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1.
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.  相似文献   

2.
Residual toxicity and leaf protection capability of five pyrethroids, professional and home garden azadirachtin formulations, and six other bioinsecticides for the home landscape were evaluated against the Japanese beetle, Popillia japonica Newman (Coleoptera: Scarabaeidae), on linden, Tilia cordata L. Capacity of intoxicated beetles to recover and subsequently feed and disperse also was evaluated to provide insight on activity characteristics of the different compounds. Intact shoots were sprayed and left in the field for varying intervals before being challenged with beetles in no-choice and choice tests. All pyrethroids except permethrin gave greater leaf protection, knockdown, and kill than did carbaryl, the standard, after 14 d of weathering. Deltamethrin, cyfluthrin, bifenthrin, and lamda-cyhalothrin gave a high level of protection for at least 19 d, and azadirachtin (Azatin XL) deterred feeding in choice tests for as long as 14 d. Home garden formulations containing pyrethrins in canola oil (Pyola) or azadiractin (Neem-Away) gave good short-term (< 3-d) protection. Formulations of capsaicin, rotenone + pyrethrins, kaolin particle film, D-limonene, or garlic extract were ineffective, the latter two formulations being highly phytotoxic to linden. Results of this study should help support updating of guidelines for insecticidal control of Japanese beetles.  相似文献   

3.
Effectiveness of companion planting, and use of nonhost masking odors were evaluated under field conditions for protecting roses against the Japanese beetle, Popillia japonica Newman. Three reputedly effective companion species, rue (Ruta graveolens L.), zonal geranium (Pelargonium x hortorum Bailey), and garlic chives (Allium scheonparum L.) were interplanted with roses in replicated garden plots. Numbers of beetles on these roses were compared with rose-only control plots on 6 d during beetle flight. The masking odor hypothesis was tested by hanging mesh bags of aromatic herbs or other sources of reputedly repellent nonhost volatiles around potted roses in the field. Treatments included crushed red pepper (Capsicum frutescens L.), fennel seeds (Foeniculm vulgare Miller), crushed spearmint (Mentha picata L.), cedar shavings (Juniperus sp.), osage orange fruits (Maclura pomifera (Raif) Schneid.), and fleshy gingko seeds (Gingko biloba L.). No treatment significantly reduced numbers of beetles relative to the controls. Interplanting with geraniums significantly increased numbers of Japanese beetles on roses. Similarly, roses surrounded by sachets with fennel seeds, cedar shavings, crushed red pepper, or osage orange fruits had significantly more beetles than the control plants on two or more sample dates. Our results suggest that the use of companion or reputedly repellent plants or plant odors probably will be ineffective for protecting roses or other highly-susceptible ornamentals from P. japonica. Use of such tactics in an effort to discourage other garden pests might even increase Japanese beetle damage in those plantings.  相似文献   

4.
The mass gain of polyphagous grasshoppers fed a mixture of intrinsically inferior plants is often greater than that of grasshoppers fed each inferior plant alone. We examined the cause of this greater mass gain by assessing the food processes of a polyphagous grasshopper in detail. Food processes were divided into five stages: consumption, fecal mass, assimilated food, approximate digestibility, and efficiency of conversion of digested food (ECD). Using final-stadium female nymphs of the polyphagous grasshopper Parapodisma subastris, we compared the mass gain of nymphs fed the inferior plants Polygonum cuspidatum and Weigela hortensis or the superior plant Artemisia princeps. The mass gain of nymphs fed a single inferior plant was less than that of nymphs fed the superior plant because of reduced ECD. However, the mass gain of nymphs fed a mixture of the two inferior plants was not only greater than that of nymphs fed a single inferior plant, but also did not differ from that of nymphs fed the superior plant. Since consumption by nymphs fed the mixture was greater than that by nymphs fed P. cuspidatum and smaller than that by nymphs fed W. hortensis, and since the ECD of nymphs fed the mixture was greater than that of nymphs fed W. hortensis and was not different from that of nymphs fed P. cuspidatum, consumption and ECD appeared to cause the greater mass gain in nymphs fed the mixture.  相似文献   

