Nonglandular leaf trichomes as short-term inducible defense of the grey alder,Alnus incana (L.), against the chrysomelid beetle,Agelastica alni L.

Summary The grey alder compensates leaf area losses due to insect grazing by continuously producing new leaves throughout the vegetative period. Different degrees of defoliation were attained experimentally by a controlled release of the oligophagous beetle Agelastica alni on arbitrarily selected trees from a homogenous population of young alders. The reduction in leaf area per tree significantly influenced the density of leaf trichomes, assessed 10–30 days later, on newly sprouting leaves only. Cross-correlations between leaf area reduction and trichome density were strongest for leaves which completed unfolding 14–21 days after damage. Dualchoice assays suggested a negative influence of trichomes on oviposition rate of A. alni. Removal of trichomes by shaving demonstrated the highly significant effect of trichomes on feeding behavior of adults and larvae in dual-choice assays. The role of the induced increase in trichome density as a possible short-term defense reaction against herbivorous insects is discussed.     

Sources of variation in rapidly inducible responses to leaf damage in the mountain birch-insect herbivore system

Summary Studies on rapidly inducible resistance in trees against insect herbivores show substantial variation in the strength of responses. Here we report the results of a study which examined causes of this variation. We bioassayed the quality of leaves of two developmental phases (young vs. mature) of the mountain birch Betula pubescens ssp. tortuosa by measuring the growth of two instars of Epirrita autumnata larvae. We used only short shoot leaves from trees of a natural stand, uniform in size and age. Damage was caused by larvae and artificial tearing of leaf lamina, varying the scale and time. We separated seasonal changes in plants from instar-dependent effects of the animals by testing experimental larvae in two subsequent growth trials. We found that only larval-made damage induced responses in leaves that made the leaves significantly poorer quality for the test larvae. Artificial damage induced only weak responses, and artificial canopy-wide damage even caused slight improvement of leaf quality. Cumulative leaf damage did not strengthen birch responses. Leaves that were in the expansion phase responded to damage while fully-expanded, mature leaves showed no response. The pattern of responses indicated that there might be physiological constraints: small-scale damage induced resistance against the larvae but largescale damage did not. Prevalent weather conditions might have modified these responses. Larvae of two instars and sexes, of low- and high-density populations responded to leaf damage similarly. However, prior experience of larvae with the host plant may have affected subsequent larval performance. Variation in rapidly inducible responses in birches was caused by plant characters rather than by test animals.     

Codling moth granulosis virus: effects of its use on some other orchard arthropods

In four field trials from 1978 to 1980, sprays of codling moth granulosis virus (CpGV) plus 1·0% skimmed milk powder did not significantly affect damage to fruit by leaf rollers (tortrix moths). In laboratory tests, survival of larvae of the leaf roller Archips podana fed on leaves sprayed with CpGV plus milk was unaffected and they grew faster than on unsprayed leaves, because of the milk deposits. This might increase damage by A. podana if CpGV plus milk were applied during the feeding period of this species. In one field trial an unusual infestation of fruit by larvae of pith moth Blastodacna atra was not affected by CpGV. Azinphos-methyl significantly reduced damage by B. atru and, in one field trial where sprays were correctly timed, that by leaf rollers. CpGV had no consistently significant effects on numbers of fruit tree red spider mite Panonychus ulmi or its predators, whereas azinphos-methyl induced outbreaks of P. ulmi by killing its predators.     

