QUANTIFYING PREDATION BY UMMELIATA INSEC-TICEPS BOES. ET STR. (ARANEAE: LINYPHIIDAE) ON RICE PLANTHOPPERS USING ELISA*

Abstract An enzyme-linked immunosorbent assay (ELISA) has been developed to quantify predation by Ummeliata insecticeps Boes. et Str. on rice planthoppers in paddy fields in Dasha Township, Guangdong Province. The assay was completely specific for rice planthopper materials. The detection periods for antigens after an adult of U. insecticeps had fed on three 3–5th instar nymphs of white-back planthopper (WBPH), Sogatella furcifera (Horvath), and brown planthopper (BPH), Nilaparvata lugens (Stal), at room temperature were 96 h and 120 h, respectively. The proportion of individual species of predators scoring positive ranged from 19. 05% to 47. 34% for WBPH, and from 9. 25% to 66. 67% for BPH in the early rice season. Although the numbers of the rice planthoppers consumed by U. insecticeps increased with increasing prey densities, the predation rates declined. When rice planthopper densities were low, the predation rates were relatively high. This kind of predation can explain why the rice planthoppers have never reached outbreak levels in rice fields in Dasha Township since 1973.     

Impacts of nitrogen fertilizer on major insect pests and their predators in transgenic Bt rice lines T2A‐1 and T1C‐19

Nitrogen is a critical factor for plant development and nitrogen input is one of the important tactics to enhance the development and yield of crops. Nevertheless, nitrogen input could influence the occurrence of insects positively or negatively. Nitrogen is also one of the main elements composing the insecticidal crystal (Cry) protein. Cry protein production could affect nitrogen partitioning in Bt plants and as such nitrogen input may influence insect pest management in transgenic Bt rice, Oryza sativa L. (Poaceae). To test this possibility, we evaluated the impacts of nitrogen regimes on the main insect pests and their predators on two Bt rice lines, T2A‐1 and T1C‐19, expressing Cry2A and Cry1C, respectively, and their non‐transgenic parental counterpart MH63. The results showed that Cry proteins with different nitrogen regimes have enough insecticidal activity on rice leaffolder, Cnaphalocrocis medinalis Guenée (Lepidoptera: Crambidae), in both laboratory and field experiments. Laboratory studies indicated that relevant parameters of ecological fitness in brown planthopper, Nilaparvata lugens (Stål) (Hemiptera: Delphacidae), a non‐target insect pest, were significantly affected by nitrogen input both on Bt and MH63 rice lines. Nymphal survival, female adult longevity, and egg hatchability in N. lugens differed significantly among rice varieties. The experiments conducted in rice fields also demonstrated that nitrogen was positively correlated with the abundance of N. lugens on Bt rice, similar to that on MH63 rice. The abundances of two predators – the wolf spider Pirata subpiraticus (Boesenberg & Strand) (Araneae: Lycosidae) and the bug Cyrthorhinus lividipennis Reuter (Hemiptera: Miridae) – were significantly affected by rice growth stages but not by nitrogen input and rice varieties. In conclusion, the above results indicate that high nitrogen regimes for Bt rice (T2A‐1 and T1C‐19) and non‐Bt rice (MH63) cannot facilitate the management of insect pests.     

用ELISA方法定量评价食虫沟瘤蛛对稻飞虱的捕食作用(英文)

对捕食性节肢动物的捕食作用进行评价是害虫生物防治研究的一个重要内容。本文利用酶联免疫吸附试验（ＥＬＩＳＡ）定量评价了食虫沟瘤蛛对稻飞虱的捕食作用。结果表明,方法特异性完全符合试验要求;食虫沟瘤蛛在捕食３头３－５龄白背飞虱或褐飞虱若虫后的检出期分别为９６小时和１２０小时;在早稻田中,捕食性天敌对白背飞虱和褐飞虱的阳性反应率分别为１９．０５％－４７．３４％和１９．０５％－６６．６７％,在飞虱密度较低时,捕食性天敌仍表现出较高的阳性率;捕食量随飞虱密度的增加而增加,但捕食率下降。捕食性节肢动物是调节褐飞虱种群动态的重要因子之一。     