5.
本文比较了龟纹瓢虫Proylea japonica和异色瓢虫Harmonia axyridis取食蚜虫和取食雄蜂蛹粉时的卵黄发生情况。当取食雄蜂蛹粉时,体内卵黄蛋白出现迟,积累速度慢,产卵前期长。但用保幼激素类似物ZR512点滴处理后则能达到与食蚜对照相当的水平。ZR512对取食雄蜂蛹粉瓢虫的作用显著大于取食岈虫者。进一步的研究表明,ZR512能促进这两种瓢虫取食雄蜂蛹粉,但对成虫的体重没有明显的影响。因此推论,雄蜂蛹粉基本能够满足这两种瓢虫生殖的营养需要,但对其内分泌有一定的影响,使瓢虫处于类似生殖滞育的状态。本文根据不同食物条件对卵黄蛋白发生的影响不同,建议用卵黄蛋白的量作为生理指标,以快速初步筛选和评估人工饲料。  相似文献   

6.
Induction of cytochrome P450, glutathione S transferase (GST), and carboxylesterase (CoE) activity was measured in guts of the scarab Popillia japonica Newman, after consumption of single or mixed plant diets of previously ranked preferred (rose, Virginia creeper, crape myrtle and sassafras) or non-preferred hosts (boxelder, riverbirch and red oak). The goal of this study was to quantify activities of P450, GST and CoE enzymes in the midgut of adult P. japonica using multiple substrates in response to host plant suitability (preferred host vs non-preferred hosts), and single and mixed diets. Non-preferred hosts were only sparingly fed upon, and as a group induced higher activities of P450, GST and CoE than did preferred hosts. However, enzyme activities for some individual plant species were similar across categories of host suitability. Similarly, beetles tended to have greater enzyme activities after feeding on a mixture of plants compared to a single plant type, but mixing per se does not seem as important as the species represented in the mix. Induction of detoxification enzymes on non-preferred hosts, or when switching between hosts, may explain, in part, the perceived feeding preferences of this polyphagous insect. The potential consequences of induced enzyme activities on the ecology of adult Japanese beetles are discussed.  相似文献   

7.
Lang A 《Oecologia》2003,134(1):144-153
Arable land typically harbours communities of polyphagous invertebrate natural enemies, among them numerous soil-surface dwelling predators such as ground beetles (Carabidae) and spiders (Lycosidae, Linyphiidae). Numbers of these predators were experimentally manipulated in a winter wheat field in order to study the predation impact of a generalist predator assemblage on herbivorous insects, the possible interferences among the predators concerned, and subsequent effects on wheat plant parameters. Removing ground beetles doubled numbers of Lycosidae indicative of intraguild interference between these two predator groups. Aphid densities were highest in carabid removal plots implying a substantial predation impact of ground beetles on the pest population. The predation impact of ground beetles was strongest earlier and disappeared later in the season. In mid-season, at intermediate aphid densities, the combined impact of carabid beetles and spiders appeared to be responsible for the reduction in aphid abundance. This result was probably due to a biomass effect rather than to a synergistic effect of the predator community. Thysanoptera decreased when spiders were removed (perhaps because spiders were preying on a predator of thrips), while Cicadellidae and Delphacidae showed no effect at all. The rise of aphid numbers in carabid removal plots corresponded to an increase in protein content of the wheat grains, while other plant parameters such as plant numbers and grain mass were not affected. In conclusion, this study provided field evidence for intraguild interference among generalist ground predators in arable land. Despite this interference the polyphagous predator community was able to depress numbers of aphids in winter wheat, a result cascading down to plant quality parameters.  相似文献   