油桐叶品质对油桐尺蠖生长发育的影响

用上年受害树和未受害树的当年新发叶在室内饲养油桐尺蠖幼虫,并用Folin－Denis分析方法测定了当年受害叶,受害后新发叶和未受害叶的总单宁含量。结果表明,取食受害叶的幼虫比取食未受害叶者存活率低20．5％,发有速度慢2天,体长短5.0mm,蛹重轻25．3％。叶内总单宁含量不因受害而上升,反而呈下降趋势,说明叶内成分中影响幼虫生长发育的是非单宁物质。     

Physiological responses of dogwood (Cornus florida) to infestation by the dogwood borer (Thamnosphecia scitula)

Studies of a healthy dogwood tree ( Cornus florida ) and one suffering from a chronic infestation of Thamnosphecia scitula revealed some physiological responses of dogwood to insect damage. In high light the stomatal resistances, measured with a diffusion porometer, of leaves from the infested tree were more than double those of leaves from the healthy tree, indicating that the stomata were not open as wide in the infested tree. The greater stomatal resistance and the curling of leaves from the infested tree implied a deficiency of water, but measurements with a pressure chamber revealed only slight differences in dehydration between the two trees. Furthermore, the leaf curling was not relieved by allowing infested leaves to absorb water and attain full turgor. Leaves from the infested tree contained 35 % more reducing sugars and 63 % more sucrose, but only about half the nitrogen and ash of the healthy counterparts. Photosynthesis was significantly depressed, but dark respiration was not modified in leaves of infested compared with healthy trees. Leaves from the infested tree were 85 % as large and exhibited a 35 % greater specific leaf weight than those from the healthy tree. The conductance of water by stems from the infested tree was only 61–78 % of that by the healthy tree. It is suggested that the inferred disruption of the vascular system by insect activity alters the distribution of minerals and metabolites, hastens senescence and stomatal closure, and modifies growth by diminishing stem conductance, slowing photosynthesis, reducing leaf area, and changing leaf morphology.     

Effects of microhabitat,time of day,and weather on predation of gypsy moth larvae

Summary Wire cages with different-sized meshes were placed on trunks and around leaves at different heights in oak trees and in forest litter. Gypsy moth, Lymantria dispar, instars II–V tethered with threads were placed in each cage (instars II–III only in leaf cages) as well as outside the cages. Predation of larvae decreased from near ground to mid-crown in trees and was highest in litter and very low on leaves. Predation in litter was not strongly related to cage type, suggesting that small, invertebrate predators were active there, but IV–V instars on trunks were mainly killed by relatively large predators, probably forest mice and shrews. Influences of the time of day and weather on predation were evaluated by observing tethered V–VI instars in litter and on tree trunks hourly. Ants, particularly carpenter ants (Camponotus ferrugineus) and Formica sp., and probably vertebrates, were conspicuous predators in the litter. Ants were most active at lower relative humidities, while other predators were apparently not influenced by humidity. No daily activity rhythms of predation were noted. Invertebrates appear to be important predators of larvae only in the litter whereas vertebrates, such as forest mice and shrews, also readily attack larvae on tree trunks.     

Defoliation of wild native box trees (Buxus sempervirens): Does box rust (Puccinia buxi) infection influence herbivory,survival and growth of the invasive Cydalima perspectalis?

The invasive box‐tree moth Cydalima perspectalis causes damage to horticultural box trees (Buxus spp.) in private gardens and parks in Europe and defoliation of large areas of European native box trees Buxus sempervirens, which grow in the understorey of deciduous forests. In some parts of their distribution area, wild native box trees are infected by the box rust Puccinia buxi, a fungus which does not occur in the native range of C. perspectalis. We examined whether the infection of P. buxi deters grazing by C. perspectalis in 31 wild box‐tree stands in forests in north‐western Switzerland and south‐western Germany. On average, 82.4% of the box trees were infested by P. buxi (among‐site variation 20%–100%), or 18.5% of their leaves. Linear mixed‐effect (LME) models revealed that the percentage of rust‐infected leaves was influenced by the local box‐tree density and the slope aspect of the forest site. 9.5% of the leaves showed grazing damage by C. perspectalis. LME showed that the percentage of leaves with grazing damage was affected by the elevation of the site and the percentage of rust‐infected leaves. The effect of box rust infection on food choice and larval performance (survival and growth rate) was examined in two experiments. In food‐choice tests, larvae of C. perspectalis did not show any preference for uninfected or rust‐infected box‐tree leaves and consumed similar amounts of leaf tissue of either type. However, when larvae were reared on leaves with three different degrees of P. buxi‐infection (and uninfected leaves as control), both survival and growth rate decreased with increasing degree of rust infection. This indicates that larval performance of C. perspectalis is reduced by the box rust. In the case of an outbreak of C. perspectalis, however, larvae may indiscriminately defoliate wild box trees.     