A false‐positive food chain error associated with a generic predator gut content ELISA

Conventional prey‐specific gut content ELISA (enzyme‐linked immunosorbent assay) and PCR (polymerase chain reaction) assays are useful for identifying predators of insect pests in nature. However, these assays are prone to yielding certain types of food chain errors. For instance, it is possible that prey remains can pass through the food chain as the result of a secondary predator (hyperpredator) consuming a primary predator that had previously consumed the pest. If so, the pest‐specific assay will falsely identify the secondary predator as the organism providing the biological control services to the ecosystem. Recently, a generic gut content ELISA was designed to detect protein‐marked prey remains. That assay proved to be less costly, more versatile, and more reliable at detecting primary predation events than a prey‐specific PCR assay. This study examines the chances of obtaining a ‘false positive’ food chain error with the generic ELISA. Data revealed that the ELISA was 100% accurate at detecting protein‐marked Lygus hesperus Knight (Hemiptera: Miridae) remains in the guts of two (true) primary predators, Hippodamia convergens Guérin‐Méneville (Coleoptera: Coccinellidae) and Collops vittatus (Say) (Coleoptera: Melyridae). However, there was also a high frequency (70%) false positives associated with hyperpredators, Zelus renardii Kolenati (Hemiptera: Reduviidae), that consumed a primary predator that possessed protein‐marked L. hesperus in its gut. These findings serve to alert researchers that the generic ELISA, like the PCR assay, is susceptible to food chain errors.     

Similarities and spatial variations of bacterial and fungal communities in field rice planthopper (Hemiptera: Delphacidae) populations

Rice planthoppers are notorious plant sap‐feeding pests which cause serious damage. While several microbes in rice planthoppers have been broadly characterized, the abundance and diversity of bacteria and fungi in field planthoppers are largely unknown. This study investigated the bacterial and fungal community compositions of Chinese wild rice planthoppers Laodelphax striatellus and Sogatella furcifera using parallel 16S rRNA gene amplicon and internal transcribed space region sequencing. The bacteria varied significantly between the species and were partitioned significantly by sex, tissues and host environments in each species. The majority of bacteria were affiliated with the genera Wolbachia, Cardinium, Rickettsia and Pantoea. The abundance of Wolbachia was negatively correlated with that of Cardinium in both planthopper species. Compared with bacteria, the abundance and diversity of fungi did not differ between sexes but both were enriched in the gut. The bacterial community as a whole showed no significant correlation with the fungal community. The majority of fungi were related to Sarocladium, Alternaria, Malassezia, Aspergillus and Curvularia. A phylogenetic analysis revealed that these fungi were closely related to botanic symbionts or pathogens. Our results provide novel insights into the bacteria and fungi of rice planthoppers.     

Transgenic Cry1Ab rice does not impact ecological fitness and predation of a generalist spider

Background

The commercial release of rice genetically engineered to express a Cry1Ab protein from Bacillus thuringiensis (Bt) for control of Lepidoptera in China is a subject of debate. One major point of the debate has focused on the ecological safety of Bt rice on nontarget organisms, especially predators and parasitoids that help control populations of insect pests.

Methodology/Principal Findings

A tritrophic bioassay was conducted to evaluate the potential impact of Cry1Ab-expressing rice on fitness parameters of a predaceous ground spider (Pardosa pseudoannulata (Bösenberg et Strand)) that had fed on Bt rice-fed brown planthopper (Nilaparvata lugens (Stål)) nymphs. Survival, development time and fecundity of this spider were not different when they were fed with Bt rice-fed or non-Bt rice-fed prey. Furthermore, ELISA and PCR gut assays, as well as a functional response trial, indicated that predation by P. pseudoannulata was not significantly different in Bt rice or non-Bt rice fields.

