Origins of Susceptibility to Insect Herbivores in High-Yielding Hybrid and Inbred Rice Genotypes.

Several studies have reported higher damage from insect herbivores to hybrid compared to inbred (pure line) rice. We used a collection of 20 hybrid and 12 inbred genotypes from diverse origins to test the hypotheses that hybrid rice susceptibility is due to (a) the hybrid plant type and/or (b) rice phylogeny. We challenged the genotypes with Nilaparvata lugans (BPH), Sogatella furcifera (WBPH) and Scirpophaga incertulas (YSB) in greenhouse and screenhouse bioassays and monitored herbivores in field plots. We used single nucleotide polymorphic (SNP) markers to assess genetic similarities between the genotypes and found that the hybrids and inbreds formed two distinct clusters regardless of origin. In the screenhouse, hybrids were more susceptible than inbreds to YSB; however, resistant hybrids and susceptible inbreds were also apparent from both the screenhouse and field plots. Plant biomass was the best predictor of susceptibility to YSB. Plant origin had a greater effect than plant type on susceptibility to BPH and WBPH. WBPH was the most abundant planthopper in the field plots where numbers were highly correlated with planthopper fitness in the greenhouse bioassays. Our results provide evidence that high-yielding hybrids that are relatively resistant to herbivores can be achieved through careful breeding. The avoidance of susceptible genotypes during breeding should remain a key element of integrated rice pest management.

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.

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.

Combined Effects of Soil Silicon and Host Plant Resistance on Planthoppers, Blast and Bacterial Blight in Tropical Rice.

Soil silicon enhances rice defenses against a range of biotic stresses. However, the magnitude of these effects can depend on the nature of the rice variety. We conducted a series of greenhouse experiments to examine the effects of silicon on planthoppers (Nilaparvata lugens [BPH] and Sogatella furcifera [WBPH]), a leafhopper (Nephotettix virescens [GLH]), blast disease (Magnaporthe grisea) and bacterial blight (Xanthomonas oryzae) in susceptible and resistant rice. We added powdered silica gel (SiO2) to paddy soil at equivalent to 0.25, 1.0, and 4.0 t ha-1. Added silicon reduced BPH nymph settling, but the effect was negligible under high nitrogen. In a choice experiment, BPH egg-laying was lower than untreated controls under all silicon treatments regardless of nitrogen or variety, whereas, in a no-choice experiment, silicon reduced egg-laying on the susceptible but not the resistant (BPH32 gene) variety. Stronger effects in choice experiments suggest that silicon mainly enhanced antixenosis defenses. We found no effects of silicon on WBPH or GLH. Silicon reduced blast damage to susceptible and resistant (Piz, Piz-5 and Pi9 genes) rice. Silicon reduced damage from a virulent strain of bacterial blight but had little effect on a less virulent strain in susceptible and resistant (Xa4, Xa7 and Xa4 + Xa7 genes) varieties. When combined with resistance, silicon had an additive effect in reducing biomass losses to plants infested with bacterial blight (resistance up to 50%; silicon 20%). We discuss how silicon-containing soil amendments can be combined with host resistance to reduce biotic stresses in rice.

Nitrogenous Fertilizer Reduces Resistance but Enhances Tolerance to the Brown Planthopper in Fast-Growing, Moderately Resistant Rice.

The brown planthopper, Nilaparvata lugens (Stål), is a key challenge to rice production in Asia. Outbreaks of planthoppers are associated with excessive fertilizer applications; consequently, we examined planthopper interactions with susceptible, tolerant and resistant varieties of rice under varying levels of soil nitrogen in a greenhouse experiment. We compared planthopper fitness (survival × reproduction) and plant tolerance (functional plant loss index) for 16 varieties at 0, 80 and 150 Kg added nitrogen ha-1. The planthoppers grew larger, developed more quickly and laid more eggs on susceptible varieties, compared with the resistant and tolerant varieties. Moreover, soil nitrogen generally increased planthopper fitness on resistant varieties, but relative resistance was maintained. Functional plant loss was highest among the susceptible varieties, but weight and growth rate reductions per mg of planthopper were often highest in the tolerant varieties. Tolerance was associated with large, fast-growing plants, with at least moderate resistance to the planthopper. Susceptibility was associated with a small size and/or an absence of resistance genes. Our results suggested that early-tillering rice plants can be both resistant and tolerant to the brown planthopper, but cannot be both susceptible and tolerant of planthoppers at high densities. This indicates that at least moderate resistance is required for tolerance against this herbivore. Furthermore, although dwarf varieties had a low tolerance of planthoppers, they could express resistance through functioning resistance genes.

