Approaching risk assessment of complex disease development in horse chestnut trees: a chemical ecologist's perspective

Abstract The chemo‐ecological predispositions were investigated for the development of a complex disease on the basis of an insect–fungus mutualism using the system of horse chestnuts (Aesculus hippocastanum and Aesculus x carnea), the horse chestnut leaf miner (Cameraria ohridella) and the biotrophic powdery mildew (Erysiphe flexuosa). Both C. ohridella and E. flexuosa can appear on the same horse chestnut leaf tissue simultaneously. The olfactory detection of fungal infection by the insect, its ability to discriminate the potentially mutualistic fungus from other fungi and the impact of fungal infection on insect oviposition were examined. Gas chromatography coupled with mass spectroscopic and electroantennographic detection by C. ohridella (GC‐MS/EAD) was used to assess the olfactory detection of fungal‐infected A. hippocastanum and A. x carnea leaves by C. ohridella. Infection‐related compounds, such as benzyl alcohol, dodecane, tridecane and methyl salicylate as well as fungus‐related C8 compounds, are perceived by C. ohridella. The discrimination of E. flexuosa from another phytopathogenic fungus, such as Guignardia aesculi, is based primarily on the differing pattern of C8 compounds of these fungi. Oviposition on fungal‐infected leaves of A. hippocastanum and leaves treated with fungal‐related compounds showed that C. ohridella is able to respond to the modifications in the leaf volatile profiles of horse chestnuts caused by the different fungal infections. Thus, from the perception point of view, the necessary predispositions for the development of a close insect–fungus relation between the biotrophic fungus E. flexuosa and the leaf‐mining insect C. ohridella are fulfilled. However, decreased oviposition on infected leaves does not enhance the selective contact between the species. As a consequence, an important predisposition for forming an insect–fungus mutualism is not fulfilled by these two species and, according to this approach, the risk of forming a complex disease can be assessed as low.     

Variations in parasitism in sympatric populations of three invasive leaf miners

Abstract:  Parasitism was investigated in sympatric populations of three invasive gracillariid leaf miners in Europe, Cameraria ohridella , Phyllonorycter robiniella and Phyllonorycter platani to test the hypotheses that C. ohridella is less heavily parasitized by native parasitoids and attacked by fewer species than the two other invasive species. In all regions investigated, C. ohridella showed a lower parasitism rate, and its parasitoid complex was poorer in species than those of either Phyllonorycter spp. Comparisons were made between sympatric populations of C. ohridella on its main host tree, Aesculus hippocastanum , and an occasional host, Acer pseudoplatanus . Parasitism rates were similarly low and composed of the same parasitoid species on both trees. In contrast, a sympatric population of Phyllonorycter geniculella , a native species mining A. pseudoplatanus , was heavily parasitized by a totally different parasitoid complex. These results suggest that the low parasitism in C. ohridella by native polyphagous leaf miner parasitoids is due neither to its host tree, nor to a problem of synchronization between the phenology of the pest and that of its parasitoids. Instead, it probably results from the inability of the native parasitoids to locate, attack, or develop on a new host that does not have any native congener in Europe.     

The invasive alien leaf miner Cameraria ohridella and the native tree Acer pseudoplatanus: a fatal attraction?

• 1 The horse‐chestnut leaf miner Cameraria ohridella is an invasive moth in Europe and a serious pest of horse‐chestnut Aesculus hippocastanum. The moth also occasionally attacks sycamore maple Acer pseudoplatanus, when situated beside infested horse‐chestnuts.
• 2 The main objective of the present study was to provide an overview of the relationship between C. ohridella and A. pseudoplatanus and to determine whether C. ohridella has the potential to shift to this native tree.
• 3 In the field, females oviposit on different deciduous tree species. Although less frequently attacked than A. hippocastanum, A. pseudoplatanus was clearly preferred for oviposition over 12 other woody species investigated.
• 4 Surveys in Europe demonstrated that the majority of A. pseudoplatanus trees found beside infested A. hippocastanum had mines of C. ohridella, even though more than 70% of the larvae died within the first two instars. Attack rates and development success greatly varied from site to site. Attack levels on A. pseudoplatanus were not always correlated with those on A. hippocastanum, and mines on A. pseudoplatanus were sometimes observed beside weakly‐infested A. hippocastanum.
• 5 Field observations, experimental exposure of A. pseudoplatanus saplings and rearing trials in a common garden study showed that individual trees may vary in their susceptibility to C. ohridella, whereas there was no evidence that C. ohridella populations vary in their performance on A. pseudoplatanus.
• 6 To date, there is little evidence that C. ohridella represents a major risk for A. pseudoplatanus.

