寄主植物叶片物理性状对潜叶昆虫的影响

潜叶昆虫广泛分布于鳞翅目、双翅目、鞘翅目和膜翅目中,其幼虫潜入叶片内部生活和取食,是一类用于研究植物-昆虫-天敌种间关系和协同进化的重要模式生物。有些潜叶昆虫是重要农林害虫。相比外食性昆虫,在叶内取食的潜叶昆虫幼虫更易受到叶片物理性状的直接影响。叶片的着生位置、朝向、大小、颜色和表皮毛等直接决定潜叶虫成虫的取食和产卵选择,从而影响幼虫的空间分布和寄主适应。叶片的某些物理性状也会直接影响幼虫的取食行为、生长发育和被寄生率。研究叶片物理性状的防御作用以及潜叶昆虫对这些防御的适应,有助于了解潜叶昆虫-寄主植物的协同进化。另一方面,外界环境和遗传育种都有可能改变植物叶片的物理特性,而对潜叶害虫产生抗性,从而实现潜叶害虫的可持续生态控制。     

多食性斑潜蝇对寄主植物的选择

综述了有关斑潜蝇对寄主植物选择的研究进展。斑潜蝇对寄主植物的选择表现为对植物组织的选择、对植物不同部位叶片的选择、对同种植物不同品种的选择、对不同寄主植物的选择四个方面。斑潜蝇对寄主植物的选择主要受叶表面的物理性质、植物叶的化学物质成分、成长经历等因素的影响。阐述了多食性斑潜蝇寄主植物范围扩大的原因。     

Fungal endophytes and phytochemistry of oak foliage: determinants of oviposition preference of leafminers?

Sedentary insect herbivores, such as gallformers and leafminers, are usually non-randomly distributed among and within host plants. Dispersion of these insects is largely a function of female oviposition choice. In field experiments and observations spanning two growing seasons, we tested the hypothesis that selective oviposition on individual leaves within trees by the dominant herbivore of Emory oak, the monophagous leaf-miner Cameraria sp. nov., is determined by the probability of colonization by endophytic fungi. These fungi are alleged to act as plant mutualists by deterring, killing, or inhibiting the growth of insect herbivores. We found that leaves selected by females for oviposition and paired, unmined leaves were equally likely to be colonized by fungal endophytes. Furthermore, condensed and hydrolyzable tannin levels, purported inhibitors of fungal infection, and protein content did not vary between leaves selected by females and unmined leaves, or between leaves with and without endophyte infections. We conclude that female Cameraria do not choose leaves within trees for oviposition on the basis of propensity for endophytic fungal infection or on phytochemical parameters that might indicate probability of future infections. At this spatial scale at least, fungal endophytes do not explain the highly aggregated distribution of Cameraria among leaves and associated costs in terms of increased larval mortality. Fungal endophytes may, nevertheless, affect leafminer dispersion and abundance at larger spatial scales, such as host plant populations or species. We did find, however, that the amount of mining activity on leaves is positively associated with increased colonization by fungal endophytes. We suggest that mining activity increases endophyte fungal infections by facilitating spore germination and hyphal penetration into the leaf or by altering leaf phytochemistry. The facilitation of endophyte colonization by leafmining activity coupled with the lack of predictability of endophyte infections based on leaf phytochemistry and almost 100% infectivity of all oak leaves during sporadic wet years may prevent female leafminers from discriminating leaves for oviposition on the basis of current or future levels of endophytes in leaves.     

Precipitation on abscised leaves modulates leaf‐mining beetle (Trachys yanoi) survivorship and population dynamics

1. We clarified the effects of early leaf abscission on the survivorship of the leaf‐mining beetle Trachys yanoi Y. Kurosawa (Coleoptera: Buprestidae) and the underlying mechanism in relation to weather conditions in Japan. Trachys yanoi is an insect pest of zelkova trees [Zelkova serrata (Thunb.) Makino (Rosales: Ulmaceae)]. Larvae burrow into zelkova leaves and feed on leaf tissue, causing early leaf abscission. 2. This study investigated the relationship between the beetle population and weather conditions over 10 years in a zelkova forest. The effects of moisture and temperature on adult emergence from early abscised leaves were examined in the laboratory and in the field. 3. The beetle population in the studied forest was negatively affected by high precipitation levels when the beetles still inhabited early abscised leaves. Fewer adults emerged from early abscised leaves under wet conditions than under dry conditions, in both laboratory and field tests. 4. These results demonstrate that early leaf abscission plays an important role in leaf‐mining beetle survivorship and population dynamics, and that the amount of precipitation when leaf‐mining beetles still inhabit early abscised leaves modulates this effect. 5. Because precipitation when leaf‐mining beetles still inhabit early abscised leaves was mainly driven by an East Asian rainy season front, the beetle population dynamics were affected by the activity of the front.     

