Landscape heterogeneity filters functional traits of rice arthropods in tropical agroecosystems.

Biological control services of agroecosystems depend on the functional diversity of species traits. However, the relationship between arthropod traits and landscape heterogeneity is still poorly understood, especially in tropical rice agroecosystems, which harbor a high diversity of often specialized species. We investigated how landscape heterogeneity, measured by three metrics of landscape composition and configuration, influenced body size, functional group composition, dispersal ability, and vertical distribution of rice arthropods in the Philippines. We found that landscape composition and configuration acted to filter arthropod traits in tropical rice agroecosystems. Landscape diversity and rice habitat fragmentation were the two main gradients influencing rice-arthropod traits, indicating that different rice arthropods have distinct habitat requirements. Whereas small parasitoids and species mostly present in the rice canopy were favored in landscapes with high compositional heterogeneity, predators and medium-sized species occupying the base of the rice plant, including planthoppers, mostly occurred in highly fragmented rice habitats. We demonstrate the importance of landscape heterogeneity as an ecological filter for rice arthropods, identifying how the different components of landscape heterogeneity selected for or against specific functional traits. However, the contrasting effects of landscape parameters on different groups of natural enemies indicate that not all beneficial rice arthropods can be promoted at the same time when using a single land management strategy. Increasing compositional heterogeneity in rice landscapes can promote parasitoids but may also negatively affect predators. Future research should focus on identifying trade-offs between fragmented rice habitats and structurally diverse landscapes to maximize the presence of multiple groups of beneficial arthropods.

Stem borers revisited: Host resistance, tolerance, and vulnerability determine levels of field damage from a complex of Asian rice stemborers.

Yield losses from rice stem borers depend on the nature of the rice variety, the timing of attack, and the composition of the stem borer species assemblage. This study uses a range of phenotyping methods to distinguish different categories of herbivore-rice interaction that determine relative damage levels (dead heart and whitehead-panicles) and yield losses to varieties exposed to stem borers. Phenotyping studies were conducted in a greenhouse, screen house and field using two stem borer species (Scirpophaga incertulas [yellow stem borer - YSB] and Chilo suppressalis [striped stem borer - SSB]) and 12 rice lines. Only YSB displayed oviposition preferences across rice varieties. Both stem borer species performed better (greater survival, shorter development times, heavier pupae) on rice at vegetative compared to reproductive growth stages, and SSB was less capable than YSB of developing on rice at reproductive growth stages. Stem borer larval survival, body weight, development time, and tiller damage across different rice varieties in greenhouse experiments was poorly correlated between the two stem borer species and for each of the species on rice at two different plant growth stages (vegetative and reproductive). In contrast, rice tillering and yield responses to the two stem borer species were often highly correlated, but only when plants were attacked at the reproductive stage. Short-term, controlled experiments revealed aspects of host resistance and relative changes in plant biomass and yield in response to damage (condition change). However, the results from controlled environments and field-plots were not generally correlated because plant vulnerability, i.e., relative exposure to stem borer attack due to crop duration, largely determined field damage. We recommend that phenotyping studies should differentiate between the resistance, tolerance and vulnerability of rice lines to complexes of stem borers in a given region. Single stem borer species experiments under controlled environments are useful to gain knowledge of the nature of rice-stem borer interactions; however, field testing with comparative treatments, particularly under high soil fertilizer levels that increase plant attractiveness, are better for assessing the relative propensities of rice varieties to incur damage and yield losses due to complexes of stem borers.

Changes in insecticide resistance and host range performance of planthoppers artificially selected to feed on resistant rice.

