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  • 1
    Electronic Resource
    Electronic Resource
    Palo Alto, Calif. : Annual Reviews
    Annual Review of Entomology 47 (2002), S. 175-205 
    ISSN: 0066-4170
    Source: Annual Reviews Electronic Back Volume Collection 1932-2001ff
    Topics: Biology
    Notes: Abstract The Japanese beetle, Popillia japonica Newman, an introduced scarab, has become the most widespread and destructive insect pest of turf, landscapes, and nursery crops in the eastern United States. It also damages many fruit, garden, and field crops. This review emphasizes recent research on the beetle's biology and management. Adults feed on leaves, flowers, or fruits of more than 300 plant species. Adaptations mediating their host finding, dietary range, mating, and oviposition are discussed. We also address abiotic and biotic factors affecting population dynamics of the root-feeding larvae. Japanese beetle grubs are widely controlled with preventive soil insecticides, but options for remedial control of adults and larvae presently are limited. Advances in understanding host plant resistance, entomopathogens, and other biorational approaches may provide more options for integrated management. Despite ongoing regulatory efforts, the Japanese beetle remains a threat as an invasive species.
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  • 2
    ISSN: 1432-1939
    Keywords: Herbivory ; Popillia japonica N. ; Nutritional ecology ; Variability ; Tilia cordata L.
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology
    Notes: Abstract Adult Japanese beetles, Popillia japonica Newman (Coleoptera: Scarabaeidae), tend to aggregate and feed most heavily in the upper canopy of host plants, defoliating them from the top down. We examined characteristics of linden (Tilia cordata L.) foliage from different canopy zones and tested two hypotheses that might account for vertical stratification of feeding by this vagile, polyphagous folivore. In the field, P. japonica caused 4–12 times more damage to upper canopy leaves in full sunlight than to lower canopy leaves in sun or shade. However, this within-tree pattern apparently cannot be explained by differences in nutritional parameters (protein, water, and sugar content) or “defensive” properties (toughness, tannins) of leaves. Furthermore, beetles did not discriminate between foliage from different canopy zones in laboratory choice tests, nor were fecundity or longevity higher for beetles fed upper canopy, sun-exposed leaves. Clonal grape plants suspended from vertical posts in the field at 3.65, 1.83 or 0 m above ground showed a top-down defoliation pattern identical to that seen in linden trees. This suggests that the height of foliage per se strongly affects initial orientation and attack by P. japonica. Recent related studies have shown that both sexes of P. japonica are strongly attracted to host volatiles induced by feeding of conspecific beetles. We suggest that Japanese beetles begin to feed in the upper canopy for reasons unrelated to host nutritional variation (e.g., behavioral thermoregulation, visual orientation to the host silhouette), and that top-down defoliation follows as additional beetles are attracted to feeding-induced volatiles acting as aggregation kairomones.
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  • 3
    Electronic Resource
    Electronic Resource
    Springer
    Oecologia 78 (1989), S. 322-329 
    ISSN: 1432-1939
    Keywords: Ilex opaca ; Allelochemicals ; Oligonychus ilicis ; Plant defense
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology
    Notes: Summary Response of the southern red mite, Oligonychus ilicis McGregor, to young and one year old leaves of Ilex opaca Aiton was studied on three dates during the period of leaf expansion in the spring. Young foliage, which is rich in nutrients but also contains high levels of saponins, was found to be unsuitable for colonization by this oligophagous herbivore until the leaves had matured and levels of saponins had declined. Mites preferentially colonized and had higher survival and reproduction on sclerophyllous, one-year old leaves than on young leaves in early spring. High levels of saponins in young, second-flush leaves of shoots that had earlier been damaged by frost were again correlated with low mite survival in June. Laboratory preference tests with the fall webworm, Hyphantria cunea Drury, and the eastern tent caterpillar, Malacosoma americanum (F.), indicated that young, saponin-rich holly foliage is especially unpalatable to these highly polyphagous caterpillars. Fifth instar fall webworms required significantly longer to complete their development, attained lower weights, and had greater mortality on artificial diet amended with low concentrations of purified holly saponins than on control diet. These results support the hypothesis that the high levels of saponins in young holly leaves provide protection from herbovores until the leaves have matured and their structural defenses are developed.