Limiting effects of low leaf-water content on the nitrogen utilization,energy budget,and larval growth ofHyalophora cecropia (Lepidoptera: Saturniidae)

Summary Hyalophora cecropia larvae were reared on leaves of wild cherry,Prunus serotina, which contained variable amounts of leaf water but otherwise did not differ in fiber, total nitrogen, and caloric content. Larvae which were fed leaves low in leaf water grew more slowly and were less efficient at utilizing plant biomass, energy, and nitrogen than those larvae fed leaves which were fully supplemented with water.Experiments were performed using excised leaves under different regimes of relative humidity and leaf water supplementation in climatic control chambers maintained at identical temperatures and photoperiod. Foodplant biomass utilization efficiencies were severely reduced by decreasing amounts of leaf water. Growth rates were halved and the efficiency of conversion of assimilated dry matter into larval biomass was reduced from 82% in the treatment with fully supplemented leaves to 34% in the driest treatment. The nitrogen utilization efficiency (N.U.E.) was reduced from 75–80% to 48%, and the relative accumulation rate of nitrogen (N.A.R.) was suppressed nearly 2-fold for larvae on low-water leaves. Relative maintenance costs (calories expended in respiration/mg tissue/day) of larvae were nearly five times higher on dry leaves than on fully supplemented leaves. Larvae on leaves which were low in water content were themselves more desiccated, and metabolized greater portions of assimilated energy, perhaps in an attempt to supplement body water with metabolic water derived from respiration.The larval rates of consumption of biomass, energy, and nitrogen were the same for all treatments, indicating that leaf water affected larval growth primarily by restricting the efficiency of utilizing these nutrients. Where water was limiting (as in tree leaves), an increased consumption rate did not appear to be a successful means of increasing growth rates. There were daily and seasonal differences in leaf water content between different trees of the same species. Although absolute differences in leaf water exist between different trees and between young and old (fully expanded) leaves of a single tree, these differences are proportional and parallel each other through daily and seasonal cycles.In spite of evolutionary adaptations of herbivores to acquire adequate water and avoid desiccation, the leaf water content naturally encountered by cecropia larvae on cherry leaves may limit their growth, especially if the R.H. is low.     

辽东栎叶片昆虫取食形状多样性及其变化模式

取食辽东栎(Quercus liaotungensis)叶片的植食性昆虫有丰富的多样性和重要的生态功能,昆虫取食叶片留下的形状变化多,易观察,可以作为昆虫物种多样性监测的手段之一。研究北京西部东灵山地区辽东栎叶片被植食性昆虫取食状况,发现昆虫对辽东栎叶片的危害非常普遍,叶片被食频率约为90％,被食面积约5％,取食状可分为9类,其中以缘食和孔食为主,共记录20余种植性昆虫,其中蛾类幼虫和甲虫是主要取食种类,植食性昆虫集中出现在叶片发育的早期阶段（5－6月份）,7月份后,大多数幼虫发育成熟,取食叶片的面积减少。各取食状所涉及昆虫种类的组成不同,面积呈不同的季节变化模式,其中缘食状和孔食状之间有显著的正相关关系;辽东栎叶片在展叶初的叶面积最低,在7月底均达到最大值,然后呈下降趋势,单位面积重量也在展叶初最低,在6月底接近最大值,不同坡向的辽东栎林和叶片面积,单位面积重量及叶片被食面积均有显著差异,偏阳坡（东南坡） 高于偏阴坡（西北坡）。以上结果表明,辽东栎叶片维持了昆虫物种多样性,昆虫的生长发育与辽东栎的物候规律相互协调,昆虫种类的取食呈季节和空间变化。     