     

Leafminers help us understand leaf hydraulic design

Leaf hydraulics of Aesculus hippocastanum L. were measured over the growing season and during extensive leaf mining by the larvae of an invasive moth (Cameraria ohridella Deschka et Dimic) that specifically destroy the palisade tissue. Leaves showed seasonal changes in hydraulic resistance (R_lamina) which were related to ontogeny. After leaf expansion was complete, the hydraulic resistance of leaves and the partitioning of resistances between vascular and extra‐vascular compartments remained unchanged despite extensive disruption of the palisade by leafminers (up to 50%). This finding suggests that water flow from the petiole to the evaporation sites might not directly involve the palisade cells. The analysis of the temperature dependence of R_lamina in terms of Q₁₀ revealed that at least one transmembrane step was involved in water transport outside the leaf vasculature. Anatomical analysis suggested that this symplastic step may be located at the bundle sheath where the apoplast is interrupted by hydrophobic thickening of cell walls. Our findings offer some support to the view of a compartmentalization of leaves into well‐organized water pools so that the transpiration stream would involve veins, bundle sheath and spongy parenchyma, while the palisade tissue would be largely by‐passed with the possible advantage of protecting cells from short‐term fluctuations in water status.     

Host size selection and sex allocation behaviour among parasitoid trophic levels

Abstract.

• 1 Sex ratio in the ectoparasitoid Chrysocharis nephereus, when attacking larvae of the lepidopteran leafminer Cameraria jacintoensis inhabiting the scrub oak Quercus dumosa, varies as a function of host size.
• 2 Female progeny are overproduced on large hosts and male progeny are overproduced on small hosts. Whether a host is perceived as large or small depends on the distribution of sizes of hosts encountered. Lastly, the primary sex ratio is consistently male biased. These patterns are not due to variation in progeny sex specific mortality, but are probably a result of maternal choice.
• 3 Female wasp size increases at a greater rate than does male wasp size with an equivalent increase in C.jacintoensis larval size. Although the average adult longevity of female C.nephereus is significantly greater than the average longevity of male C.nephereus, male wasp longevity increases at a greater rate than does female wasp longevity with an equivalent increase in C.jacintoensis larval size.
• 4 Chrysocharis nephereus developing on large C.jacintoensis are more likely to be hyperparasitized by Closterocerus sp. than are C.nephereus developing on smaller sized C.jacintoensis. Closterocerus sp. hyperparasitize female C.nephereus progeny with significantly greater frequency than male progeny.
• 5 Female C.nephereus overproducing daughters on large hosts and sons on small hosts may experience fitness trade-offs between the benefits associated with producing offspring on large hosts and costs associated with these offspring becoming hyperparasitized.

     

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.     

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.     

Density-related foraging behaviour in Closterocerus tricinctus, a parasitoid of the leaf-mining moth, Cameraria hamadryadella

• 1 We examined the foraging behaviour of the parasitoid wasp, Closterocerus tricinctus (Ashmead) (Chalcidoidea: Eulophidae), as it visited larvae of the leaf-mining moth, Cameraria hamadryadella (Lepidoptera: Gracillariidae), in an outbreak population.
• 2 We tracked females of C.tricinctus, recording the time spent searching for mines and handling host larvae. The density of leaf-mines (host larvae) and their condition were recorded for each leaf visited. A subset of leaves visited by C.tricinctus was enclosed in fine mesh bags so that foraging success could be determined by rearing or dissection. The average density of mines and the average leaf-area mined was estimated for a random sample of leaves from each tree.
• 3 The selection of leaves upon which to forage appears to be density-dependent. C.tricinctus visits leaves with leaf-mine densities twice the average, and when switching leaves lands directly on leaf-mines 5 times more often than expected assuming random landings.
• 4 The total time spent foraging on a leaf, the average time spent handling hosts, and the total search time within leaves tend to decline on leaves with many hosts, but the observed declines are not statistically significant.
• 5 The proportion of leaf-mines visited within a leaf is strongly inversely density-dependent. 30% of visits to leaf-mines are re-visits and 29% of handling time is spent re-handling previously visited hosts. Furthermore, only 21% of visits to mines lead to successful parasitism. We suggest that self-interference and the avoidance behaviour of the host may reduce the number of visits of leaf-mines by C.tricinctus within a leaf.
• 6 The effect of the strongly inversely density-dependent foraging investment within leaves is to offset the observed density-dependent pattern of leaf visitation making the overall spatial pattern of visitation by C.tricinctus to mines of C.hamadryadella inversely density-dependent.
• 7 We suggest that the uncertainty of C.tricinctus surviving on multiply mined leaves because of density-dependent host mortality due to intraspecific competition in high-density host populations, the rarity of high-density host populations, and the rarity of multiply-mined leaves in low-density host populations combine to select against an aggregative response within leaves by C.tricinctus.

     

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

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