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  • 4
    Electronic Resource
    Electronic Resource
    Springer
    Oecologia 75 (1988), S. 216-221 
    ISSN: 1432-1939
    Keywords: Schlerophylly ; Spinescence ; Ilex opaca ; Plant defense
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology
    Notes: Summary Although spinose teeth of holly leaves have been widely cited as an example of a physical defense against herbivores, this assumption is based largely on circumstantial evidence and on general misinterpretation of a single, earlier experiment. We studied the response of third and fifth instar larvae of the fall webworm, Hyphantria cunea Drury, a generalist, edge-feeding caterpillar, to intact American holly leaves and to leaves that had been modified by blunting the spines, by removing sections of leaf margin between the spines, or by removing the entire leaf margin. The results suggest that the thick glabrous cuticle and tough leaf margin of Ilex opaca are more important than the spinose teeth in deterring edge-feeding caterpillars. Microscopic examination of mature leaves revealed that the epidermis is thickened at the leaf margin, and that the leaf is cirucumscribed by a pair of fibrous veins. In simple choice tests neither domesticated rabbits nor captive whitetailed deer discriminated between spinescent holly foliage and foliage from which spines were removed. Nevertheless, we found little evidence of herbivory by mammals in the field, either on small experimental trees or in the forest understory. While it is possible that spinose teeth contribute to defense by reducing acceptibility of holly relative to other palatable plant species, we suggest that the high concentrations of saponins and poor nutritional quality of holly foliage may be more important than spines in deterring vertebrate herbivores. The degree of leaf spinescence and herbivory was compared at different heights with the tree canopy to test the prediction that lower leaves should be more spinescent as a deterrent to browsers. Leaves on lower branches of mature forest trees were slightly more spinescent than were upper leaves, and juvenile trees were slightly more spinescent than were mature trees. However, there was no relationship between degree of spinescence and feeding damage. The greater spinescence of holly leaves low in the canopy is probably an ontogenetic phenomenon rather than a facultative defense against browsers.
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  • 5
    ISSN: 1432-1939
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology
    Notes: Summary Three sources of mortality, intraspecific competition, parasitism, and early leaf abscission, were investigated to determine their importance in regulating populations of the leafminer Phytomyza ilicicola Loew (Diptera: Agromyzidae) on cultivated American holly trees. Spatial aggregation of leafminers among leaves resulted in density-dependent mortality from interference and from intraspecific competition for nutritional resources. Pupal weight of survivors decreased significantly as within-leaf density increased. Within-tree parasitism of P. ilicicola by Opius striativentris (Hymenoptera: Braconidae) ranged from 3%–58%, but parasitism was independent of host density both for individual leaves and on a tree-wide spatial scale. The incidence of early leaf abscission was significantly higher for mined than for unmined leaves, with about 14% of the mined leaves abscising before the adult leafminers emerged. There was high mortality of both P. ilicicola and O. striativentris within fallen leaves. These results are discussed in relation to recent hypotheses concerning the role of competition, parasitism, and early leaf abscission in population regulation of leafminers.