Larval gregariousness and neonate establishment of the eucalypt-feeding beetle Chrysophtharta agricola (Coleoptera: Chrysomelidae: Paropsini)

The selective advantage offered to individuals living within groups may relate to natural enemy defence, but in leaf feeding insects may also relate to overcoming plant defences, especially with respect to feeding establishment. We conducted a series of experiments focusing on neonate larval survival, examining the effect of group size and leaf age on the survival of a eucalypt-feeding beetle, Chrysophtharta agricola , which formed groups of up to 43 larvae on the foliage of Eucalyptus nitens in the field. In the laboratory, in the absence of natural enemies, we found that initial density, leaf age and damage to the leaf margin significantly affected larval survival. Survival of solitary first-instar larvae on young foliage was around 80% whereas on older foliage it was around 11%. Prior damage to the leaf margin significantly increased survival on older leaves to around 61%. Initial larval density also affected survival, although mortality was always significantly higher on older leaves. On older leaves the larval group size above which mortality increased no further was over two-fold that on young leaves. Observations of group feeding behaviour at each instar showed that the majority of larvae (75.7%) were aligned facing away from the feeding site and that only around 7.5%, or just 1–2 larvae per group, fed at any one time. Feeding larvae chewed the leaf edge by straddling the leaf margin. Measurements of leaf margins showed that older leaves had significantly thicker leaf margins and 'thickness' ratios (leaf margin to leaf lamina proper). In the field, approximately 85% of all larvae occurred on the first two expanded leaf pairs, and larval mortality was highest between eclosion and establishment of the first instar. However, beetles apparently did not adjust clutch size according to leaf age.     

Development and Leaf Consumption by Spodoptera cosmioides (Walker) (Lepidoptera: Noctuidae) Reared on Leaves of Agroenergy Crops

Spodoptera cosmioides (Walker) (Lepidoptera: Noctuidae) is a polyphagous pest that threatens more than 24 species of crop plants including those used for biodiesel production such as Ricinus communis (castor bean), Jatropha curcas (Barbados nut), and Aleurites fordii (tung oil tree). The development and leaf consumption by S. cosmioides reared on leaves of these three species were studied under controlled laboratory conditions. The egg-to-adult development time of S. cosmioides was shortest when reared on castor bean leaves and longest when reared on tung oil tree leaves. Larvae reared on castor bean and Barbados nut leaves had seven instars, whereas those reared on tung oil tree leaves had eight. Females originating from larvae reared on castor bean and Barbados nut leaves showed greater fecundity than did females originating from larvae reared on tung oil tree leaves. Insects fed on castor bean leaves had shorter life spans than those fed on tung oil tree and Barbados nut leaves although the oviposition period did not differ significantly. The intrinsic and finite rates of increase were highest for females reared on castor bean leaves. Total leaf consumption was highest for larvae reared on tung oil tree leaves and lowest for those reared on Barbados nut leaves. We conclude that castor bean is a more appropriate host plant for the development of S. cosmioides than are Barbados nut and tung oil tree.     

Pattern of defoliation and its effect on photosynthesis and growth of Goldenrod

1. Leaf area was removed from Solidago altissima in either a dispersed pattern (half of every leaf removed) or a concentrated pattern (every other leaf removed) and effects on leaf gas exchange, vegetative growth and flowering were examined relative to undefoliated controls. Gas exchange was measured for leaves remaining after defoliation and for regrowth leaves that developed post-damage (at 7, 16 and 26 days post-defoliation).
2. Area-based photosynthetic rates of leaves remaining after defoliation were not affected by either dispersed or concentrated damage, but damage of both types enhanced area-based photosynthesis of regrowth leaves at 16 days post-defoliation and to a lesser extent at 26 days post-defoliation.
3. Dispersed damage, but not concentrated damage, stimulated mass-based photosynthesis of undamaged leaves remaining after defoliation. Undamaged leaves remaining after defoliation and regrowth leaves on damaged plants had higher specific leaf area (leaf area/leaf mass) than comparable leaves on control plants. Mass-based photosynthesis was more strongly elevated by defoliation than area-based photosynthesis because of this increase in specific leaf area.
4. Plants with dispersed damage recovered more quickly from defoliation; they had higher relative growth rates in the first week post-defoliation than plants with concentrated damage. Both types of defoliation caused similar reductions in flower production.
5. These results add to accumulating evidence that dispersed damage is generally less detrimental to plants than concentrated damage and suggest that physiological changes in leaves may be part of the reason.     