Host plant resistance has received considerable attention for the management of insect herbivores on crop plants. However, resistance is threatened by the rapid adaptation of target herbivores towards virulence (the ability to survive, develop and damage a host with major resistance genes). This study examines the potential costs and benefits of adaptation for virulence in herbivores. We continuously reared planthoppers, Nilaparvata lugens, on two susceptible and three resistant rice, Oryza sativa, varieties for 20 + generations. We then assessed the performance of selected planthoppers across a range of rice lines with distinct resistance genes. We found that planthoppers with long-term exposure to resistant hosts (particularly IR62 with the Bph3(t) and BPH32 gene loci, and PTB33 with the Bph3(t), BPH32 and BPH26 gene loci) gained virulence against related varieties with the same and different resistance genes, but planthoppers adapted to the resistant host IR65482-4-136-2-2 (BPH10 locus) had reduced performance on phylogenetically distant plants with distinct resistant genes. In choice bioassays, avirulent planthoppers showed marked preferences for susceptible lines, whereas virulent planthoppers were less selective of varieties. We also examined whether virulence was associated with insecticide susceptibility. We tested susceptibility to three insecticides using a topical application method. Populations selectively reared on IR65482-4-136-2-2 had increased susceptibility to imidacloprid and fipronil, representing a possible trade-off with virulence. In contrast, a population with virulence to the highly resistant variety PTB33 was 4.88 × more resistant to imidacloprid and 3.18 × more resistant to BPMC compared to planthoppers of the same origin but reared on the susceptible variety IR22. Our results suggest complex relations between insecticide resistance and virulence that vary according to insecticidal toxins and resistance genes, and include potentially increased insecticide-susceptibility (a trade-off) as well as common detoxification mechanisms (a benefit).

Use and Avoidance of Pesticides as Responses by Farmers to change Impacts in Rice Ecosystems of Southern Sri Lanka.

Since the beginning of the millennium, pesticide use has increased dramatically across Asia. However, it is still unclear what drives pesticide use among low-income farmers. We surveyed 275 farmers to investigate their decisions to apply or avoid pesticides on rice farms in southern Sri Lanka. Based on the information from the survey, we applied the Drivers-Pressures-State-Impacts-Responses framework to assess farmers' decisions as 'Responses' to changes in the rice ecosystem. Organic farmers mainly planted traditional varieties (64-77%) that they perceived as resistant to environmental stresses, including insect herbivores. Pesticide users, including integrated pest management farmers, and insecticide-free farmers (without insecticide use for ≥5 years) mainly planted modern rice varieties. There were no significant differences in reported rice yields among farmers using different management practices. However, fewer organic and insecticide-free farmers reported insect pests, particularly planthoppers, as a production constraint. Insecticide use by farmers was mainly an adaptive response to perceived risks of pest damage to high-yielding varieties. But farmers' responses were largely unsupported by any previous experience, suggesting that an increased availability and marketing of agrochemicals, and pressures to modernize production have influenced their decisions to use insecticides. Organic and insecticide-free farming practices were mainly restorative responses to perceived health risks associated with agrochemicals. We suggest that attention could be placed on promoting 'pesticide-free' or 'insecticide-free' farming through certification schemes as a viable option to break the lock-in of pesticide use on rice farms in the region and to provide additional product choices for rice consumers.

Unanticipated benefits and potential ecological costs associated with pyramiding leafhopper resistance loci in rice.

We tested the hypotheses that increasing the number of anti-herbivore resistance loci in crop plants will increase resistance strength, increase the spectrum of resistance (the number of species affected), and increase resistance stability. We further examined the potential ecological costs of pyramiding resistance under benign environments. In our experiments, we used 14 near-isogenic rice lines with zero (T65: recurrent parent), one, two or three resistance loci introgressed through marker-assisted selection. Lines with two or more loci that were originally bred for resistance to the green rice leafhopper, Nephotettix cincticeps, significantly reduced egg-laying by the green leafhopper, N. virescens. Declines in egg-number and in nymph weight were correlated with the numbers of resistance loci in the rice lines. To test the spectrum of resistance, we challenged the lines with a range of phloem feeders including the zig-zag leafhopper, Recilia dorsalis, brown planthopper, Nilaparvata lugens, and whitebacked planthopper, Sogatella furcifera. There was an increase in the number of tested species showing significant declines in egg-laying and nymph survival on lines with increasing numbers of loci. In a screen house trial that varied rates of nitrogenous fertilizer, a line with three loci had stable resistance against the green leafhopper and the grain yields of infested plants were maintained or increased (overcompensation). Under benign conditions, plant growth and grain yields declined with increasing numbers of resistance loci. However, under field conditions with natural exposure to herbivores, there were no significant differences in final yields. Our results clearly indicate the benefits, including unanticipated benefits such as providing resistance against multiple herbivore species, of pyramiding anti-herbivore resistance genes/loci in crop plants. We discuss our results as part of a review of existing research on pyramided resistance against leafhoppers and planthoppers in rice. We suggest that potential ecological costs may be overcome by the careful selection of gene combinations for pyramiding, avoidance of high (potentially redundant) loci numbers, and introgression of loci into robust plant types such as hybrid rice varieties.