Conclusions/Significance

The transgenic Cry1Ab rice lines tested in this study had no adverse effects on the survival, developmental time and fecundity of P. pseudoannulata in the laboratory or on predation under field conditions. This suggests that this important predator would not be harmed if transgenic Cry1Ab rice were commercialized.     

Planthopper‐rice interactions: unequal stresses on pure‐line and hybrid rice under similar experimental conditions

Hybrid and pure‐line (inbred) rice [Oryza sativa L. (Poaceae)] varieties have distinct physiologies, particularly as related to their nutrient requirements. These differences could confound the results and interpretation of experiments that compare rice varieties grown in pots for their resistance and responses to herbivores. In this study, a series of experiments was conducted to identify potentially confounding interactions between pot size (soil volume), fertilizer regime, and the use of acetate insect cages with rice line type (hybrid, fertile parental inbred, and male sterile inbred) during bioassays with the brown planthopper, Nilaparvata lugens (Stål) (Hemiptera: Delphacidae). The growth of hybrid rice was often limited (relatively low biomass and low grain production) compared to fertile inbred lines even in large pots (7 200 ml) and when grown without added fertilizer. Several interactions between the effects of growth conditions and line type were detected. Acetate cages caused a significant reduction in grain yield in hybrids, but not in inbreds, mainly resulting from a cage‐induced decrease in grain weight (smaller grains). Hybrids and male sterile lines often had higher root or above‐ground biomass in the largest caged pots compared with fertile inbred lines, but biomass was similar in smaller pots, indicating that the large pots allowed longer unimpeded growth of fertile inbreds, but not other line types. There were no interactions between the presence or absence of planthoppers with line type or experimental conditions. All line types were equally susceptible to planthoppers. Often the effects of planthopper feeding on plant fitness (i.e., tolerance) were apparent when plants were grown in large pots but not in small pots, particularly for hybrid lines and under high nitrogen regimes. On the basis of these results we recommend that researchers ensure that plants are not unequally stressed by inadequate growth conditions during comparative studies with herbivores on physiologically distinct rice varieties or rice species. We recommend the use of large pots (soil volume) and lower fertilizer levels with young, non‐reproductive plants during comparative bioassays with planthoppers. Field cages are recommended for hybrid‐inbred comparisons during older plant stages but these are subject to a range of other confounding variables.     

Electrical penetration graphs indicate that tricin is a key secondary metabolite of rice,inhibiting phloem feeding of brown planthopper,Nilaparvata lugens

Tricin (5,7,4′‐trihydroxy‐3′,5′‐dimethoxyflavone) is a valuable secondary metabolite which is widely present in gramineous plants, including cultivated rice (Oryza sativa L.) (Poaceae). It can defend the rice plant against damage by the brown planthopper (BPH), Nilaparvata lugens (Stål) (Hemiptera: Delphacidae), one of the most important pests of rice. This study was conducted to elucidate the mechanisms of action of tricin on BPH feeding behavior. BPH feeding behavior in resistant (Rathu Heenati, RHT) and susceptible (Taichuang native 1, TN1) rice varieties and artificial diets was monitored using the electrical penetration graph (EPG) technique. Tricin concentrations in leaves of varieties RHT and TN1 were quantitatively analyzed by liquid chromatography, coupled to tandem mass spectrometric techniques. Six (NP and N1‐5) and four (NP, N1, N2, and N4) types of waveforms occurred during feeding on rice plants and artificial diets, respectively. The tricin concentration of rice varieties was correlated with total and average durations of N4. Moreover, EPG data indicated that tricin significantly increased the duration of non‐probing and pathway periods and strongly inhibited phloem ingestion (N4). The inhibition was strongly dose dependent, resulting in complete suppression of activity in the phloem region when the tricin concentration was increased to 1 g l^?1. This study revealed that tricin disturbed the feeding behavior of BPH mainly by increasing the non‐probe period and inhibiting phloem ingestion. We confirmed the hypothesis that tricin is a ‘stylet probing stimulant’ of rice planthoppers as proposed in previous studies. The information on the ecological effect of tricin from this study may be useful to clarify the resistance mechanism against BPH of RHT and other tricin‐containing rice varieties.     