Evaluation of systemic insecticides as a treatment option in integrated pest management of the elm leaf beetle, Xanthogaleruca luteola (Müller) (Coleoptera: Chrysomelidae)

This study evaluates the efficacy of two systemic insecticides (imidacloprid and abamectin) in an operational setting and their suitability to be incorporated into an integrated pest management program. Elm leaf beetle abundance and leaf damage were compared between treated trees and untreated control trees from 1995 through 1999. Laboratory bioassays using first-instar larvae were also used to measure the toxicity of leaves collected from treated trees at varying times after treatment. Trunk injections of abamectin and imidacloprid reduced the defoliation caused by elm leaf beetle when applied after monitoring at the peak density of elm leaf beetle eggs. Treatment in the first generation appeared to provide protection against damage in that generation as well as the second and third beetle generations. Both of these materials become active within the tree canopy very quickly and are therefore compatible with a management program that determines the need for treatment based on monitoring for egg clusters at peak density of eggs. Laboratory bioassays showed no toxicity of leaves in the year following treatment.     

Leaf variation in a tree of Pourouma tomentosa (Cecropiaceae) in French Guiana

In the flora of French Guiana we find considerable within-plant variation in leaf form. We observed entire, two-lobed, and three-lobed leaves within five separate levels (tiers) of the canopy of a single individual ofPourouma tomentosa subsp.maroniensis. Five branches from each of the five tiers of the tree were collected around the axis of the trunk. From these branches five secondary branchlets were selected and all leaves excised with information recorded as to nodal position, number of leaf nodes, and fertility status of the main branch. This design produced 1015 leaves representing about 20 m² of foliar area and about 2.4 kg of blade dry weight. Our objectives were to determine if statistically significant patterns exist for leaf variation and to suggest improvements for future, general collections. The four lower tiers had 62% entire, 10% 2-lobed, and 28% 3-lobed leaves, in contrast to the top tier with 38% entire, 11% 2-lobed, and 51% 3-lobed leaves. The top tier had no fertile branches. in the lower tiers, fertile branches produced 68% entire leaves whereas nonfertile branches produced only 46% entire leaves. In the top tier, lobed leaves made up 73% of surface area, while in the lower tiers, lobed leaves made up only 48% of total surface area. We selected a random subset of 75 leaves from the 1015, for morphometric analysis using two-way ANOVA (tier×leaf type). The boundaries of leaf images were digitized and rendered into Fourier coefficients, yielding leaf surface area and two variables that quantify aspects of shape: dissection index and leaf complexity. The Fourier coefficients were averaged by tier and by leaf type to reconstruct synthetic, average leaf images. Logistic regression was used to predict the position of leaves on the tree and to provide visualization of the relationships between leaf position on the tree and leaf morphological variables. Within the tree crown, leaf surface area and leaf specific mass (LSM) increases with height, although leaf shape does not change with height. LSM does not vary with leaf form; and sun leaves are larger than shade leaves on this tree. We conducted computer sampling experiments based on exact randomization to simulate the process of obtaining all leaf shapes present in an individual tree when making field collections of varying numbers of duplicates. This also points out the importance of noting the presence of within-tree variation in leaf form on herbarium labeds. Failure to recognize leaf variation can lead to incorrect delimitation of species as well as cause overestimates of the number of species in diversity studies.     