Spatial distributions of the leafminer Ophiomyia maura (Diptera: Agromyzidae) in host plant Aster ageratoides

The seasonal occurrence and among-plant and within-plant spatial distribution of the multivoltine leafminer Ophiomyia maura Meigen (Diptera: Agromyzidae) on the herbaceous plant Aster ageratoides Turcz. subsp. ovatus (Asteraceae) were investigated in the field. O. maura has at least four generations a year and mines per leaf fluctuate with a mean of 0.007 throughout the occurrence period. Seasonal occurrence is associated with abundance of new host leaves, suggesting O. maura females prefer to oviposit in newly emerged leaves. The among-plant distribution of O. maura is described by a Poisson distribution early in the season but tends to be weakly clumped later. The within-plant vertical distribution of larval mines increased from middle to upper leaves during plant development, because mined leaves in the middle position early in the season move downward with the emergence of new leaves, shifting mined leaves from the position where O. maura oviposits eggs. Later in the season, mined leaves remain where they are deposited because few new leaves emerge. The spatial distribution of O. maura , resource utilization patterns, and host plant characteristics are discussed.     

Leaf mines as visual defensive signals to herbivores

Leaf‐mining insects produce conspicuous and distinct leaf mines on various types of plant leaves. The diversity of leaf‐mine morphology has typically been explained by several factors, such as selective feeding on plant tissues, improvement of microclimate, faecal disposal, reduction in the efficiency of parasitoid search behaviour and leafminer phylogeny. Although these factors are certainly associated with mining patterns, masking the mines, rather than making them conspicuous, appears to be more advantageous for deterring parasitoids and predators of leafminers. However, here, I propose that prominent leaf mines may serve to signal or cue herbivores to avoid feeding on the mined leaves. Because most leafminers are sessile and complete their development within a single leaf, herbivory of mined leaves is detrimental to leafminer survival. Other herbivores appear to avoid consuming mined leaves for a variety of reasons: leaf mines mimic leaf variegation or mottling; mined leaves induce chemical and physical defences against herbivores; and leaf mines mimic fungal infection, animal excrement, and necrosed plant tissues. Hence, natural selection may have favoured leafminers that produce conspicuous mines because of the increased survival and fecundity of thereby reducing herbivory on mined leaves.     

江苏地区三叶斑潜蝇和美洲斑潜蝇的发生危害及种群动态

【目的】三叶斑潜蝇Liriomyza trifolii和美洲斑潜蝇Liriomyza sativae是一类危害蔬菜、花卉的世界性害虫,也是我国重要的外来入侵害虫。了解其发生危害特征,可为这两种斑潜蝇的防治提供科学依据。【方法】于2015年对江苏地区的两种斑潜蝇的发生分布、危害程度以及在扬州地区的发生动态进行了调查,采集含有潜道的叶片,待蛹羽化为成虫后鉴定斑潜蝇种类,同时统计叶片上的潜道数量划分危害等级。【结果】美洲斑潜蝇主要在苏北地区发生危害,三叶斑潜蝇主要在苏南、苏中地区发生危害;两种斑潜蝇在江苏地区的发生危害程度均未超过3级。在扬州地区的蔬菜上仅调查到三叶斑潜蝇的危害,其危害的寄主植物有6科、12种,主要包括茄科、豆科、葫芦科、十字花科的蔬菜。三叶斑潜蝇的发生危害从3月开始,11月结束,全年有3个发生危害高峰,其中在7月下旬至8月初的发生危害最重。【结论】两种斑潜蝇在江苏地区的种类分布呈现出明显的地理差异,同时扬州地区三叶斑潜蝇的虫口密度随寄主和时间不同呈现的明显,因此生产上需加强对斑潜蝇发生危害进行监测,并指导斑潜蝇防治。     