用ElISA研究稻田节肢类捕食者对稻飞虱的捕食作用

应用酶联免疫吸附试验(ELlSA)来研究捕食作用能较准确地评价捕食性天敌对害虫的控制作用。ELlSA检测表明,不同种类的捕食性天敌对稻飞虱的捕食作用有较大差别,其中以优势种食虫沟瘤蛛和拟水狼蛛的捕食作用最大,阳性反应率与稻飞虱的种群密度有一定的相关关系,但在稻飞虱密度较低的情况下也表现出较高的阳性反应率。     

Chromosomal‐level genomes of three rice planthoppers provide new insights into sex chromosome evolution

The brown planthopper Nilaparvata lugens, white‐backed planthopper Sogatella furcifera, and small brown planthopper Laodelphax striatellus are three major insect pests of rice. They are genetically close; however, they differ in several ecological traits such as host range, migration capacity, and in their sex chromosomes. Though the draft genome of these three planthoppers have been previously released, the quality of genome assemblies need to be improved. The absence of chromosome‐level genome resources has hindered in‐depth research of these three species. Here, we performed a de novo genome assembly for N. lugens to increase its genome assembly quality with PacBio and Illumina platforms, increasing the contig N50 to 589.46 Kb. Then, with the new N. lugens genome and previously reported S. furcifera and L. striatellus genome assemblies, we generated chromosome‐level scaffold assemblies of these three planthopper species using HiC scaffolding technique. The scaffold N50s significantly increased to 77.63 Mb, 43.36 Mb and 29.24 Mb for N. lugens, S. furcifera and L. striatellus, respectively. To identify sex chromosomes of these three planthopper species, we carried out genome re‐sequencing of males and females and successfully determined the X and Y chromosomes for N. lugens, and X chromosome for S. furcifera and L. striatellus. The gene content of the sex chromosomes showed high diversity among these three planthoppers suggesting the rapid evolution of sex‐linked genes, and all chromosomes showed high synteny. The chromosome‐level genome assemblies of three planthoppers would provide a valuable resource for a broad range of future research in molecular ecology, and subsequently benefits development of modern pest control strategies.     

Interactions between a hunting spider and a web-builder: consequences of intraguild predation and cannibalism for prey suppression

Abstract. 1. Antagonistic interactions among invertebrate predators such as intraguild predation and cannibalism have the potential to dampen top‐down impacts on shared prey at lower trophic levels. Two abundant spider predators, the large wolf spider Pardosa littoralis and the small sheet‐web builder Grammonota trivitatta co‐occur on the salt marshes of eastern North America where they both attack planthoppers (Prokelisia spp.), the dominant herbivores on the marsh. Experiments both in the laboratory and field were used to assess the incidence of intraguild predation and cannibalism in these spiders and elucidate how such antagonistic interactions influence planthopper suppression. 2. Functional response experiments showed that with an increase in planthopper prey density, Grammonota captured more prey but not a higher proportion of that offered. Pardosa exhibited the same response when Grammonota were offered as intraguild prey. Both functional responses were type I over the range of prey densities offered. 3. Grammonota is moderately cannibalistic, and the presence of planthopper prey reduced the incidence of cannibalism. 4. Factorial experiments in the laboratory showed that Pardosa but not Grammonota reduced planthopper prey populations when prey density was low. By contrast, at high prey densities, both Pardosa and Grammonota had significant adverse effects on planthopper populations. Moreover, there was an interactive effect such that Grammonota reduced planthopper populations relatively more when Pardosa was absent than when it was present. 5. There was direct evidence for the intraguild predation of Grammonota by Pardosa such that fewer Grammonota survived in the presence of Pardosa than when it was absent. This result occurred whether planthopper prey were abundant or not. 6. Field releases of Grammonota in open plots resulted in significant but small decreases in the density of planthopper prey, both nymphs and adults. 7. Enhancing densities of Pardosa in open plots resulted in Grammonota suppression. The intraguild predation of Grammonota at this enhanced Pardosa density, however, did not preclude Pardosa from significantly reducing planthopper populations. 8. Although there was evidence that Grammonota reduced planthopper populations and that the intraguild predation of Grammonota by Pardosa occurred, the strength of these interactions was relatively weak given the low consumption rate of planthoppers by Grammonota (< 3 day^–1) and Grammonota by Pardosa (≈ 2 day^?1). Thus, weak asymmetric intraguild predation among spiders on the marsh likely dampens but does not eliminate the ability of Pardosa to exert significant top‐down control on planthopper populations.     