Impact of the leaf miner<Emphasis Type="Italic"> Cameraria ohridella</Emphasis> on photosynthesis,water relations and hydraulics of<Emphasis Type="Italic"> Aesculus hippocastanum</Emphasis> leaves

The mining of leaves of Aesculus hippocastanum caused by the larvae of Cameraria ohridella leads to precocious defoliation of trees. Damage to plant productivity was estimated in terms of the photosynthetic performance as well as of leaf water relations and hydraulics of increasingly mined leaves from infested plants in comparison with the same variables measured in non-mined leaves (controls). Electron microscopy and photosynthesis measurements revealed that chloroplasts within the green portions of mined leaves were still intact and photosynthesis of these areas was close to that of non-mined leaves, i.e. damage to functional integrity of the photosynthetic system did not extend beyond the mines. Stomata below the mines were functional as they maintained their physiological kinetics but most chloroplasts in the spongy parenchyma below the mines were degraded so that a 1:1 relationship existed between photosynthesis loss and loss of leaf green areas. Leaf conductance to water vapour and transpiration rate were 60% lower in mined leaf areas but equal to controls in green portions of mined leaves. Leaf water potential was insensitive to the amount of mined leaf area and so was leaf hydraulic conductance. Anatomical observations of leaf minor veins revealed that they were structurally and functionally intact even in leaves with 90% mined surface area. Our conclusion was that the actual damage to A. hippocastanum plants in terms of loss of photosynthates and water and nutrient transport was less than that visually estimated in recent studies.     

Complex feeding tracks of the sessile herbivorous insect Ophiomyia maura as a function of the defense against insect parasitoids

Because insect herbivores generally suffer from high mortality due to their natural enemies, reducing the risk of being located by natural enemies is of critical importance for them, forcing them to develop a variety of defensive measures. Larvae of leaf-mining insects lead a sedentary life inside a leaf and make conspicuous feeding tracks called mines, exposing themselves to the potential risk of parasitism. We investigated the defense strategy of the linear leafminer Ophiomyia maura Meigen (Diptera: Agromyzidae), by focusing on its mining patterns. We examined whether the leafminer could reduce the risk of being parasitized (1) by making cross structures in the inner area of a leaf to deter parasitoids from tracking the mines due to complex pathways, and (2) by mining along the edge of a leaf to hinder visually searching parasitoids from finding mined leaves due to effective background matching of the mined leaves among intact leaves. We quantified fractal dimension as mine complexity and area of mine in the inner area of the leaf as interior mine density for each sample mine, and analyzed whether these mine traits affected the susceptibility of O. maura to parasitism. Our results have shown that an increase in mine complexity with the development of occupying larvae decreases the probability of being parasitized, while interior mine density has no influence on parasitism. These results suggest that the larval development increases the host defense ability through increasing mine complexity. Thus the feeding pattern of these sessile insects has a defensive function by reducing the risk of parasitism.     

Using classification tree analysis to reveal causes of mortality in an insect population

• 1 Invasive species pose significant threats to native and managed ecosystems. However, it may not always be possible to perform rigorous, long‐term studies on invaders to determine the factors that influence their population dynamics, particularly when time and resources are limited. We applied a novel approach to determine factors associated with mortality in larvae of the sawfly Profenusa thomsoni Konow, a leafminer of birch, and a relatively recent invader of urban and rural birch forests in Alaska. Classification tree analysis was applied to reveal relationships between qualitative and quantitative predictor variables and categorical response variables in a large data set of larval mortality observations.
• 2 We determined the state (living or dead) of sawfly larvae in samples of individual leaves. Each leaf was scored for variables reflecting the intensity of intra‐specific competition and leaf quality for leafminers, year of collection and degree‐days accumulated were recorded for each sample. We explored the association of these variables with larval state using classification tree analysis.
• 3 Leafminer mortality was best explained by a combination of competition and resource exhaustion and our analysis revealed a possible advantage to group feeding in young larvae that may explain previously observed patterns of resource overexploitation in this species. Dead larvae were disproportionately found in smaller leaves, which highlights the potential effect of competition on mortality and suggests that smaller‐leaved species of birch will better able to resist leafminer damage.
• 4 We show that classification tree analysis may be useful in situations where urgency and/or limited resources prohibit traditional life‐table studies.