has experienced an explosive westward range expansion, progressively colonizing all of Central and Western Europe. We used cytochrome oxidase I sequences (DNA barcode fragment) and a set of six polymorphic microsatellites to assess the genetic variability of C. ohridella populations, and to test the hypothesis that C. ohridella derives from the southern Balkans (Albania, Macedonia and Greece). Analysis of mtDNA of 486 individuals from 88 localities allowed us to identify 25 geographically structured haplotypes. In addition, 480 individuals from 16 populations from Europe and the southern Balkans were genotyped for 6 polymorphic microsatellite loci. High haplotype diversity and low measures of nucleotide diversities including a significantly negative Tajima's D indicate that C. ohridella has experienced rapid population expansion during its dispersal across Europe. Both mtDNA and microsatellites show a reduction in genetic diversity of C. ohridella populations sampled from artificial habitats (e.g. planted trees in public parks, gardens, along roads in urban or sub-urban areas) across Europe compared with C. ohridella sampled in natural stands of horse-chestnuts in the southern Balkans. These findings suggest that European populations of C. ohridella may indeed derive from the southern Balkans.     

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.     

Factors favouring the development and maintenance of outbreaks in an invasive leaf miner Cameraria ohridella (Lepidoptera: Gracillariidae): a life table study

1 Life tables were constructed for seven generations of the invasive horse‐chestnut leaf miner Cameraria ohridella in Switzerland, to assess the factors allowing the moth to build and maintain outbreak densities and to identify ‘gaps’ among their mortality factors that could be targeted by new control methods. The fecundity of females was calculated and the mortality factors affecting all developmental stages determined. 2 Females contained 106.6 eggs. In captivity, the mean number of eggs laid per female varied between 34 and 82 eggs, with a maximum of 184 eggs. Egg mortality was 18–75% and was mostly due to unknown factors. 3 The main mortality factors affecting larvae and pupae in the leaves were predation by birds and arthropods in all generation and leaf senescence in the autumn generation. Parasitoids killed less than 5% of the population. 4 Nearly 90% of overwintering pupae died in dead leaves, the majority of them being killed by earthworms and other leaf decomposers. As a result, the overwintering generations were the only ones showing a net rate of increase less than one. Non‐overwintering generations had net rates of increase between four and 11, allowing populations to build up from spring to autumn. 5 The persistence of high outbreak densities in Europe is explained by a combination of at least three factors: (i) high fecundity; (ii) multivoltinism; (iii) the unusually low impact of traditional natural enemies of leaf miners and particularly parasitoids. 6 Yearly variations in population densities could be at least partly explained by differences in larval mortality due to leaf senescence and intraspecific competition in the last generation in autumn.     

Within and between seasonal changes of detoxifying capabilities of Cameraria ohridella (Lepidoptera : Gracillariidae) larvae

Cameraria ohridella is an invasive leaf miner, a severe pest of horse chestnut trees. Chemical control needs recognition of intrinsic metabolic capability to cope with external stressing factors. Our tasks were to check annual effects of generation, and host tree age on detoxifying abilities of the last larval stage. Activities of CAT, SOD, GSTPX, GST CarE and AChE were assayed in the midgut of larvae from two localities during 3 years and three generations, annually. Activities of GSTPX and GST were high, but CAT and SOD were low in relation to other lepidopteran larvae. In general, the second generation larvae had the highest activity, indicating an effective defence against allelochemicals produced by the host. Effects of host tree age were significant for SOD, GSTPX and CarE activities. Significant annual differences on enzyme profiles of CAT, SOD, CarE and AChE reflected differences of temperature and precipitation between consecutive years.     

Visual, semi-quantitative assessments allow accurate estimates of leafminer population densities: an example comparing image processing and visual evaluation of damage by the horse chestnut leafminer Cameraria ohridella (Lep., Gracillariidae)

Abstract: Qualitative or semi-quantitative visual assessments are most often used for estimating population size of herbivorous insects. The precision of these estimates, however, is often difficult to establish. A 'simulation game' with the horse chestnut leafminer, Cameraria ohridella Deschka & Dimic (Lep., Gracillariidae) shows that visual, semi-quantitative assessments can provide accurate information. Damaged areas of 411 horse chestnut leaves collected in 100 sites were closely related to mine numbers despite some variability in mine and leaf size ( R ² = 0.915; n  = 411; P < 0.001). On the basis of this relationship, two methods of population assessment are compared: (i) digital image processing of leaf damage and (ii) visual assessment using a damage key reflecting the relative infested area on each leaf (0, 0%; 1, 0–2%; 2, 2–5%; 3, 5–10%; 4, 10–25%; 5, 25–50%; 6, 50–75%; 7, 75–100%). Both methods used to estimate damage presented a similar, close relationship to the 'real' numbers of mines ( R ² = 0.858; n  = 777; P < 0.001 for image processing and R ² = 0.905; n  = 777; P < 0.001 for visual assessment). The potential of using visual assessments as an accurate and fast method in situ at the tree scale is discussed.     