Genetic and biochemical mechanisms of rice resistance to planthopper

Key message

This article presents a comprehensive review on the genetic and biochemicalmechanisms governing rice-planthopper interactions, aiming to contribute substantialplanthopper control and facilitate breeding for resistance to planthoppers in rice.

Abstract

The rice planthopper is the most destructive pest of rice and a substantial threat to rice production. The brown planthopper (BPH), white-backed planthopper (WBPH) and small brown planthopper (SBPH) are three species of delphacid planthoppers and important piercing-sucking pests of rice. Host-plant resistance has been recognized as the most practical, economical and environmentally friendly strategy to control planthoppers. Until now, at least 30, 14 and 34 major genes/quantitative trait loci for resistance to BPH, WBPH and SBPH have been identified, respectively. Recent inheritance and molecular mapping of gene analysis showed that some planthopper-resistance genes in rice derived from different donors aggregate in clusters, while resistance to these three species of planthoppers in a single donor is governed not by any one dominant gene but by multiple genes. Notably, Bph14, Bph26, Bph3 and Bph29 were successfully identified as BPH-resistance genes in rice. Biological and chemical studies on the feeding of planthoppers indicate that rice plants have acquired various forms of defence against planthoppers. Between the rice-planthopper interactions, rice plants defend against planthoppers through activation the salicylic acid-dependent systemic acquired resistance but not jasmonate-dependent hormone response pathways. Transgenic rice for the planthopper-resistance mechanism shows that jasmonate and its metabolites function diversely in rice’s resistance to planthopper. Understanding the genetic and biochemical mechanisms underlying resistance in rice will contribute to the substantial control of such pests and facilitate breeding for rice’s resistance to planthopper more efficiently.

     

Intraspecific Variation in Chemical Attraction of Rice to Insect Predators

The olfactory response of predators of the brown planthopper,Nilaparvata lugensStål, to different genotypes of rice (14 cultivars and breeding lines ofOryza sativaL. and 1 wild species,Oryza nivaraSharma et Shastry) was measured in an airflow olfactometer. Odor from rice plants attracted more females of the mirid predatorCyrtorhinus lividipennisReuter than plain air (control) on only 6 of the 15 rice genotypes. Orientation ofC. lividipennistoward volatiles of certain rice genotypes was apparent even when the plants were free of the brown planthopper. However, the predator distinguished between prey-infested and uninfested plants and preferred plants with eggs over plants with nymphs. The predator did not distinguish different stages of plant growth (vegetative, booting, or flowering). Plants artificially injured to simulate brown planthopper oviposition wounds were not as attractive to the predator as plants on which the planthopper had oviposited. The preassay preconditioning on the cultivar TN1 did not produce a predator bias for this genotype. This suggests that rearing effects or chemically mediated associative learning reported for some natural enemies did not influenceC. lividipennis'host response. Results with another predator, the coccinellidMicraspis hirashimaiSasaji, produced less consistent behavior. Planthopper-infested plants attracted more females ofM. hirashimaithan unifested plants in only 1 of the 12 rice genotypes evaluated. Implications for augmenting predators by rice cultivar selection and modification are discussed.     