8.
人工饲料对龟纹瓢虫生长和繁殖的影响   总被引:2,自引:0,他引:2  
张丽莉  李恺  张天澍  王斌 《昆虫知识》2007,44(6):871-876
龟纹瓢虫Propylaea japonica(Thunberg)能捕食多种农业害虫,并且具有食量大、产卵多、适应性强、年发生代数多等特点,是我国农业生产中一种重要的捕食性天敌昆虫。对龟纹瓢虫人工饲料的研究则是实现其规模化饲养的关键。作者研究不同饲料对龟纹瓢虫生长和繁殖的影响,结果表明:添加橄榄油后幼虫发育历期明显增长,添加0.3%的橄榄油能显著提高幼虫的成活率;在成虫体重增长上,雌雄虫均表现为在刚羽化后几天内体重增长显著,随后增长变缓;并且添加蔗糖能显著提高雌虫的体重增长率而对雄虫作用不显著;对雌虫繁殖性能的研究发现,蚜虫喂养组有明显的持续时间较长的产卵高峰期,而人工饲料的产卵高峰期持续时间较短,并且出现时间较晚或分为2个高峰期;添加0.3%的橄榄油能显著缩短龟纹瓢虫的产卵前期,几乎达到蚜虫喂养组的水平;而添加蔗糖在一定程度也能缩短产卵前期,同时还能显著提高龟纹瓢虫的产卵量。  相似文献   

9.
Nearby fruit and vegetable fields in Islamabad, Pakistan were surveyed for phytoplasma infection. ' Candidatus Phytoplasma asteris' (Group 16SrI) was found infecting mango, citrus, loquat, geranium, periwinkle, radish, blackberry and potato. Results suggest that a polyphagous vector may be involved in phytoplasma transmission to these plant species, which are first host records of 16SrI phytoplasma infection in Pakistan.  相似文献   

10.
DNA barcoding facilitates many evolutionary and ecological studies, including the examination of the dietary diversity of herbivores. In this study, we present a survey of ecological associations between herbivorous beetles and host plants from seriously threatened European steppic grasslands. We determined host plants for the majority (65%) of steppic leaf beetles (55 species) and weevils (59) known from central Europe using two barcodes (trnL and rbcL) and two sequencing strategies (Sanger for mono/oligophagous species and Illumina for polyphagous taxa). To better understand the ecological associations between steppic beetles and their host plants, we tested the hypothesis that leaf beetles and weevils differ in food selection as a result of their phylogenetic relations (within genera and between families) and interactions with host plants. We found 224 links between the beetles and the plants. Beetles belonging to seven genera feed on the same or related plants. Their preferences were probably inherited from common ancestors and/or resulted from the host plant's chemistry. Beetles from four genera feed on different plants, possibly reducing intrageneric competition and possibly due to an adaptation to different plant chemical defences. We found significant correlations between the numbers of leaf beetle and weevil species feeding on particular plants for polyphagous taxa, but not for nonpolyphagous beetles. Finally, we found that the previous identifications of host plants based on direct observations are generally concordant with host plant barcoding from insect gut. Our results expand basic knowledge about the trophic relations of steppic beetles and plants and are immediately useful for conservation purposes.  相似文献   

11.
1. The food plant quality influences feeding preferences and various life history traits of herbivorous insects. However, the effects of different host plant qualities on the behavioural phenotype have rarely been studied in behavioural ecology, especially in a pest‐crop‐framework. 2. Behavioural phenotypes of insects may not only be affected by external environmental factors, such as the host plant quality but are also shaped by internal factors, such as the sex and the age of individuals. 3. To study host plant effects on behavioural phenotypes, we reared mustard leaf beetles (Phaedon cochleariae Fabricius) either on their natural host watercress or on the crop cabbage, on which this beetle can be a pest. The behavioural phenotype was characterised twice in the adult lifetime by measuring six behavioural traits tested in distinct contexts. 4. Depending on the context, different behavioural traits were specifically affected by the host plant, the sex and/or the age. Beetles fed on cabbage became more active with age. Furthermore, the boldness tested in an unprotected environment context was influenced by the host, with beetles fed on cabbage being bolder, whereas the boldness in a hiding or predator attack context was affected by the age and/or the interaction of host plant × sex. 5. In conclusion, beetles fed on the crop cabbage develop a different behavioural phenotype compared to beetles fed on watercress. Previous results showed that beetles reared on cabbage have a higher reproductive output. Thus, beetles fed on the crop potentially express a faster pace‐of‐life.  相似文献   

