Direct and indirect effects of plant genetic variation on enemy impact

Abstract.

• 1 The Tritrophic and Enemy Impact concepts predict that natural enemy impact varies: (a) among plant genotypes and (b) may depend on the abundance of heterospecific herbivores, respectively. I tested these predictions using three herbivore species on potted, cloned genotypes of Salik sericea Marshall in a common garden experiment.
• 2 Densities of the leaf miner (Phyllonorycter salicifoliella (Chambers)) and two leaf galling sawflies (Phyllocolpa nigrita (Marlatt) and Phyllocolpa eleanorae Smith and Fritz) varied significantly among willow clones, indicating genetic variation in resistance.
• 3 Survival and natural enemy impact caused by egg and larval parasitoids and/or unknown predators differed significantly among willow clones for each of the three herbivore species, indicating genetic variation in survival and enemy impact.
• 4 Survival of Phyllonorycter was negatively density-dependent among clones.
• 5 Survival of Phyllonorycter and Phyllocolpa eleanorae were positively correlated with densities of heterospecific herbivores among clones and parasitism of these species were negatively correlated with densities of the same heterospecific herbivores among clones.
• 6 At least for Phyllonorycter this positive correlation may suggest either facilitation of survival between herbivore species, which do not share natural enemies, or an apparent interaction caused by host plant genetic variation.
• 7 Among clones, egg parasitism of Phyllocolpa eleanorae was weakly positively correlated with density of Phyllocolpa nigrita. Since these species share the same Trichogramma egg parasitoid, this interaction could support the hypothesis of apparent competition.

     

Analysis of searching movements of a leafminer parasitoid in a structured environment

ABSTRACT. Pnigalio soemius (Hymenoptera, Eulophidae) is a polyphagous ectoparasitoid of the apple leafminer Phyllonorycter cydo-niella (Lepidoptera, Gracillaridae). The tracks of females searching on the unmined leaf lamina and on the mine were segmented into 2 mm steps and analysed for angle distributions and correlations between successive steps. The movements on the mine are characterized by a higher mean and higher variance of turn angles compared to the turn angles on the leaf (kinesis-reaction family). Both klinokinesis and schemakinesis, for which a more precise definition is proposed, are present. Varying the step length has a great influence on the result and can be used to identify different orientation reactions within one track. The insect turns sharply once the mine border has been reached, a behaviour shown to be a crude taxis. An ecologically related species, Sympiesis sericeicornis (Hymenoptera, Eulophidae), moves in the same way. For this species a taxis towards the host could also be observed. A complex mixture of these orientation behaviours allows the parasitoid to remain on the mine and to localise the host.     

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.     

Seasonal biological variation in some leaf-miner parasites in the genus Achrysocharoides (Hymenoptera, Eulophidae)

ABSTRACT.

• 1 Most Achrysocharoides species and their Phyllonorycter hosts (Lepidoptera, Gracillariidae) have two generations per year in Britain.
• 2 In those species with separate sex broods, peak male emergence tends to be earlier than peak female emergence. This female emergence lag is shorter in the second generation.
• 3 The mean brood size in the second generation is significantly smaller than in the first in A.cilla males and females, A.latreilli females, mixed sex broods of A.atys, and A.carpini females. A.cilla, A.latreilli and A.niveipes generally have a significantly greater proportion of males in the second generation, but A.atys does not.
• 4 There is a shift to killing later instar Phyllonorycter larvae in the second generation, when a much higher percentage parasitism is generally achieved.
• 5 The intergeneration differences in sex ratio and brood size may be explained by a change in oviposition behaviour of females of the first and second generations.

     

Response of native parasitoids to a range-expanding host

Abstract 1. As species shift their geographic distributions, new feeding interactions with natural enemies such as parasitoids, and resources such as host plants, may be established, and existing interactions may be severed. 2. The leaf mining moth Phyllonorycter leucographella (Zeller, 1850) (Lep.: Gracillariidae) first colonised the southern United Kingdom in the mid 1980s associated with its ancestral host plant Pyracantha coccinea M. Roem. (Rosaceae), which is widely cultivated in the U.K. The moth has since spread northwards to central Scotland and has been recorded feeding on a novel host plant, Crataegus monogyna L. 3. The combined effects of latitude and time since colonisation on parasitoid community responses to the arrival of this novel host were investigated across its U.K. range. The response of parasitoids to colonisation of C. monogyna was also investigated. 4. Both the observed richness of parasitoid species associated with P. leucographella, and the proportion of P. leucographella parasitised declined with latitude and towards the current range margin. A combination of a latitudinal gradient in parasitoid and alternative host species richness is likely to lead to the trends in species richness and parasitism observed. 5. Experimental host patches exposed to parasitism beyond the current range margin of P. leucographella experienced low levels of parasitism consistent with range‐margin populations, indicating an instantaneous response by native parasitoids to availability of the novel host. Parasitism levels and numbers of associated species in the U.K. were similar to those observed in the species’ native range in Turkey. 6. The host plant switch to C. monogyna was not associated with an altered parasitoid assemblage, but rates of parasitism were significantly higher on the novel host plant. 7. Alterations in the incidence and frequency of victim‐enemy interactions as species shift their geographic ranges may be key in determining rates of range expansion and the impact invading species have on ecological communities.     

