Despite over 30 years of deployment, varieties with the Bph3 gene for resistance to the brown planthopper (BPH), Nilaparvata lugens (Stål) (Hemiptera: Delphacidae), are still effective in much of the Philippines. In the present study, we determined the effects of adaptation to one resistant variety, IR62 – assumed to possess the Bph3 gene – on (1) resistance against a series of varieties with similar biotypical responses (presumed to contain the same major resistance genes), and (2) a differential variety with the bph4 gene that occurs at the same chromosome position as Bph3. We also examined the effects of high soil nitrogen on the effectiveness of Bph3. Feeding, planthopper biomass, and development times were reduced in a wild BPH population when reared on IR62 compared with the susceptible standard variety TN1. However, nitrogen application increased the susceptibility of IR62. After 13 generations on IR62, BPH had adapted to the plant’s resistance. Virulence of the adapted BPH against the variety ‘Rathu Heenati’ supports the idea that Bph3 is present in IR62. Across similar IR varieties (IR60, IR66, IR68, IR70, IR72, and IR74), feeding, planthopper biomass, and development rates were generally higher for IR62‐adapted than for non‐adapted BPH; however, contrary to expectations, many of these varieties were already susceptible to wild BPH. Fitness was also higher for IR62‐adapted BPH on the variety ‘Babawee’ indicating a close relation between Bph3 and bph4. The results indicate that the conventional understanding of the genetics behind resistance in IR varieties needs to be readdressed to develop and improve deployment strategies for resistance management.

The brown planthopper (Nilapavata lugens: BPH) and whitebacked planthopper (Sogatella furcifera: WBPH) co-occur as the principal pests of rice in Asia. A review of previous studies suggests that the two species have similar temperature tolerances and similar temperature thresholds for development. However, the distribution and seasonality of WBPH suggest that its temperature optima for performance (survival, oviposition and growth) may be lower than for BPH. We compared adult longevity, oviposition, nymph survival and development success, as well as nymph biomass in both species across a gradient of constant temperatures from 15˚C-40˚C, at 5˚C intervals. The most suitable temperatures for oviposition, nymph biomass and development success were 5-10˚C lower for WBPH than for BPH. Furthermore, compared to BPH, WBPH demonstrated clear differences in oviposition on different rice subspecies and on rice at different growth stages at 25˚C and 30˚C, but not at other temperatures. The results suggest that aspects of herbivore performance within tolerable temperature ranges, which are not often included in temperature models, may be more useful than thermal tolerances or development thresholds in predicting the effects of global warming on pest damage to crops.

This study compares the effects of temperature (constant at 15, 20, 25, 30 and 35 °C) on adult longevity, oviposition, and nymph development of the brown planthopper, Nilaparvata lugens, on susceptible and resistant rice varieties. The resistant variety contained the BPH32 gene. In our experiments, nymphs failed to develop to adults at 15, 20 and 35 °C on either variety. Host resistance had its greatest effect in reducing adult survival at 20–25 °C and its greatest effect in reducing nymph weight gain at 25 °C. This corresponded with optimal temperatures for adult survival (20–25 °C) and nymph development (25–30 °C). At 25 and 30 °C, adult females achieved up to three oviposition cycles on the susceptible variety, but only one cycle on the resistant variety. Maximum egg-laying occurred at 30 °C due to larger numbers of egg batches produced during the first oviposition cycle on both the susceptible and resistant varieties, and larger batches during the second and third oviposition cycles on the susceptible variety; however, resistance had its greatest effect in reducing fecundity at 25 °C. This revealed a mismatch between the optimal temperatures for resistance and for egg production in immigrating females. Increasing global temperatures could reduce the effectiveness of anti-herbivore resistance in rice and other crops where such mismatches occur.