12.
Abstract: Plants have developed defensive mechanisms to minimize predation by insect pests. Proteinase inhibitors are an example of plant compounds synthesized as a mechanism for defence. The objective of this study was to determine the impact of trans‐epoxysuccinyl‐l ‐leucylamido (4‐guanidino) butane (E‐64), phenylmethylsulfonyl fluoride (PMSF‐serine protenase inhibitor) and Kunitz trypsin inhibitors on the pre‐ovipositional and ovipositional periods, the mean number of eggs laid per female, and the longevity of western corn rootworm (WCR), Diabrotica virgifera virgifera LeConte, adults. This study provides information on the effectiveness of proteinase inhibitors as a host‐plant resistance tool for managing WCR beetles. The study was conducted in 1997, 1998, and 2000. In 1997, E‐64 was added to an artificial diet at the concentrations of 0.05, 0.025 and 0.0125% (w/w), corresponding to 500, 250, and 125 ppm respectively. In 1998, PMSF was added to the artificial diet at the same concentrations. In 2000, Kunitz trypsin inhibitor was added to the artificial diet at concentrations of 0.2, 0.1 and 0.05% (w/w), corresponding to 2000, 1000, and 500 ppm respectively. The mean fecundity of beetles fed the untreated diet was between 67 and 111 eggs per female. The fecundity of beetles fed E‐64 and PMSF at different concentrations, ranged between 162 and 246 eggs per female for E‐64 and 61 and 80.5 eggs per female for PMSF. The fecundity of the beetles fed Kunitz trypsin inhibitor was between 155 and 225 eggs per female. When beetles fed on the diet which consisted of the lowest dosage of Kunitz trypsin inhibitor (500 ppm), fecundity was higher than that on untreated control. The proteinase inhibitors investigated did not show a negative impact on WCR adults. Beetle fecundity, the length of the pre‐ovipositional and ovipositional periods and the longevity of the beetles fed with proteinase inhibitors were not lower than that of the beetles fed only the artificial diet. This study does not support the use of investigated proteinase inhibitors at applied concentrations as effective host‐plant resistance tools for managing WCR beetles.  相似文献   

13.
研究了取食转Bt-cry1Ah基因玉米花粉对龟纹瓢虫Propylaea japonica(Thunberg)体内解毒酶和中肠蛋白酶活性的影响。利用饲喂结合比色方法,比较龟纹瓢虫取食转Bt-cry1Ah基因玉米花粉和非转基因玉米花粉后体内α-乙酸萘酯酶、乙酰胆碱酯酶、谷胱甘肽-S-转移酶、中肠总蛋白酶、类胰蛋白酶和类胰凝乳蛋白酶的酶活性。结果发现:在解毒酶方面,取食Bt玉米花粉的龟纹瓢虫4龄幼虫和蛹的α-乙酸萘酯酶活性显著低于取食非Bt玉米花粉的龟纹瓢虫(对照组),取食Bt玉米花粉的龟纹瓢虫的乙酰胆碱酯酶和谷胱甘肽-S-转移酶活性在各个发育时期与对照相比均无显著差异。在中肠蛋白酶方面,与对照组相比,取食Bt玉米花粉的龟纹瓢虫的总蛋白酶和强碱性类胰蛋白酶活性在各个发育时期均无显著差异;但取食Bt玉米花粉的龟纹瓢虫的弱碱性类胰凝蛋白酶和类胰凝乳蛋白酶活性在蛹期显著低于取食非Bt玉米花粉的龟纹瓢虫。由此可见,龟纹瓢虫取食含有Cry1Ah杀虫蛋白的玉米花粉后,体内代谢解毒酶和中肠蛋白酶与Cry1Ah杀虫蛋白相互作用,可能会引起某些酶活性的变化。因此,转cry1Ah基因玉米花粉对龟纹瓢虫的潜在影响还需要进一步的研究。  相似文献   

