Holly leaf‐miners on two continents: what makes an outbreak species?

1. Some herbivore species periodically undergo damaging, high‐density outbreak phases followed by less damaging low‐density phases. Others maintain steady, low to moderate density levels that do little damage to their hosts. 2. Two closely related holly leaf‐miner species were compared that share many ecological traits and have very similar life cycles, but only one of which exhibits outbreaks. Phytomyza ilicicola in the eastern U.S.A. varied widely in mortality and infestation levels, reaching local densities of over 10 mines per leaf. In contrast, Phytomyza ilicis in the U.K. showed low infestation and high mortality at all sites. Using data from the literature and from field studies, the factors responsible for these contrasting dynamics were sought. 3. Phytomyza ilicicola oviposits into the leaf lamina, and experiences weak larval competition only at high densities. Phytomyza ilicis oviposits into the leaf midrib, which leads to high mortality of young larvae before mine formation. Multiply mined leaves were therefore very common in P. ilicicola but rare in P. ilicis. 4. Differences in the parasitoid complexes of the two systems accounted for further differences in survival to adulthood. The main (larval) parasitoid, which was found to impose high, density‐dependent mortality on P. ilicis, is missing on P. ilicicola. It is replaced by an egg–pupal parasitoid, which varies in its impact at differe～t sites. Multiple emergence of adults from multiply mined leaves is therefore widespread in P. ilicicola but does not occur in P. ilicis. 5. The differences in oviposition behaviour and in the parasitoid complexes are likely to allow P. ilicicola to outbreak when habitat conditions are favourable, while P. ilicis is always tightly regulated.     

Carbon dioxide-induced changes in beech foliage cause female beech weevil larvae to feed in a compensatory manner

The phenology of Fagus sylvatica was unaffected by exposure to an atmosphere of elevated CO₂ (600 μL L^-1) after two years of fumigation. Non-significant changes in nitrogen and phenolic content of the leaves decreased the nutritional status of beech for female larvae in elevated CO₂ such that they responded by eating in a compensatory manner; males were unaffected. Rates of development, mortality and adult biomass of Rhynchaenus fagi were no different from those in ambient CO₂ concentrations (355 μL L ^-1). It is possible that, with the changes in leaf chemistry affecting the females, fecundity will be altered, with important consequences for populations of beech weevil.     

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.

     

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.

     

Population dynamics and functional response of Citrostichus phyllocnistoides (Narayanan) (Hym., Eulophidae) on citrus leaf-miner, Phyllocnistis citrella Stainton (Lep., Phyllocnistidae) in Fuzhou region of south-east China

Abstract:  Population dynamics of the citrus leaf-miner (CLM) Phyllocnistis citrella Stainton, and its major parasitoid Citrostichus phyllocnistoides (Narayanan) were investigated from June to September 1994 at the teaching and research farm of the Fujian Agricultural and Forestry University (FAFU), south-east China. Based on the field investigation, the functional and numerical responses of C. phyllocnistoides were studied in laboratory experiments. During the survey period, four peaks of CLM and C. phyllocnistoides populations were observed in the growing stage of summer shoots and two peaks of CLM and C. phyllocnistoides populations, respectively, in the growing stage of fall shoots. C. phyllocnistoides is the most abundant parasitoid both in seasonal and spatial dynamics, and demonstrates a Holling type III functional response to CLM. Parasitism varied with the densities of both parasitoid and host. A 1 : 10 ratio was recorded to be an optimal proportion of C. phyllocnistoides to CLM.     

INFLUENCE OF RELATIVE HUMIDITY ON SURVIVAL OF THE CITRUS LEAF-MINER, PHYLLOCNZSTZS CITRELLA STAINTON (LEPIDOPTERA: PHYLLOCNISTIDAE)

Abstract The relationshLps between relative humidity (RH) and survival rates of eggs, all larval stages and pupae of the citrus leaf-miner, Phyllooiistis citrella Stainton, were determined by laboratory experiments. The survival of the citrus leaf-miner was observed at seven levels of relative humidity from 35% RH to 95% RH at intervals of 10% RH, with 12 L: 12 D photoperiod and temperatiure (29±0.5) C. The relative humidity was controlled by saturated solutions of MgCl₂ 6H₂O, K₂CO₃ 2H₂O, C₆H₁₂O₆, NaNO₂, NaCl, KCl, and Pb(NO₃)₂. The results showed that lower relative humidity is unfavorable for incubation of the eggs, survival of the larvae and eclosion of the pupae. The survival rates increased generally with rising of relative humidity within the range of 35% - 85% RH, and the maximum survival rates occurred at 85% RH for different life stages. The variations in hatching rates of the eggs, survival rates of the larvae and emergence rates of the pupae were great, but unimodal at different relative humidity. The effect of relative humidity on survival rates of the citrus leaf-miner could be simulated by regression analysis, using a polynomial function of three orders, and the results of fitting the model to the observed data are presented and discussed.     