Development of a TaqMan Real‐Time Polymerase Chain Reaction Assay for Detection and Quantification of Fusarium oxysporum f. sp. lycopersici in Soil

Fusarium wilt, caused by Fusarium oxysporum f. sp. lycopersici (FOL), is an important disease of tomato. Pathogenicity and vegetative compatibility tests, although reliable, are laborious for the identification of FOL isolates and cannot efficiently quantify population densities of FOL in the soil. The objective of this study was to develop a rapid, sensitive and quantitative real‐time polymerase chain reaction (PCR) assay for detecting and quantifying FOL in soil. An inexpensive and relatively simple method for soil DNA extraction and purification was developed based on bead‐beating and a silica‐based DNA‐binding method. A TaqMan probe and PCR primers were designed using the DNA sequence of the species‐specific virulence gene SIX1, which is only present in isolates of FOL, not in isolates of other formae speciales or non‐pathogenic isolates of F. oxysporum. The real‐time PCR assay successfully amplified isolates of three races of FOL used in this study and quantified FOL DNA in soils, with a detection limit of 0.44 pg of genomic DNA of FOL in 20 μl of the real‐time PCR. A spiking test performed by adding different concentrations of conidia to soil showed a significant linear relationship between the amount of genomic DNA of FOL detected by the real‐time PCR assay and the concentration of conidia added. In addition, the real‐time PCR assay revealed a significant quadratic regression for a glasshouse experiment between disease severity and DNA concentration of FOL. The soil DNA extraction method and real‐time PCR assay developed in this study could be used to determine population densities of FOL in soil, develop threshold models to predict Fusarium wilt severity, identify high‐risk fields and measure the impact of cultural practices on FOL populations in soils.     

An analysis of the predator-prey system in the paddy field

A predator-prey system involving Lycosa pseudoannulata as predators and both the green rice leafhopper, Nephotettix cincticeps and the brown planthopper, Nilaparvata lugens as prey was studied in an experimental paddy field. Daily predation rates on the prey and weekly disappearance rates of Lycosa females were related to densities of both predators and prey by multiple regression. During the off-season of reproduction for predators and prey, it proved possible to simulate the interaction between Lycosa and its prey.     

稻田穗期主要捕食性天敌对两种盲蝽集团内捕食的初步研究

【目的】集团内捕食是影响农业系统广食性天敌对靶标害虫控制作用的重要因素,全面揭示稻田广食性天敌对稻飞虱重要天敌黑肩绿盲蝽Cyrtorhinus lividipennis Reuter和中华淡翅盲蝽Tytthus chinensis(St?l)(半翅目:盲蝽科)的集团内捕食作用有助于更好地保护利用天敌。【方法】建立以两种盲蝽为猎物对象的种特异性二重定量PCR系统,检测比较不同抗性品种穗期田间主要捕食者对两种盲蝽的集团内捕食作用。【结果】(1)本研究建立的二重定量PCR具有的物种专化性,两种检测靶标灵敏度高且相似,可用于田间捕食者猎物分析;(2)感虫水稻品种上捕食者对盲蝽的捕食强度高于抗性水稻品种;不同种类捕食者对盲蝽的集团内捕食强度有显著差异;捕食者对黑肩绿盲蝽集团内捕食强度显著高于中华淡翅盲蝽。【结论】捕食性盲蝽在稻田生态系统遭遇集团内捕食,其集团内捕食强度与水稻品种抗性、捕食者种类和猎物丰富度有关。     

Absence from the nest due to human disturbance induces higher nest predation risk than natural recesses in Common Eiders Somateria mollissima