Resistance and tolerance to the brown planthopper, Nilaparvata lugens (Stål), in rice infested at different growth stages across a gradient of nitrogen applications.

High resource availability can reduce anti-herbivore resistance (a plant's ability to defend against herbivores and reduce damage) in rice, Oryza sativa L, but may also increase tolerance (a plant's ability to withstand damage by, for example, compensatory growth). Through a series of greenhouse, screenhouse and field experiments, this study examines fitness (survival and development × reproduction) of the brown planthopper, Nilaparvata lugens (Stål), on resistant (IR62) and susceptible (IR22) rice varieties and age-related rice tolerance to planthopper damage under varying resource (nitrogenous fertilizer) availability. Planthoppers reared on IR62 in the greenhouse had lower fitness than planthoppers on IR22. IR62 became increasingly resistant as plants aged. IR22 was generally more tolerant of planthopper damage, and tolerance increased in IR22, but declined in IR62, as the plants aged. Rice plants infested at pre-tillering stages (3-4 leaf stage) in the screenhouse had greater losses to root, shoot and grain yield per unit weight of planthopper than plants infested at tillering stages, particularly in IR22. These trends were mainly due to the impact of planthoppers during pre-tillering stages and the length of exposure to the planthoppers. High nitrogen compromised IR62 resistance, particularly in tillering plants in the greenhouse study; however, high nitrogen did not increase planthopper biomass-density on IR62 in greenhouse or field cages. Tolerance to damage in IR62 at mid-tillering stages declined under increasing levels of nitrogen, but nitrogen increased tolerance during late-tillering stages. Planthopper damage to IR22 in field cages was severe and hopperburn (plant death) occurred in 83% of IR22 plants under high nitrogen (60-150 kg N ha-1). In contrast, despite planthopper infestations, damage to IR62 was low in field-grown plants and productivity (tillers, roots, shoots and grain) increased in IR62 under increasing nitrogen. Our results indicate that, whereas nitrogenous fertilizer increases planthopper fitness on susceptible and resistant varieties, the net effects of high nitrogen on IR62 include decreased planthopper biomass-density (apparent in all experiments) and higher tolerance to damage during later growth stages (observed in the greenhouse, and during one of two seasons in field cages).

Virulence adaptation in a rice leafhopper: Exposure to ineffective genes compromises pyramided resistance.

Pyramiding resistance genes is predicted to increase the durability of resistant rice varieties against phloem-feeding herbivores. We examined responses by the green leafhopper, Nephotettix virescens (Hemiptera: Cicadellidae), to near-isogenic rice lines with zero, one and two resistance genes. The recurrent parent (T65) and monogenic lines (GRH2-NIL and GRH4-NIL) with genes for resistance to the green rice leafhopper, Nephotettix cincticeps (Hemiptera: Cicadellidae), were susceptible to the green leafhopper, but the pyramided line (GRH2/GRH4-PYL) was highly resistant to the green leafhopper. We selected green leafhoppers, N. virescens, from five sites in the Philippines for over 20 generations on each of the four lines. Populations selected on GRH2/GRH4-PYL gained partial virulence (feeding and development equal to that on T65) to the pyramided line within 10 generations and complete virulence (egg-laying equal to that on T65) within 20 generations. After 20 generations of rearing on the susceptible monogenic lines, green leafhoppers were also capable of developing and laying eggs on GRH2/GRH4-PYL. Furthermore, green leafhoppers reared on the susceptible GRH4-NIL for 20 generations showed equal preferences for T65 and GRH2/GRH4-PYL in choice bioassays. Our results indicate that previous long-term exposure to ineffective genes (including unperceived resistance genes) could dramatically reduce the durability of pyramided resistance. We suggest that informed crop management and deployment strategies should be developed to accompany rice lines with pyramided resistance and avoid the build-up of virulent herbivore populations.

Effects of nitrogen on egg-laying inhibition and ovicidal response in planthopper-resistant rice varieties.