How to safely compost Cameraria ohridella-infested horse chestnut leaf litter on private compost heaps

Abstract:  The horse chestnut leafminer Cameraria ohridella (Lep., Gracillariidae) is a recently introduced invasive pest, which frequently defoliates horse chestnut trees Aesculus hippocastanum already in summer. Currently, the removal of leaf litter, in which pupae of C. ohridella hibernate, is the only effective short-term control measure available. In the past the deposition of pest-infested litter on private compost heaps was dissuaded because of the risk of leafminer emergence in the following spring. Thus, the aim of this study was to test safe ways to compost pest-infested horse chestnut leaf litter on small heaps so that private gardeners can save the costs for leaf disposal. The number of C. ohridella adults emerging from heaps covered with either soil or uninfested foliage was reduced by 96% compared with controls. Only single leafminers emerged from the top covered heaps and there was no difference between the use of soil or uninfested foliage as cover. However, cover did not prevent the emergence of a few leafminers over the sides of the heaps. Overall, we conclude that it is relatively safe to compost C. ohridella -infested A. hippocastanum leaf litter on small compost heaps provided they are properly covered.     

The Compatibility of Control Agents used for the Control of the South American Leafminer, Liriomyza huidobrensis

The compatibility of infective juveniles (IJs) of the entomopathogenic nematode, Steinernema feltiae, the parasitic wasps Diglyphus isaea and Dacnusa sibirica, and chemical insecticides to control larval stages of the South American leafminer, Liriomyza huidobrensis, on leafy salad crops was investigated. In intact leaf mines, leafminer larvae already parasitized by D. isaea which had developed to the larval or pupal stage, and D. sibirica in the larval stage were subsequently also infected by S. feltiae following a foliar application. This reduced the potential of the wasps to survive until the adult stage. In similar tests foliar applied chemical insecticides reduced the survival to adulthood of D. sibirica and their host larvae. However, a soil drench of imidaclorpid did not cause a significant reduction in the number of D. sibirica which survived the treatment and developed to adult emergence. Tests to investigate the utilisation of parasitoids following foliar applications of nematodes found that adult D. isaea did not discriminate between the healthy and nematode-infected leafminer larvae for host feeding, although 98% of eggs laid by the female wasps were deposited alongside healthy larvae.     

Leaf abscission phenology of a scrub oak: consequences for growth and survivorship of a leaf mining beetle

Brachys tessellatus is a leaf-mining beetle that attacks Quercus laevis (turkey oak), a deciduous scrub oak in the fall line Sandhills of the southeastern United States. This oak species varies substantially in leaf abscission phenology. In the fall of 1994 we examined leaf abscission patterns at three sites in central South Carolina and found that leaves containing active miners abscised earlier than leaves without active miners. To quantify the effect of leaf abscission phenology on the survivorship and body size of leaf miners within the abscised leaves, we marked naturally and manually abscised leaves, left leaves on the ground to over-winter, and then collected leaves before beetles emerged the following spring. We found that beetles inside leaves that were abscised early in the season were more likely to be killed by predators, and that they produced smaller pupae than beetles within leaves abscised later in the season. We also evaluated the relationship between adult beetle body size and fecundity; body size of B. tessellatus is positively correlated with lifetime fecundity and daily oviposition rate indicating that effects of leaf-abscission phenology on beetle body size likely translate into effects on beetle fitness. We discuss how B. tessellatus has adapted to variation of leaf abscission phenology of oaks by evolving developmental plasticity that permits successful completion of larval development over a range of season lengths.     