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  • 6
    ISSN: 1432-1939
    Keywords: Phytomyza ilicicola ; Ilex opaca ; Herbivory-Plant stress ; Demographic variation
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology
    Notes: Summary Phytomyza ilicicola (Diptera: Agromyzidae), a univoltine specialist leafminer, typically reaches higher population densities on cultivated host planted in sunny, urban sites than it does on native trees in the forest understory. I tested the hypothesis that environmental factors, more specifically differences in light intensity and soil fertility, are responsible for this observed pattern, either by affecting leaf morphology or nutritional quality, or indirectly through early leaf abscission. Clonal trees were planted at a common site and experimentally shaded and fertilized for 14 months before being exposed to oviposition by adult flies. Differences in leaf thickness, nitrogen and water status, and in the abundance, developmental rate, survival, pupal weights and abscission-related mortality of leafminers were compared among trees. Leaves from experimentally shaded trees were larger and thinner, with only two palisade cell layers as compared to three to four layers in leaves from unshaded trees. Crystals, probably of calcium oxalate, are abundant in the abaxial cell layer, and it was hypothesized that these might set mechanical constraints on larvae feeding within shaded leaves. However, there was little or no difference in leafminer abundance, developmental rate, survival to pupation, area of finished mines, or pupal weight between shaded and unshaded trees. Leafminers compensated for feeding within thinner, shaded leaves by consuming portions of the abaxial and adaxial palisade layers and leaving a thinner roof on the mine. Furthermore, there was no difference in thickness between leaves with successful or aborted mines in either shade or sun. Leaves from fertilized trees contained 37% higher total nitrogen than those from unfertilized control trees. Fertilization did not significantly affect leafminer abundance, developmental rate, mine area or pupal weight, but survival to pupation was slightly lower on fertilized trees. Abscission-related mortality of P. ilicicola was much higher in full sun than for shaded trees, opposite of what would be predicted if early abscission per se were a primary determinant of relative abundance between habitats. These results indicate that structural constraints on leafminer larvae within shaded leaves, differences in leaf nitrogen related to soil fertility, and/or differential mortality resulting from early leaf abscission are probably not the proximate causes of density variation of this leafminer between woods and urban habitats.
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  • 7
    ISSN: 1432-1939
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology
    Notes: Summary Phytomyza ilicicola (Diptera: Agromyzidae), a univoltine specialist leafminer, is one of the few insect herbivores of American holly. Adult emergence is closely synchronized with leaf flush in spring, and females make numerous feeding punctures on and oviposit in new leaves. Larvae hatch in late May and June, but their feeding period and development are prolonged so that more than 80% of the mine enlargement occurs from January until March of the following year. We propose that this unusual life cycle reflects adaptation to constraints imposed by seasonal and age-related changes in chemical and structural defenses, and in nutritional quality of holly foliage. As holly leaves age, there is a shift in allocation of defense investment away from allelochemicals, including phenolic compounds and saponins, toward leaf sclerophylly, spinose teeth, and low foliar nitrogen and water. Rapid increases in leaf toughness and decreases in nutritional quality limit availability of leaf tissues for adult feeding and oviposition to a two-to threeweek phenological window during leaf flush. Mature holly foliage is a nutritionally poor resource by nearly all criteria known to affect food quality for herbivores. This may be the main reason for the prolonged larval development of P. ilicicola. Alternatively, winter feeding and pupation in spring may be adaptations which help to ensure synchrony of adult emergence with leaf flush. Repeated puncturing by female P. ilicicola does not render leaves more suitable for larvae, nor is it a means by which females sample leaf exudate to assess leaf quality prior to oviposition. Rather, leaf puncturing occurs mostly on leaves that are relatively high in soluble nitrogen, and is apparently a means by which females obtain protein and sugars prior to and during oviposition.
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  • 8
    ISSN: 1432-1939
    Keywords: Carbon allocation ; Buprestidae ; Quercus ; Agrilus bilineatus ; Prionoxystus robiniae
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology
    Notes: Summary The twolined chestnut borer, Agrilus bilineatus (Weber) (Coleoptera: Buprestidae), attacks oaks (Quercus spp.) that have been weakened by prior environmental or biotic stress. Our earlier work showed that trees with relatively low winter starch reserves are more likely to be attacked by A. bilineatus the following summer. We hypothesized that such trees may have less energy available for defense (Callus formation and allelo-chemical synthesis) in tissues wounded by borer larvae. However, wounding experiments showed little or no relationship between winter or summer carbohydrate reserves, callus formation, radial growth, or concentrations of tannins and phenolics in wounded or nonwounded phloem tissues. Trees with relatively low winter carbohydrate reserves were again found to be attractive to adult A. bilineatus, although not all low starch trees were attacked or successfully colonized by borers. There was a trend for carpenterworm larvae, Prinoxystus robiniae (Lepidoptera: Cossidae), a generalist bark and wood borer, to be more successful in establishing galleries on low starch trees. Carpenterworms gained significantly more weight when fed phloem from trees attractive to A. bilineatus. Oaks that attracted large numbers of A. bilineatus or that were successfully colonized by the borer produced significantly less callus than did non-attacked trees when experimentally wounded at about the time of Agrilus egg hatch. Callus formation may limit the establishment of small larvae that feed slowly in the cambial region. These results indicate that current theory regarding relationships between increased tree stress and decreased allocation of energy reserves to radial growth and defense against phloem borers may be an oversimplification. We suggest that tree growth and the defensive response of phloem tissues may be limited more by the rate of carbohydrate utilization or by changes in source-sink relationships than by storage levels. Callus formation and synthesis of allelochemicals in wounded phloem may be under the same control as cambial activation, which is mediated by plant growth regulators and can be influenced by environmental conditions.