A series of experiments was set up to examine the effects of nitrogen on rice (Oryza sativa L.) resistance against Nilaparvata lugens (Stål) and Sogatella furcifera (Horváth). Egg laying by N. lugens was reduced on the indica variety IR60. Nymph biomass (N. lugens and S. furcifera) was also lower on IR60: this was associated with low honeydew production and a high proportion of xylem-derived honeydew in N. lugens but not in S. furcifera. Nitrogen increased egg-laying by S. furcifera and increased N. lugens nymph biomass on all varieties tested. Oviposition and egg mortality in both planthopper species were examined on plants at 15, 30 and 45 days after sowing (DAS). Sogatella furcifera laid more eggs on plants at 15 DAS, but laid few eggs during darkness; N. lugens continued to lay eggs on older rice plants (30 DAS) and during darkness. Egg mortality was high on cv. Asiminori, highest at 45 DAS, and higher for S. furcifera than for N. lugens. Mortality of S. furcifera eggs was associated with lesions around the egg clusters. These were more common around clusters laid during the day and suggested induction by Asiminori of an ovicidal response. Egg mortality declined under higher soil nitrogen levels. Results are discussed in the light of improving rice resistance against planthoppers and reducing rates of planthopper adaptation to resistance genes.

Population genetic structure of Bombus terrestris in Europe: Isolation and genetic differentiation of Irish and British populations.

The genetic structure of the earth bumblebee (Bombus terrestris L.) was examined across 22 wild populations and two commercially reared populations using eight microsatellite loci and two mitochondrial genes. Our study included wild bumblebee samples from six populations in Ireland, one from the Isle of Man, four from Britain and 11 from mainland Europe. A further sample was acquired from New Zealand. Observed levels of genetic variability and heterozygosity were low in Ireland and the Isle of Man, but relatively high in continental Europe and among commercial populations. Estimates of Fst revealed significant genetic differentiation among populations. Bayesian cluster analysis indicated that Irish populations were highly differentiated from British and continental populations, the latter two showing higher levels of admixture. The data suggest that the Irish Sea and prevailing south westerly winds act as a considerable geographical barrier to gene flow between populations in Ireland and Britain; however, some immigration from the Isle of Man to Ireland was detected. The results are discussed in the context of the recent commercialization of bumblebees for the European horticultural industry.

Heterosis for Interactions between Insect Herbivores and 3-Line Hybrid Rice under Low and High Soil Nitrogen Conditions.

Hybrid rice results from crossing a male-sterile line (the A line) with a pollen doner (the restorer or R line). In 3-line hybrid breeding systems, a fertile B line is also required to maintain A line populations. Heterosis is defined as a condition of traits whereby the hybrid exceeds the average of the parental lines. Heterobeltiosis is where the hybrid exceeds both parents. Hybrid rice may display heterosis/heterobeltiosis for growth, yield and resistance to herbivores, among other traits. In a greenhouse experiment, we assessed the frequency of heterosis for resistance to the brown planthopper (Nilaparvata lugans (BPH)), whitebacked planthopper (Sogatella furcifera (WBPH)) and yellow stemborer (Scirpophaga incertulas (YSB)) in eight hybrids under varying soil nitrogen conditions. We also assessed plant biomass losses due to herbivore feeding as an approximation of tolerance (the plant's capacity to compensate for damage). Nitrogen reduced resistance to all three herbivores but was also associated with tolerance to WBPH and YSB based on improved plant survival, growth and/or yields. Plant biomass losses per unit weight of WBPH also declined under high nitrogen conditions for a number of hybrids, and there were several cases of overcompensation in rice for attacks by this herbivore. There was one case of nitrogen-related tolerance to BPH (increased grain yield) for a hybrid line with relatively high resistance, likely due to quantitative traits. Heterosis and heterobeltiosis were not essential to produce relatively high herbivore resistance or tolerance across hybrids.

Heterosis for Resistance to Insect Herbivores in a 3-Line Hybrid Rice System.