Lydella thompsoni Herting (Dipt., Tachinidae), a parasitoid of the European corn borer, Ostrinia nubilalis Hbn. (Lep., Pyralidae) in Slovakia, Czech Republic and south-western Poland

This paper shows that, within Central Europe, parasitism of the European corn borer (ECB) by the tachinid parasitoid, Lydella thompsoni (LT) increased from 0.47 to 1.49% in south-western Poland (51°03'N), to 4.31–21.95% in eastern Slovakia (48°20'N). The synchrony between the parasitoid LT and its primary host, the ECB, was studied in Central Europe under conditions where the host is univoltine, but the parasitoid is bivoltine. A cumulated total of more than 400 LT was field-collected from overwintering ECB larvae. The parasitoid hibernated as larva inside the host. Pupation started in the second half of the following March and 50% of pupation was surpassed in the first half of April. The first parasitoid adults emerged at the end of April and the majority at the beginning of May. Development threshold temperatures for 50% pupation was determined to be 2.7°C, and for 50% adult emergence 5.0°C; the respective thermal constants were 178.8–179.8 and 237.7–251.8 Celsius degree-days. Emerged adults did not parasitize overwintered ECB larvae in spring, hence there must be an alternate host for the first generation of LT in areas of univoltine life cycle of the ECB. Parasitization of the ECB larvae by LT continued until the end of July. The first parasitoid adults from this second generation emerged in the second half of August. By the end of the season, nearly one-third of LT adults had emerged. The rest of this generation apparently overwintered in the larval stage.     

Quantitative food webs of lepidopteran leafminers and their parasitoids in a Japanese deciduous forest

Quantitative food webs were constructed to explore the community structure of leaf-mining moths in the family Gracillariidae and their parasitoid wasps in a deciduous forest in Hokkaido, Japan. A whole food web was constructed from data collected from June to October 2001. In the web, 16 leafminer species on seven tree species were attacked by 58 species of hymenopteran parasitoid; 376 links between leafminers and parasitoids were observed. Leafminers were specialist herbivores, but most parasitoids were generalists. Five webs were constructed for the seasonal prevalence of leafminers over the one-year period to reveal the temporal dynamics in community structure. Among the seasonal webs, the first web in June was distinctive because two tree species, Japanese umbrella tree Magnolia obovata and Japanese magnolia M. kobus, supported the community. Second to fourth webs from July to September were dominated by the leafminer species on Japanese oak Quercus crispula, and the fifth web was marked by that on Carpinus cordata. The extent of potential apparent competition among leafminers was evaluated using quantitative parasitoid overlap diagrams. These diagrams suggested that abundant host species are likely to have large indirect effects on less abundant species. Moreover, the potential for apparent competition between leafminer species inhabiting different host tree species can occur, although leafminers sharing the same tree species are prone to interact via shared parasitoids. In this system, particular leafminer species, acting as potential sources of apparent competition, can affect other species as sinks, and control whole-community dynamics. Directed apparent competition may potentially occur around oak trees.     

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.

     

First record of a walnut shield bearer Coptodisca (Lepidoptera: Heliozelidae) in Europe

A leafminer of the Nearctic genus Coptodisca Walsingham (Lepidoptera: Heliozelidae), a species of potential economic interest, is reported for the first time from Europe, infesting the black (Juglans nigra L.) and the common walnut (Juglans regia L.). Mines were collected since September of 2010 in several sites of two Italian regions (Campania and Lazio). The species is rather similar to Coptodisca juglandella (Chambers), the only Coptodisca known to attack walnuts, but at present, an unambiguous identification cannot be provided because of the unsatisfactory characterization of this leafminer and congeneric species. Three generations were recorded per year and leafminers overwinter as mature larvae. The first adults emerged in May–June while mature larvae of the last generation started the overwintering in September. During the last generation of the year, infestation levels of leaves were 100% in all sampled localities. Several species of parasitoids were reared from infested mines, with specimens belonging to the genus Chrysocharis (Hymenoptera: Eulophidae) being the most frequent parasitoids.     

Maintenance of the phenology of the winter moth (Lepidoptera: Geometridae)

Adult winter moths ( Operophtera brumata (L.)) are active in late autumn or early winter. The eggs overwinter in the canopy of trees and hatch simultaneously with the bursting of host tree buds. Many young larvae disperse on the wind on silk strands. Larvae are polyphagus and feed until late spring when they pupate in soil or leaf litter. The duration of the egg and pupal stages is genetically determined and varies with latitude. The egg stage is long in the north and short in the south, while the pupal stage is short in the north and long in the south.
The literature on the ecology and physiology of winter moth is reviewed. The factors maintaining the unusual phenology are discussed. It is concluded that the larval stage is early because mature leaves of many host trees are unsuitable as food, because parasitism against later larvae is more intense, and because summer temperatures may be injurious to larvae. The adult period is late in the year so that the final stages of pupal development occur in cool conditions and so that adults emerge after most insect predators have ceased activity. Throughout most of the range retarding the adult emergence period would cause activity to be impeded by severe winter weather; in the south this is not so and it is suggested that eggs must be on the trees for a minimum period to ensure synchronization of egg hatch with bud burst. The protracted adult emergence period may be an adaptation reducing predation by birds.     