14.
As fundamentally different as phytopathogenic microbes and herbivorous insects are, they enjoy plant‐based diets. Hence, they encounter similar challenges to acquire nutrients. Both microbes and beetles possess polygalacturonases (PGs) that hydrolyze the plant cell wall polysaccharide pectin. Countering these threats, plant proteins inhibit PGs of microbes, thereby lowering their infection rate. Whether PG‐inhibiting proteins (PGIPs) play a role in defense against herbivorous beetles is unknown. To investigate the significance of PGIPs in insect–plant interactions, feeding assays with the leaf beetle Phaedon cochleariae on Arabidopsis thaliana pgip mutants were performed. Fitness was increased when larvae were fed on mutant plants compared to wild‐type plants. Moreover, PG activity was higher, although PG genes were downregulated in larvae fed on PGIP‐deficient plants, strongly suggesting that PGIPs impair PG activity. As low PG activity resulted in delayed larval growth, our data provide the first in vivo correlative evidence that PGIPs act as defense against insects.  相似文献   

15.
Production of sweetpotatoes, Ipomoea batatas (L.) Lam. (Convolvulaceae), is limited by several insect pests, including Diabrotica spp. (Coleoptera: Chrysomelidae), and new integrated pest management (IPM) techniques for this crop are needed. Host plant resistance is one attractive approach that fits well into IPM programs. A host plant resistance research program typically depends on reliable bioassay procedures to streamline evaluation of germplasm. Thus, a bioassay technique was developed for evaluating sweetpotato germplasm by using adults of the banded cucumber beetle, Diabrotica balteata LeConte, and spotted cucumber beetle, Diabrotica undecimpunctata howardi Barber. A single beetle was placed on a piece of sweetpotato peel (periderm and cortex with stele removed) that was embedded periderm-side up in plaster in a petri dish. Feeding and longevity of insects on 30 sweetpotato genotypes were evaluated in two experiments by using this procedure. Adult longevity ranged from 7 to 11 d for starved individuals to 211 d for beetles fed a dry artificial diet. Longevity of banded cucumber beetles that fed on sweetpotato peels ranged from 12 d for the most-resistant genotype to 123 d for SC1149-19, a susceptible control cultivar. Longevity of spotted cucumber beetles was slightly shorter than longevity of banded cucumber beetles. For the most resistant sweetpotato genotypes, both Diabrotica species exhibited a significant delay in initiation of feeding, and more beetles died on these genotypes before they had fed. Both antibiosis and nonpreference (antixenosis) are important mechanisms of resistance in sweetpotato genotypes. This bioassay was consistent with field results, indicating that this technique could be useful for evaluating resistance to Diabrotica spp. in sweetpotato genotypes.  相似文献   

16.
The flight response of Cotesia kariyaiWatanabe (Hymenoptera: Braconidae), a parasitoid of the polyphagous herbivore, Mythimna separataWalker (Lepidoptera: Noctuidae), to pairs of different plant species infested by M. separatalarvae was tested under a dual choice condition in the laboratory. The oviposition-inexperienced (naive) wasps showed preference in the order: corn > kidney bean > Japanese radish. Wasps that had previously oviposited on the less preferred plant in a pair were found to have shifted their preference to this plant at 2 h after oviposition. However, this shift became indistinct at 17 h after oviposition. Prior oviposition on a plant species other than those being compared also affected the preference. These data suggest that learning is involved in the wasp's flight response. Prior oviposition was also observed to have an effect on the antennal searching behavior of the wasp on corn leaves. Such behavioral plasticity may enhance the efficiency by which C. kariyaisearches for polyphagous hosts.  相似文献   

17.
The relative impact of cannibalism and predation on the development and survival of fourth instar larvae was assessed in two species of aphidophagous ladybird beetles, Coccinella septempunctata and Harmonia axyridis. The effect of eating aphids, conspecific larvae or heterospecific larvae on larval performance differed in the two species: aphids were the best food for C. septempunctata and survival of C. septempunctata larvae was significantly lower when offered heterospecific larvae rather than conspecific larvae or aphids as food, indicating that H. axyridis larvae were not suitable food for C. septempunctata. However, as the different foods did not affect the larval performance of H. axyridis, this species appears to be more polyphagous. Both intraguild predation by the aggressive larvae of H. axyridis and the polyphagous food habit of this species may account for its dominance in ladybird assemblages and its displacing other ladybird beetles in several places in the world.  相似文献   