Efficacy and Cost-Effectiveness of Phenotyping for Rice Resistance and Tolerance to Planthoppers.

The standard seedling seedbox test (SSST) is the most prevalent phenotyping test in research on the genetics and breeding of planthopper-resistant rice. Using 16 rice lines that included plants susceptible, resistant and tolerant to the brown planthopper (Nilaparvata lugens), we compared the SSST to modified seedling seedbox tests (MSSTs) and the days-to-wilt (DTW) test. We also conducted a series of performance tests to assess nymph survival and development; adult longevity and egg-laying; egg survival; honeydew production; and plant weight loss. We also assessed the relative costs of the different phenotyping tests to better recommend test protocols that are suitable for high-throughput phenotyping. The SSST was found to be highly robust but fails to identify late-stage resistance; tolerance; or ovicidal responses. MSSTs improved phenotyping by identifying plants with low damage from planthoppers at later growth stages. Herbivore performance tests such as population or biomass build-up tests reduce space requirements and reduce setup and evaluation costs compared with bulk tests. They can also facilitate the assessment of plant tolerance; albeit with added costs. The DTW test most clearly segregates resistant and susceptible plants, thereby facilitating gene discovery and marker-assisted selection. We recommend that bulk testing be improved by switching from the SSST to a suitable MSST and that donor variety and pre-release lines be assessed for the nature of rice-planthopper interactions using biomass build-up tests-including the DTW test.

Immature stages, phenology, distribution and host plants of the Andean Moon Moth Cercophana frauenfeldii Felder, 1862 (Lepidoptera: Saturniidae)

Abstract Cercophana frauenfeldii Felder (Lepidoptera: Saturniidae), also known as the "Andean Moon Moth", is a Neotropical species native to continental Chile whose larvae feed on species of the families Gomortegaceae, Laureaceae and Winteraceae. We describe and document C. frauenfeldii immature stages, namely, egg, its four larval instars, and chaetotaxy of the last instar, pupa and cocoon for the first time. In terms of its phenology, we extend its larval activity, originally described to occur between November and mid-December, to June until the end of January. We report the adult flight period depends on the species' distributional range following two well-differentiated patterns: February to mid-April in Central-North Chile and April to June in Central-South Chile. Furthermore, we provide a unified view of its current distributional range and host plants (including the endangered tree Gomortega keule) through bibliographic data, field observations and laboratory rearing. Finally, we discuss aspects of the species' conservation as part of the unique ecosystems found in the temperate forests of southern South-America.

Larva, pupa and DNA barcodes of the Neotropical geometrid moth Glena mielkei (Lepidoptera: Geometridae: Ennominae: Boarmiini)

ABSTRACT Glena mielkei Vargas, 2010 (Lepidoptera: Geometridae: Ennominae: Boarmiini) is a Neotropical geometrid moth native to the Atacama Desert of northern Chile whose larvae are folivorous on the shrub Trixis cacalioides (Asteraecae). The last instar and pupa are described and illustrated, and DNA barcode sequences are provided for the first time for G. mielkei. Descriptions are made based on larvae collected in the type locality. Comparisons with the available descriptions of congeneric species suggest that the chaetotaxy of the SV group of the abdominal segment and the morphology of the cremaster could be useful tools to species identification based on last instar and pupa, respectively. A search in BOLD (Barcode of Life Data System) showed that the only DNA barcode haplotype found in the two specimens sequenced was closest to Physocleora Warren, 1897 than Glena Hulst, 1896. These results coincide with the morphological peculiarities of the genitalia highlighted in the original description of G. mielkei, suggesting that a definitive assessment of the generic status of this geometrid moth deserves further integrative studies.