Three-line hybrid rice is produced by crossing male sterile (A line) rice with a fertility-restorer (R line). Fertile lines (B lines) are also required to maintain A line seed for breeding programs. We used a range of hybrids and their parental lines to assess the frequency and nature of heterosis for resistance to the whitebacked planthopper (Sogatella furcifera), brown planthopper (Nilaparvata lugens) and yellow stemborer (Scirpophaga incertulas). Heterosis is defined as trait improvement above the average of the parental lines as a result of outbreeding. Based on the results from a greenhouse study that challenged hybrids and their parental lines with each herbivore species, we found that susceptibility to planthoppers was associated with one of the eight A lines tested, but resistance was improved by crossing with a relatively resistant restorer. Higher frequencies of heterosis for susceptibility in comparisons between hybrids and their B lines suggest that susceptibility was not related to the cytoplasmic genomes of the associated sterile A lines. Furthermore, because none of the parental lines possessed currently effective resistance genes, improved resistance against planthoppers was probably due to quantitative resistance. In a related field trial, hybrids had generally higher yields than their fertile parents and often produced larger grain; however, they were often more susceptible to stemborers, leaffolders (Cnaphalocrocis medinalis) and other caterpillars (Rivula atimeta). This was largely a consequence of hybrid heterosis for plant biomass and was strongly affected by crop duration. We make a series of recommendations to improve hybrid breeding to reduce the risks of herbivore damage.

Pathogen prevalence in commercially reared bumble bees and evidence of spillover in conspecific populations.

Worldwide, wild bumble bees (Bombus spp.) are experiencing marked declines, with potentially up to 11% of species currently under threat. Recent studies from North America suggest that disease transmission from commercially reared bumble bees to wild populations has led to marked range contractions in some species. In Europe, data on the prevalence of pathogen spillover from commercial to wild bumble bee populations is lacking, despite the widespread production and transport of hives within the EU since the early 1980s. We determined the permeability of cropping systems to commercial bumble bees, and quantified the prevalence of four pathogens in commercial Bombus terrestris hives and adjacent conspecific populations at increasing distances from greenhouses in Ireland. Commercial bumble bees collected from 31% to 97% of non-crop pollen, depending on the cropping system, and hives had markedly higher frequencies of two gut parasites, Crithidia spp. and Nosema bombi, compared to adjacent populations, but were free of tracheal mites. The highest prevalence of Crithida was observed within 2 km of greenhouses and the probability of infection declined in a host sex- and pathogen-specific manner up to 10 km. We suggest implementing measures that prevent the interaction of commercially reared and wild bumble bees by integrating the enforcement of national best management practices for users of commercial pollinators with international legislation that regulates the sanitation of commercial hives in production facilities.

Slowing virulence adaptation in Asian rice planthoppers through migration-based deployment of resistance genes.

Advances in molecular biology have accelerated rice breeding for resistance to Asian planthoppers. However, experience shows that planthoppers quickly adapt to resistance in tropical overwintering areas. With only limited sources available, the large-scale deployment of resistance genes can rapidly reduce the utility of these public goods. Planthoppers that migrate from tropical to temperate Asia carry virulence against many resistance genes, but adapt more slowly to resistant rice in temperate regions. Therefore, by restricting deployment of selected genes to temperate regions, virulence-adaptation rates and the volume of migrants returning to overwintering sites could be reduced. The current open exchange of breeding materials throughout Asia urgently requires an international, multidisciplinary, stakeholder coalition to promote a more sustainable deployment of planthopper-resistant rice.

The Structure of Rice Stemborer Assemblages: A Review of Species' Distributions, Host Ranges, and Interspecific Interactions.

This review describes global rice stemborer assemblages based on published species distributions, apparent host preferences, and reported shifts in assemblage composition in response to environmental factors. At least 56 moth (Lepidoptera: Crambidae, Pyralidae, Noctuidae) and fly (Diptera: Diopsidae, Chloropidae) species have been associated with rice; however, only 21 species are of potential, large-scale economic importance with a further 2 species of localized concern; most of the remaining species' associations with rice are based on dubious records without economic impacts on rice production. A list of stemborer-host associations indicates that rice stemborers are largely oligophagous on grasses (Poaceae), but a few species are polyphagous (also attacking Cyperaceae, Typhaceae, and some Eudicotyledon plants). Total stemborer abundance is determined by rice cropping patterns and management. Assemblage species richness is determined by geographical location, surrounding habitat (particularly as regards secondary and occasional species), and season. Evidence suggests that stemborer assemblage structure is largely determined through conditional interspecific competition. Regional assemblages typically include a single dominant lepidopteran species (primary species) that is largely restricted to rice and for which the climate is optimal; one or more secondary species that vary based on the age of rice attacked, rice anatomy, and the proximity to other habitats (including other crops); and occasional species that probably spill over from adjacent grasslands. The co-occurrence of lepidopteran with dipteran rice stemborers requires further research attention.