Pathogenicity of entomopathogenic fungi on hibernating pupae of Cameraria ohridella Deschka & Dimic 1986 (Lepidoptera, Gracillariidae). Part 1: Pathogenicity against the naked pupa

Strains from Paecilomyces fumosoroseus, Lecanicillium muscarium, Metarhizium anisopliae and Beauveria bassiana were examined in standardized Biotest to control the horse-chestnut leaf miner (Cameraria ohridella) in her pupal stage in winter. The fungi were pathogenic against the hibernating pupae of Cameraria ohridella at dose of 1.9 x 10(7) conidia/ml. They were aggressive, led to infection, death and mouldiness of naked pupae. Even at low temperature of 5 degrees C and 12 degrees C. L. muscarium strain V24 showed the highest pathogenicity after 4 weeks against this host, close followed by P. fumosoroseus strain P6. M. anisopliae strain 72 and 8. bassiana strain B412 were also pathogen but after a long-time period. Experiments gave information for general susceptibility of naked pupae of C. ohridella under low temperatures against entomopathogenic fungi. In further examinations it has to be tested, whether fungi can infected, when the pupae stay in their natural surroundings, the pupal cell in the leaf.     

Native parasitoids and their potential to control the invasive leafminer, Cameraria ohridella DESCH. & DIM. (Lep.: Gracillariidae)

In spite of the fact that since the end of the eighties, the horse chestnut leafminer, Cameraria ohridella, has established itself throughout Europe, native predators such as ants and birds are not attuned to this neozoic species. In contrast, several parasitic wasp species already started to exploit the invasive horse chestnut leafminer, but until now parasitation rates are quite low, mainly because of asynchrony in the lifecycles of parasitoids and host. Only the removal of leaf litter, in which pupae hibernate, is at the moment a strategy to reduce the infestation level in the next year. Unfortunately, not only hibernating horse chestnut leafminers but also parasitoids are removed, and important resources for biocontrol are unused. In the current study, we investigated the potential efficiency of the horse chestnut leafminer parasitoid complex extracted from leaf litter in defined environments. Parasitoids were released at different densities to investigate density dependence in parasitation rates.Although seven different species were released in our experiments, only Pnigalio agraules turned out to be responsible for biocontrol of C. ohridella. We recorded parasitation rates of up to 35%. Overall, parasitation rates were independent of the leafminer density but increased fourfold if ten times more parasitoid individuals were released. Unfortunately, none of the parasitoid species could be established in the experimental units in the long run. Results are compared to other parasitoid-leafminer systems, and promotion of horse chestnut leafminer parasitoids to support natural selection and biological control of the horse chestnut leafminer is discussed.     

Female sex pheromone of Cameraria ohridella Desch. and Dim. (Lepidoptera: Gracillariidae): structure confirmation,synthesis and biological activity of (8E,10Z)-8,10-tetradecadienal and some analogues

Mass spectrometric investigations confirmed the structure of the female produced sex pheromone of the horse-chestnut leafminer Cameraria ohridella Desch. and Dim. to be (8E,IOZ)-8,10-tetradecadienal. Pure samples, prepared in a straightforward synthesis, were highly attractive in field tests and proved to be suitable for monitoring of flight activities and population dynamics. In mixtures with the synthetic pheromone, analogues like 9-tridecynal and 7-dodecynyl formate were shown to reduce trap catches. In electroantennographic experiments, pheromone analogues were less active than the pheromone. 9-Tridecynal was the most EAG active analogue tested, followed by 7-dodecyn-1-yl formate and 7-undecyn-1-yl formate.     

How to safely compost Cameraria ohridella-infested horse chestnut leaf litter on private compost heaps

Abstract:  The horse chestnut leafminer Cameraria ohridella (Lep., Gracillariidae) is a recently introduced invasive pest, which frequently defoliates horse chestnut trees Aesculus hippocastanum already in summer. Currently, the removal of leaf litter, in which pupae of C. ohridella hibernate, is the only effective short-term control measure available. In the past the deposition of pest-infested litter on private compost heaps was dissuaded because of the risk of leafminer emergence in the following spring. Thus, the aim of this study was to test safe ways to compost pest-infested horse chestnut leaf litter on small heaps so that private gardeners can save the costs for leaf disposal. The number of C. ohridella adults emerging from heaps covered with either soil or uninfested foliage was reduced by 96% compared with controls. Only single leafminers emerged from the top covered heaps and there was no difference between the use of soil or uninfested foliage as cover. However, cover did not prevent the emergence of a few leafminers over the sides of the heaps. Overall, we conclude that it is relatively safe to compost C. ohridella -infested A. hippocastanum leaf litter on small compost heaps provided they are properly covered.