Egg laying site selection by a host plant specialist leaf miner moth at two intra-plant levels in the northern Chilean Atacama Desert

Egg laying site selection by a host plant specialist leaf miner moth at two intra-plant levels in the northern Chilean Atacama Desert. The spatial distribution of the immature stages of the leaf miner Angelabella tecomae Vargas & Parra, 2005 was determined at two intra-plant levels (shoot and leaflet) on the shrub Tecoma fulva fulva (Cav.) D. Don (Bignoniaceae) in the Azapa valley, northern Chilean Atacama Desert. An aggregated spatial pattern was detected for all the immature stages along the shoot, with an age dependent relative position: eggs and first instar larvae were clumped at apex; second, third and fourth instar larvae were mostly found at intermediate positions; meanwhile the spinning larva and pupa were clumped at basis. This pattern suggests that the females select new, actively growing leaflets for egg laying. At the leaflet level, the immature stages were found more frequently at underside. Furthermore, survivorship was higher for larvae from underside mines. All these results highlight the importance of an accurate selection of egg laying site in the life history of this highly specialized leaf miner. By contrast, eventual wrong choices in the egg laying site selection may be associated with diminished larval survivorship. The importance of the continuous availability of new plant tissue in this highly human modified arid environment is discussed in relation with the observed patterns.

Observations on the biology and distribution of Uresiphita reversalis (Lepidoptera, Crambidae), a defoliator of the native tree Calia secundiflora in México

Observations on the biology and distribution of Uresiphita reversalis (Lepidoptera, Crambidae), a defoliator of the native tree Calia secundiflora in México. Uresiphita reversalis (Guenée, 1854) feeding on Calia secundiflora (Ortega) Yakovlev is recorded for the first time in the state of Hidalgo, Mexico. New aspects regarding the life cycle, feeding behaviour, geographical distribution and host plant damage by U. reversalis on C. secundiflora are here presented and discussed.

A replacement name for Hualpenia Mundaca, Parra & Vargas (Lepidoptera, Gracillariidae)

A replacement name for Hualpenia Mundaca, Parra & Vargas (Lepidoptera, Gracillariidae). Vihualpenia nom. nov. is proposed as a replacement name for Hualpenia Mundaca, Parra & Vargas, 2013 (Lepidoptera, Gracillariidae), in order to remove homonymy with Hualpenia Franz, 1996 (Coleoptera, Staphylinidae).

A new genus and species of leaf miner (Lepidoptera, Gracillariidae) for Chile associated to the native tree Lithraea caustica

A new genus and species of leaf miner (Lepidoptera, Gracillariidae) for Chile associated to the native tree Lithraea caustica. We propose the new genus and species of Gracillariidae (Lepidoptera) Hualpenia lithraeophaga Mundaca, Parra &Vargas gen. nov., sp. nov., leaf miner of Lithraea caustica (Mol.) H. et Arn (Anacardiaceae) occurring in southern central Chile. Aspects of the life cycle, adult and larval morphology, development and feeding habits of the new genus and species are also presented. We emphasise the uniqueness and importance of this new species for broadening the current knowledge on the Chilean fauna of Gracillariidae.

First remarks on the nesting biology of Hypodynerus andeus (Packard) (Hymenoptera, Vespidae, Eumeninae) in the Azapa valley, northern Chile / Primeiras observações sobre a biologia da nidificação de Hypodynerus andeus (Packard) (Hymenoptera, Vespidae, Eumeninae) no vale de Azapa, norte do Chile

First remarks on the nesting biology of Hypodynerus andeus (Packard) (Hymenoptera, Vespidae, Eumeninae) in the Azapa valley, northern Chile. Some aspects about the nesting biology of the potter wasp Hypodynerus andeus (Packard, 1869) are reported for the first time. Observations were carried out at the Azapa valley, coastal desert of northern Chile. A total of sixty nests were collected and examined, each composed by 1-14 cells, most of them found attached to concrete lamp posts. The only preys recorded in the cells were Geometridae (Lepidoptera) caterpillars and the presence of the parasitoid Anthrax sp. (Diptera, Bombyliidae) was also recorded. A number of arthropods belonging to different groups, mainly spiders, were found occupying empty nests.

Primeiras observações sobre a biologia da nidificação de Hypodynerus andeus (Packard) (Hymenoptera, Vespidae, Eumeninae) no vale de Azapa, norte do Chile. Alguns aspectos da biologia da nidificação da vespa Hypodynerus andeus (Packard, 1869) são registrados pela primeira vez. As observações foram conduzidas no vale de Azapa, deserto litoral do norte do Chile. Sessenta ninhos foram coletados e examinados, cada um composto por 1-14 células. A maioria dos ninhos estava aderida a postes de concreto. As únicas presas registradas nas células foram larvas de Geometridae (Lepidoptera). Um parasitóide, Anthrax sp. (Diptera, Bombyliidae), foi também registrado. Vários artrópodes, principalmente aranhas, foram encontrados utilizando ninhos vazios.