Diversified Rice Farms with Vegetable Plots and Flower Strips Are Associated with Fewer Pesticide Applications in the Philippines.

Ecological engineering is defined as the design of sustainable ecosystems for the benefit of both human society and the environment. In Southeast Asia, researchers have applied ecological engineering by diversifying farms using flower strips to restore regulatory services to rice ecosystems and thereby reduce herbivore-related yield losses and overall pesticide use. We conducted a survey of 302 rice farmers across four regions of the Philippines to assess their farm diversification practices and determine possible associations with pesticide use. Rice was the main product on all farms; however, the farmers also produced fruits and vegetables, either rotated with rice (47% of the farmers) or in small plots in adjacent farmland. In addition, 64% of the farmers produced flowers, herbs, and/or vegetables on rice bunds. Vegetables were cultivated mainly to supplement household food or incomes, but 30% of the farmers also believed that the vegetables reduced pest and weed damage to their rice. We found that 16% of the farmers grew flowers on their bunds to reduce pest damage to rice and vegetables, and many farmers applied botanical extracts, growth stimulants, and insect traps to reduce damage to the vegetables. Some farmers avoided insecticides on rice by using Trichogramma cards. Planting flowers on rice bunds, rearing ducks in the rice fields, and farmers' recognition of beneficial rice arthropods were statistically significantly associated with lower pesticide (particularly, insecticide) applications to rice. Our results indicate that farm diversification to produce supplementary foods for rural households and access to alternative pest management options can reduce pesticide use on rice farms in tropical Asia.

Agriculture without paraquat is feasible without loss of productivity-lessons learned from phasing out a highly hazardous herbicide.

A small proportion of the thousands of pesticides on the market today are associated with a disproportionately high incidence of severe acute pesticide poisoning and suicide. Paraquat stands out as one of the most lethal pesticides in common use, frequently involved in fatal incidents due to suicides or accidental exposure. Even though paraquat has been banned in over 67 countries, it is still widely used in many others, particularly in Asia and Latin America. Based on a literature review and consultations, this paper identifies options for replacing paraquat and distils practical lessons from numerous successes around the world. Our aim is to support regulators, policymakers, agronomists and the supply chain sector with practical information related to phasing out paraquat. Production data consistently failed to show any negative effects of banning paraquat on agricultural productivity. A wide range of alternative approaches to weed management and crop defoliation are available, many of which do not rely on herbicides. Over 1.25 million farmers in low- and middle-income countries (LMICs) successfully produce a range of crops for private voluntary standards (PVS) in food and fiber supply chains which prohibit paraquat use. We conclude from the findings of this study that eliminating paraquat will save lives without reducing agricultural productivity. Less hazardous and more sustainable alternatives exist. To enhance successful adoption and uptake of these methods on a wide scale, farmers require training and support within an enabling policy environment.

Dung removal increases under higher dung beetle functional diversity regardless of grazing intensification.

Dung removal by macrofauna such as dung beetles is an important process for nutrient cycling in pasturelands. Intensification of farming practices generally reduces species and functional diversity of terrestrial invertebrates, which may negatively affect ecosystem services. Here, we investigate the effects of cattle-grazing intensification on dung removal by dung beetles in field experiments replicated in 38 pastures around the world. Within each study site, we measured dung removal in pastures managed with low- and high-intensity regimes to assess between-regime differences in dung beetle diversity and dung removal, whilst also considering climate and regional variations. The impacts of intensification were heterogeneous, either diminishing or increasing dung beetle species richness, functional diversity, and dung removal rates. The effects of beetle diversity on dung removal were more variable across sites than within sites. Dung removal increased with species richness across sites, while functional diversity consistently enhanced dung removal within sites, independently of cattle grazing intensity or climate. Our findings indicate that, despite intensified cattle stocking rates, ecosystem services related to decomposition and nutrient cycling can be maintained when a functionally diverse dung beetle community inhabits the human-modified landscape.

Compatibility of Insecticides with Rice Resistance to Planthoppers as Influenced by the Timing and Frequency of Applications.