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.     

Regional climate modulates the canopy mosaic of favourable and risky microclimates for insects

1. One major gap in our ability to predict the impacts of climate change is a quantitative analysis of temperatures experienced by organisms under natural conditions. We developed a framework to describe and quantify the impacts of local climate on the mosaic of microclimates and physiological states of insects within tree canopies. This approach was applied to a leaf mining moth feeding on apple leaf tissues. 2. Canopy geometry was explicitly considered by mapping the 3D position and orientation of more than 26 000 leaves in an apple tree. Four published models for canopy radiation interception, energy budget of leaves and mines, body temperature and developmental rate of the leaf miner were integrated. Model predictions were compared with actual microclimate temperatures. The biophysical model accurately predicted temperature within mines at different positions within the tree crown. 3. Field temperature measurements indicated that leaf and mine temperature patterns differ according to the regional climatic conditions (cloudy or sunny) and depending on their location within the canopy. Mines in the sun can be warmer than those in the shade by several degrees and the heterogeneity of mine temperature was incremented by 120%, compared with that of leaf temperature. 4. The integrated model was used to explore the impact of both warm and exceptionally hot climatic conditions recorded during a heat wave on the microclimate heterogeneity at canopy scale. During warm conditions, larvae in sunlight-exposed mines experienced nearly optimal growth conditions compared with those within shaded mines. The developmental rate was increased by almost 50% in the sunny microhabitat compared with the shaded location. Larvae, however, experienced optimal temperatures for their development inside shaded mines during extreme climatic conditions, whereas larvae in exposed mines were overheating, leading to major risks of mortality. 5. Tree canopies act as both magnifiers and reducers of the climatic regime experienced in open air outside canopies. Favourable and risky spots within the canopy do change as a function of the climatic conditions at the regional scale. The shifting nature of the mosaic of suitable and risky habitats may explain the observed uniform distribution of leaf miners within tree canopies.     

Foraging behaviour of a leafminer parasitoid in the field

Abstract. 1. The searching behaviour of the parasitoid Sympiesis sericeicornis Nees (Hymenoptera, Eulophidae) is analysed under field conditions.
2. Females were tracked when flying around the canopy of apple trees infested with the tentiform leafminer Phyllonorycter cydoniella (D. & S.)(Lepidoptera, Gracillariidae) and their behaviour studied at the level of a single mine and at the level of several mines on the same leaf (patch level).
3. The presence of mines is detected during flight. The foraging behaviour at the patch level is efficient: the female is able to distinguish the suitable from the unsuitable hosts and handle the suitable hosts first. Not all mines are visited before departure from the leaf and those left unvisited are exclusively mines without suitable hosts. The patch leaving rule is discussed.
4. An ethogram of the behaviour on the mine is constructed. Selected behavioural categories are analysed for different combinations of mine content and outcome of the visit to the mine. The assessment of the mine content is done in a differential way, quickly and without mistakes.
5. The results are discussed in the context of two environmental factors, wind speed and temperature, which reduce the available time for searching. They are also related to the growing percentage of unsuitable hosts during each host generation. Tentiform mines remain attractive for a long period of time, even after parasitism or emergence of the adult, and their examination reduces the rate of host discovery.     

Mortality factors affecting the leaf-mining stages of Phyllonorycter (Lepidoptera: Gracillariidae) on oak and birch 1. Analysis of the mortality factors

Mortality of life stages of Phyllonoycter species in mined leaves of oak and birch was investigated in a Cheshire locality during 1974. Phyllonotycter species are mostly bivoltine, but P. cavella is univoltine. Parasitism and host-feeding (predation) by Hymenoptera caused most mortality. Parasitism was heaviest in the first generations, with ectoparasitism predominating, although endoparasitism caused inore mortality in the second generations. The incidence of host-feeding increased to leaf-fall when more mines contained host-fed remains than either parasites or healthy Phyllonoycter . Host-feeding occurred particularly in the first three (sap-feeding) larval instars; ectoparasitism affected mainly the two subsequent (tissue-feeding) larval stages and pupae; and endoparasitism occurred in all stages although affected hosts are killed only from the fourth larval instar onwards. Highest mortality was suffered by tissue feeders in the first generation but by sap feeders in the second. Total mortality was greatest in second generation mines and, in a given generation, survival curves for Phyllonotycter were similar on both tree species. The observed density of second generation mines relative to first in the sample was higher than predicted from first generation mortality levels, and this is discussed.