Resistance to locust leafminer (Coleoptera: Chrysomelidae) in black locust

Locust leafminer, Odontota dorsalis (Thunberg), activity was observed for 2 yr on 70 black locust, Robinia pseudoacacia L., trees grown from seed collected from its native range. Four black locust seedlings from Tennessee had low numbers of eggs, larvae, pupae, and mines, and were selected for clonal propagation. These clonal propagules were resistant to locust leafminers collected from three different counties in Maryland. This resistance was also observed in controlled choice and no-choice tests in outdoor screen houses. Adults showed different ovipositional preferences for black locust trees from different seed sources, which were reflected in lower numbers of larvae, pupae, and mines in resistant clones. It was concluded that antixenosis and antibiosis were involved in the mechanisms of resistance to the locust leafminer in black locust.     

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.     

Group size effects on survivorship and adult development in the gregarious larvae of Euselasia chrysippe (Lepidoptera, Riodinidae)

Caterpillars living in aggregations may derive several benefits that outweigh the costs, including better survivorship and improved growth rates. I tested whether larval group size had an effect on these two vital rates in Euselasia chrysippe. These caterpillars feed gregariously during all instars and move in processionary form over the host plant and even pupate together. There was a positive relationship between group size and larval survivorship in the field, although genetic variability was not taken into account in this experiment. Under laboratory conditions, there was also a positive relationship between group size, and larval growth rate and adult weight. This supports the hypothesis that aggregations facilitate feeding and larval growth. Single sixth instar larvae in the laboratory also had a lower survivorship than larvae in groups. These results provide further evidence of the benefits of group living for gregarious caterpillars.     

Introduction,establishment, and impact of Lathrolestes thomsoni (Hymenoptera: Ichneumonidae) for biological control of the ambermarked birch leafminer,Profenusa thomsoni (Hymenoptera: Tenthredinidae), in Alaska

The ambermarked birch leafminer, Profenusa thomsoni Konow (Hymenoptera: Tenthredinidae), was first discovered in Alaska in 1991 but was not correctly identified until 1996 when it invaded Anchorage and became a widespread and damaging pest to forest and urban birches. In 2003, the parasitoid wasp, Lathrolestes thomsoni Reshchikov (Hymenoptera: Ichneumonidae), was selected as a candidate for a classical biological control program against P. thomsoni. Parasitized leafminer larvae were collected from the Northwest Territories and Alberta, Canada, where it was previously introduced and causing injury to Betula spp., and transferred to Alaska in soil as pre-pupae for emergence. From 2004 to 2008, 3636 adult L. thomsoni adults were released in birch stands in Alaska. Parasitoids were later recovered at all six release sites in Anchorage and the Kenai Peninsula. While percent leaves mined declined by 30–40% at parasitoid release sites over the period of this study, a similar decline was also observed at paired, non-release long-term monitoring sites. Therefore, the decline in the pest density cannot be ascribed entirely to L. thomsoni.     

美丽青背姬小蜂生物学特性研究

美丽青背姬小蜂Chrysonotomyiaformosa（Westwood）是美洲斑潜蝇的优势天敌,在美洲 斑潜蝇的自然控制中发挥着非常重要的作用。本文对其生物学进行了研究,结果表明：在实验 温度范围内,随着温度的升高,寄生蜂羽化趋早,羽化时间更集中,羽化高峰也更明显;随着 温度的升高,成蜂的寿命逐渐缩短。在提供清水时,寄主可以显著地延长雌蜂的寿命;在有寄 主时,提供10%蜂蜜水,雌蜂的寿命显著延长。美丽青背姬小蜂对3龄寄主幼虫有偏好,对3龄寄 主幼虫的致死率和寄生率都高于对1～2龄寄主幼虫的,且产下后代的雌雄性比为5.11∶1。在 实验温度范围内,发育历期随温度的升高而缩短。