The brown planthopper, Nilaparvata lugens (Stål)(BPH) is a pest of rice in Asia. We examined the effects of seven insecticides combined with host resistance against BPH. In a screenhouse environment, we treated BPH-infested and non-infested resistant (IR62) and susceptible (IR64) rice with buprofezin, carbofuran, cartap hydrochloride, cypermethrin, deltamethrin, fipronil, or thiamethoxam + chlorantraniliprole. In one experiment, plants received one, two or three applications. In a second experiment, plants received one early or late insecticide application. Carbofuran and fipronil reduced planthopper biomass densities but resistance did not contribute to these effects (i.e., resistance was redundant). Single applications of cartap hydrochloride (at 20 or 50 days after sowing (DAS)), cypermethrin (20 DAS), or buprofezin (50 DAS) reduced BPH biomass densities on IR62 (i.e., synergies); other insecticides and application times, and multiple applications of all insecticides did not reduce BPH biomass densities on IR62 more than on IR64 (i.e., either resistance or insecticides were redundant). Deltamethrin (three applications) was antagonistic to resistance, but host resistance tended to buffer against the negative effects of single deltamethrin applications. Yields of infested IR62 were not statistically improved by insecticide applications. Late applications reduced yields of non-infested rice. We discuss how prophylactic insecticide applications could destabilize BPH populations and reduce the productivity and profitability of resistant rice.

Interactions between Rice Resistance to Planthoppers and Honeydew-Related Egg Parasitism under Varying Levels of Nitrogenous Fertilizer.

Host plant resistance is the most researched method for the management of planthoppers and leafhoppers in tropical rice. For optimal effects, resistance should be resilient to fertilizer inputs and work in synergy with natural enemies. In field plot experiments, we examined how rice resistance and fertilizer inputs affect mortality of planthopper and leafhopper eggs by hymenopteran parasitoids. We used IR62 as a variety with resistance to Nilaparvata lugens (Stål) [BPH], Sogatella furcifera (Horváth) [WBPH] and Nephotettix virescens (Distant) [GLH], and IR64 as a susceptible control. The herbivores were more abundant during wet season sampling in low-nitrogen plots. During this study, parasitoids killed between 31 and 38% of BPH eggs and 24 and 52% of WBPH eggs during four days of field exposure. Parasitism, mainly due to Oligosita spp., was generally higher in high-nitrogen and IR64 plots. Similar densities of eggs in exposed plants suggest that these trends were mediated by semiochemicals and therefore support the Optimal Defense Hypothesis. Honeydew from BPH on IR62 had more xylem-derived wastes than honeydew on IR64. We applied honeydew from both varieties to sentinel plants. Parasitism by Anagrus spp. was higher on plants of either variety treated with honeydew derived from IR62; however, the effect was only apparent in high-nitrogen plots. Results suggest that Anagrus spp., by responding to honeydew, will counter the nitrogen-induced enhancement of planthopper fitness on resistant rice.

Death in the paddy field: Carcass decomposition and associated arthropods in subunits of a rice field landscape.

Although rice production landscapes are often the scene of homicides, vertebrate decomposition and associated arthropods have never been described from rice paddies. Tropical rice landscapes are typically composed of irrigated/non-irrigated rice fields, fallow land (i.e., uncultivated fields), and low earthen levees (bunds) used as access pathways. The specific microclimatic and environmental conditions associated with each subunit of rice field landscapes are likely to impact carcass decomposition as well as the arthropod species associated with carrion. Here, we tested the hypothesis that the combined effects of constraints on arthropod colonization and survival, scarcity of necromass, limited habitat architecture, and recurrent disturbance limit the ability of carrion-related arthropods to colonize corpses and large carcasses in tropical rice-fields. Our results from monitoring pig carcasses in Philippine rice fields indicated that vertebrate decomposition in irrigated fields was slower and incomplete when compared to non-irrigated fields and bunds. Carcasses were colonized by a small complex of carrion-related arthropods that differed in composition and relative species abundance between dry bunds and relatively humid rice paddies. Fire ants (Solenopsis germinata) were observed frequently on carcasses exposed on bunds, rarely in non-irrigated fields, and almost never in irrigated fields. The presence of fire ants was associated with reduced blow fly (Chrysomya megacephala) abundance. Taken together, this indicates that the arthropod fauna associated with carcasses in tropical rice fields is relatively simple in contrast to the generally high arthropod diversity reported for other ecosystems at tropical latitudes. The limited richness of the community also means that an understanding of the development of one abundant calliphorid blow fly, C. megacephala, may be sufficient to investigate deaths and homicides under conditions similar to the ones described in this study.