18.
The effects of three cotton cultivars with low ('ZMZ13'), medium ('HZ401'), and high ('M9101') gossypol contents on the development, reproduction, and survival of Aphis gossypii Glover and its predator Propylaea japonica (Thunberg) were investigated. Developmental duration and immature survivorship did not vary between aphids on the three cultivars, whereas A. gossypii feeding on M9101 (high gossypol cultivar) displayed significantly shorter adult longevity and lower fecundity than aphids fed on 'ZMS13' and 'HZ401'. Free fatty acid content in cotton aphids reared on 'M9101' was greater than in those reared on 'HZ401' and 'ZMS13'. No significant differences in survival and lifetime fecundity of P. japonica were observed between P. japonica fed cotton aphids reared on the three different cultivars. P. japonica fed aphids from 'M9101' showed a significantly shorter developmental period and greater adult weight than those fed aphids from the other two cultivars. The decreased larval developmental duration and increased adult weight of P. japonica fed cotton aphids reared on the high gossypol-containing cultivar might have been caused by the high fatty acid content of the prey aphids. Our results indicate that high gossypol in host cotton had an antibiotic effect on A. gossypii and showed a positive effect on growth and development of P. japonica at the third trophic level. This suggests compatibility between one form of host plant resistance and biological control by predators. The allelochemical contents should be taken into account in integrated pest management for their effects on both herbivores and entomophagous insects.  相似文献   

19.
The Phytophagous Insects Data Base (PIDB) was used to summarize information about 6933 species of British insects/mites and their food-plant families. Total species in 183 insect families on 127 vascular plant families were correlated with numbers of plant species in each family in Britain and Europe, accounting for 41.8% of the variation. Families with trees had more insects, while ferns and aquatic, uncommon and ephemeral plants, particularly orchids, had fewer. Seven individual insect families were analysed separately: Aphididae were more closely correlated with plant species numbers than were the more polyphagous Geometridae. Agromyzidae were uncommon on families with trees. Of insect species 75.8% fed on only one plant family and 10.1% were recorded on two families. Species feeding on three families or more were considered to be polyphagous. In the 30 largest insect families, eight had species feeding on only one plant family, while 12 had more than 20% of their species polyphagous. Proportions of family-restricted insects were high on Pinaceae, Gramineae and Compositae, but low on Corylaceae, Primulaceae and Tiliaceae. More pest species were polyphagous. Polyphagy was related to large larvae, larvae overwintering, wood-feeding, ephemeral food sources, poor dispersal abilities and omnivory. Some polyphagous species might be actively evolving in exploitation of vacant niches, leading to later progressive specialization to food-plants.  相似文献   

20.
Host plant cues are known to shape insect–host plant association in many insect groups. More pronounced associations are generally manifested in specialist herbivores, but little is known in generalist herbivores. We used a polyphagous native beetle from New Zealand, bronze beetle, Eucolaspis sp. ‘Hawkes Bay’ (Chrysomelidae: Eumolpinae) to explore the role of olfaction in locating host plants and local adaptation. We also tested the role of other cues in the degree of acceptance or rejection of hosts. Adult Eucolaspis beetles were attracted to fresh leaf volatiles from apple and blackberry (Rosaceae). Male and female beetles responded similarly to olfactory cues of host plants. An indication of evolutionary affiliation was observed in olfactory preferences of geographically isolated conspecific populations. We found that geographically isolated populations of the beetles differ in their olfactory responses and exhibit some degree of local adaptation. However, irrespective of geographical and ecological associations, blackberry was preferred over apple as a feeding plant, and another novel plant, bush lawyer (Rubus australis), was readily accepted by 53.25% of the tested beetles. We show that plant volatiles play an important role in host location by Eucolaspis, but the acceptance or rejection of a particular host could also involve visual and contact cues.  相似文献   

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