Human disturbance of nesting birds may cause reduced breeding success. It is therefore necessary to assess the impact of disturbance to identify steps that minimize negative impacts. We carried out a study of nesting success at two adjacent colonies of Common Eider Somateria mollissima on the islands of Grindøya and Håkøya in northern Norway between 2006 and 2011. Over the study period, nesting success was consistently higher on Håkøya (69–82%) than on Grindøya (35–60%). Between 2009 and 2011 we used camera monitoring of individual nests to identify determinants of nest survival and predation, focusing in particular on the effect of departures from the nest due to human disturbance, which differed between the colonies due to a long‐term research project on Grindøya. Overall, absence of Common Eiders from nests due to disturbance increased the predation risk by a factor of 6.42 for an increase of one additional daily disturbance. In contrast, absence due to natural recesses did not increase nest losses. Under high levels of human disturbance, camera monitoring indicated that the main cause of breeding failure was predation, primarily by Hooded Crows Corvus cornix, but also to some extent Great Black‐backed Gulls Larus marinus. The presence of cameras did not increase the predation risk. Both the presence of researchers and the sight of Common Eider females conspicuously departing from nests are likely to have provided cues to these predators. We suggest management trials to reduce nesting disturbance through the guarding of unoccupied nests to mitigate the effects of human disturbance on reproductive success.     

Benefits and potential trade‐offs associated with yeast‐like symbionts during virulence adaptation in a phloem‐feeding planthopper

Insect herbivores form symbioses with a diversity of prokaryotic and eukaryotic microorganisms. A role for endosymbionts during host feeding on nutrient‐poor diets – including phloem – is now supported by a large body of evidence. Furthermore, symbiont‐herbivore associations have been implicated in feeding preferences by host races (mainly aphids) on multiple plant species. However, the role of symbionts in mediating herbivore preferences between varieties of the same plant species has received little research attention despite the implications for virulence adaptation to resistant crops. This study investigates the role of yeast‐like symbionts (YLS) in virulence adaptation and host plant switching among populations of the brown planthopper, Nilaparvata lugens (Stål) (Hemiptera: Delphacidae), that were selected on various rice [Oryza sativa L. (Poaceae)] lines differing in their resistance against herbivores. Planthopper fitness (nymph weight) declined where YLS densities were depleted through heat treatment. However, compared to normal symbiotic planthoppers, the depletion of symbionts did not generally change the relative fitness of planthoppers (each ‘adapted’ to a single natal host) when switched to feed on a range of rice lines (exposed hosts). In some cases, this occurred despite differences in YLS density responses to the various hosts. Furthermore, we detected no fitness costs associated with YLS in adapted populations. Therefore, the results of this study suggest that, whereas YLS are essential for planthopper nutrition, changes in YLS density play little role during virulence adaptation and host plant switching by the brown planthopper.     

Elevational contrast in predation and parasitism risk to caterpillars in a tropical rainforest

Invertebrate predators and parasitoids are among the most important natural enemies of insect herbivores. Yet, the strength of natural enemy pressure along an altitudinal gradient and interactions between the groups of natural enemies (such as predation on parasitized prey) are not well known. Various methods are used to reveal the mortality factors of herbivores. Predation pressure is usually assessed through exposure of artificial prey. However, this method cannot provide information about the attacks of parasitoids, or their eventual interactions with predators. Furthermore, artificial or dead prey might not attract predators because they do not show expected host behavior, and this method mostly cannot distinguish between predation and scavenging. For the first time in a tropical rainforest, we quantified elevational contrast in mortality factors using exposure of live caterpillars. We exposed a total of 800 live caterpillars of Talanga excelsalis moresbyensis Strand (Lepidoptera: Crambidae) on saplings of Ficus copiosa Steud. (Moraceae) at two elevations in primary tropical rain forest in Papua New Guinea (200 and 1 200 m a.s.l.). We exposed the caterpillars in two treatments: exposed to and protected from invertebrate predators and parasitoids. Disappearance of caterpillars was significantly higher in the exposed treatment. Furthermore, caterpillar disappearance was significantly higher in lowlands than in highlands (43 vs. 12%). We consider the vast majority of the disappearance to be due to predation, as migration of the caterpillars from the focal trees was not observed (except one caterpillar). This estimate of invertebrate predation rate corresponds with studies which used artificial caterpillar models. No significant difference in parasitism rate between the two elevations was observed (12 vs. 13%). The combination of the disappearance and parasitism rate patterns means that larval parasitoids face stronger pressure from invertebrate predators through higher predation of their hosts in the lowlands than in the highlands.