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  • 9
    Electronic Resource
    Electronic Resource
    Springer
    Oecologia 81 (1989), S. 192-197 
    ISSN: 1432-1939
    Keywords: Phytomyza ilicicola ; Ilex opaca ; Herbivory ; Plant defense
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology
    Notes: Summary Adults of Phytomyza ilicicola (Diptera: Agromyzidae), a univoltine specialist leafminer, emerge in close synchrony with leaf flush of American holly and feed on and oviposit in soft, partially expanded leaves. Early spring defoliation, such as commonly results from freezing injury to young shoots, is followed several weeks later by a second flush of young leaves from lateral buds. We simulated this phenomenon by manually defoliating whole small trees and individual shoots of large trees to test the hypothesis that freezing injury can encourage leafminer outbreaks by inducing an abundance of soft, protein rich young leaves late in the adult activity period, when availability of vulnerable leaves becomes limited. Defoliation of small trees one or two weeks after bud break resulted in six- to 13-fold increases in the incidence of feeding punctures and larval mines on second flush leaves as compared with densities on original young leaves of control trees. Similarly, we induced significant increases in feeding punctures and larval mines on second flush leaves of individual defoliated shoots, although leaves that did not open until after the flight period escaped this injury. These observations underscore the capability of adult female P. ilicicola to locate and exploit a small number of phenologically available leaves among many hundreds of older leaves on the same tree. By altering the phenology of leaf flush, certain kinds of environmental stress may predispose perennial plants to outbreaks of early season folivores that restrict their feeding or oviposition to very young leaves.
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  • 10
    ISSN: 1432-1939
    Keywords: Cucumis sativus ; Plant defense ; Herbivory ; Insect/microbial interactions
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology
    Notes: Summary Restricted (non-systemic) inoculation of cucurbits, green bean, tobacco, and other plants with certain viruses, bacteria, or fungi has been shown to induce persistent, systemic resistance to a wide range of diseases caused by diverse pathogens. The non-specificity of this response has fueled speculation that it may also affect plant suitability for arthropod herbivores, and there is limited evidence, mainly from work with tobacco, which suggests that this may indeed occur. Young cucumber plants were immunized by restricted infection of a lower leaf with tobacco necrosis virus (TNV), and upper leaves were later challenged with anthracnose fungus, Colletotrichum lagenarium, to confirm induction of systemic resistance to a different pathogen. The response of arthropod herbivores was simultaneously measured on non-infected, systemically protected leaves of the same plants. As has been reported before, immunization with TNV gave a high degree of protection from C. lagenarium, reducing the number of lesions and the area of fungal necrosis by 65–93%. However, there was no systemic effect on population growth of twospotted spider mites, Tetranychus urticae Koch, on upper leaves, nor did restricted TNV infection of leaf tissue on one side of the mid-vein systemically affect mite performance on the opposite, virus-free side of the leaf. Similarly, there were no effects on growth rate, pupal weight, or survival when fall armyworm larvae were reared on systemically protected leaves from induced plants. In free-choice tests, greenhouse whiteflies oviposited indiscriminately on induced and control plants. Feeding preference of fall armyworms was variable, but striped cucumber beetles consistently fed more on induced than on control plants. There was no increase in levels of cucurbitacins, however, in systemically-protected foliage of induced plants. These findings indicate that pathogen-activated induced resistance of cucumber is unlikely to provide significant protection from herbivory. The mechanisms and specificity of induced resistance in cucurbits apparently differ in response to induction by pathogens or herbivores.
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