Colonization, survival, and causes of mortality of Camerariu hamadryadella (Lepidoptera: Gracillariidae) on four species of host plants

Abstract.

• 1 The abundance, survival, and causes of mortality of Cameraria hamadryadella (Clemens) (Lepidoptera: Gracillariidae) were examined on four host plant species in Virginia, U.S.A. Quercus alba L. and Q.rubra L. are native within the geographic range of C.hamadryadella, and Q.robur L. and Q.benderi Baenitz are exotic. Q.robur is native to Europe, North Africa, and Asia and was probably introduced prior to 1850, and Q.benderi is of hybrid origin and introduced to cultivation before 1900. Q.alba and Q.robur are in the subgenus Lepidobalanus (white oaks), and Q.rubra and Q.benderi are in the subgenus Erythrobalanus (red oaks).
• 2 Larval mines of C. hamadryadella were abundant on both white oak species, including the exotic Q.robur, but were rare on host plants in the red oak subgenus. Un-hatched eggs of C.hamadryadella were not observed on red oaks. Other observations on host distribution indicate that C.hamadryadella is rarely found on red oaks. These observations are interpreted as circumstantial evidence that C. hamadryadella generally avoids ovipositing on red oaks.
• 3 Survival of C.hamadryadella to the adult stage was similar among all host species, but larvae tended to survive longer on hosts in the red oak subgenus. The observation of higher survival rates of early instar larvae on red oaks suggests that no nutritional or secondary chemical barrier reinforces the observed pattern of oviposition.
• 4 Significant differences in the distribution of the causes of mortality were detected between native and exotic host plant species. Larvae and pupae on native hosts were more likely to die because of predation, while those on exotic host plants were more likely to die because of parasitism and host feeding by adult female parasitoids. This pattern could arise because parasitoids prefer to forage on exotic host plants or because predators avoid foraging on exotic plants.
• 5 This study shows for C. hamadryadella that the only barriers to colonization and use of exotic hosts, in the white and red oak subgenera, are the presence of cues sufficient to stimulate oviposition and/or the absence of cues to deter oviposition. It also suggests that the presence of closely related native host plants in the region of introduction will increase the probability that exotic plants will be colonized by phytophagous insects.

     

Elevated carbon dioxide affects leaf-miner performance and plant growth in docks (Rumex spp.)

Exposure of R. crispus and R. obtusifolius to elevated CO₂ (600 ppm) resulted in an increased C:N ratio of leaf tissue and greater leaf areas. Larvae of P. nigritarsis mining leaves of R. obtusifolius during exposure produced significantly bigger mines in elevated than in ambient (350 ppm) conditions. There were no significant treatment effects on pupal weight although in both host species mean weight was greater in ambient than in elevated conditions. These results are consistent with the hypothesis that insect herbivores compensate for increased C:N ratios by increased food consumption. This response by herbivores may partially offset predicted increases in plant biomass in a future high CO₂ environment.     

Plant green-island phenotype induced by leaf-miners is mediated by bacterial symbionts

The life cycles of many organisms are constrained by the seasonality of resources. This is particularly true for leaf-mining herbivorous insects that use deciduous leaves to fuel growth and reproduction even beyond leaf fall. Our results suggest that an intimate association with bacterial endosymbionts might be their way of coping with nutritional constraints to ensure successful development in an otherwise senescent environment. We show that the phytophagous leaf-mining moth Phyllonorycter blancardella (Lepidoptera) relies on bacterial endosymbionts, most likely Wolbachia, to manipulate the physiology of its host plant resulting in the ‘green-island’ phenotype—photosynthetically active green patches in otherwise senescent leaves—and to increase its fitness. Curing leaf-miners of their symbiotic partner resulted in the absence of green-island formation on leaves, increased compensatory larval feeding and higher insect mortality. Our results suggest that bacteria impact green-island induction through manipulation of cytokinin levels. This is the first time, to our knowledge, that insect bacterial endosymbionts have been associated with plant physiology.