1. Global warming is often predicted to increase damage to plants through direct effects on insect herbivores. However, the indirect impacts of rising temperatures on herbivores, mediated through interactions with their biotic environment, could dampen these effects. 2. Using a series of reciprocal density experiments with gravid females and developing nymphs, we examined interspecific competition between two coexisting phloem feeders Nilaparvata lugens (BPH) and Sogatella furcifera (WBPH), on rice at 25 and 30°C. 3. WBPH performed better (i.e. adults survived longer, nymphs developed faster and grew larger) at 25°C and BPH (i.e. nymphs developed faster) at 30°C. However, contrary to predictions, WBPH had a greater effect in reducing oviposition and nymph performance in BPH at 30°C. 4. A decoupling of resource use by WBPH and its antagonistic effects on BPH at the higher temperature suggests that WBPH feeding induces host defences that reduce BPH fitness (i.e. interference competition). Meanwhile, BPH facilitated WBPH oviposition at 30°C and facilitated WBPH nymph performance at 25 and 30°C. Greater facilitation of feeding in WBPH nymphs by BPH at high densities suggests that mechanical damage and host responses to damage increased the fitness of the heterospecific nymphs. 5. Although BPH also facilitated egg-laying by WBPH, intra-and interspecific crowding countered this facilitation at both temperatures. Simulated life tables for planthoppers at 25 and 30°C depicted significantly lower offspring numbers on rice infested by WBPH alone and from mixed BPH-WBPH infestations than from infestations by BPH alone. 6. Our results indicate how interference competition-mediated through host plant defences-can increase ecosystem resilience to the warmer temperatures predicted under global climate change.

The direct effects of rising global temperatures on insect herbivores could increase damage to cereal crops. However, the indirect effects of interactions between herbivores and their biotic environment at the same temperatures will potentially counter such direct effects. This study examines the potential for intraspecific competition to dampen the effects of optimal temperatures on fitness (survival × reproduction) of the brown planthopper, Nilaparvata lugens [BPH] and whitebacked planthopper, Sogatella furcifera [WBPH], two phloem-feeders that attack rice in Asia. We conducted a series of experiments with increasing densities of ovipositing females and developing nymphs on tropical and temperate rice varieties at 25, 30 and 35°C. Damage from planthoppers to the tropical variety was greater at 30°C compared to 25°C, despite faster plant growth rates at 30°C. Damage to the temperate variety from WBPH nymphs was greatest at 25°C. BPH nymphs gained greater biomass at 25°C than at 30°C despite faster development at the higher temperature (temperature-size rule); however, the effect was apparent only at high nymph densities. WBPH survival, development rates and nymph weights all declined at ≥ 30°C. At about the optimal temperature for WBPH (25°C), intraspecific crowding reduced nymph weights. Temperature has little effect on oviposition responses to density, and intraspecific competition between females only weakly counters the effects of optimal temperatures on oviposition in both BPH and WBPH. Meanwhile, the deleterious effects of nymph crowding will counter the direct effects of optimal temperatures on voltinism in BPH and on body size in both BPH and WBPH. The negative effects of crowding on BPH nymphs may be decoupled from resource use at higher temperatures.

The adaptation by planthoppers to feed and develop on resistant rice is a challenge for pest management in Asia. We conducted a series of manipulative experiments with the brown planthopper (Nilaparvata lugens (Stål)) on the resistant rice variety IR62 (BPH3/BPH32 genes) to assess behavioral and bionomic changes in planthoppers exhibiting virulence adaptation. We also examined the potential role of yeast-like symbionts (YLS) in virulence adaptation by assessing progeny fitness (survival × reproduction) following controlled matings between virulent males or females and avirulent males or females, and by manipulating YLS densities in progeny through heat treatment. We found virulence-adapted planthoppers developed faster, grew larger, had adults that survived for longer, had female-biased progeny, and produced more eggs than non-selected planthoppers on the resistant variety. However, feeding capacity—as revealed through honeydew composition—remained inefficient on IR62, even after 20+ generations of exposure to the resistant host. Virulence was derived from both the male and female parents; however, females contributed more than males to progeny virulence. We found that YLS are essential for normal planthopper development and densities are highest in virulent nymphs feeding on the resistant host; however, we found only weak evidence that YLS densities contributed more to virulence. Virulence against IR62 in the brown planthopper, therefore, involves a complex of traits that encompass a series of behavioral, physiological, and genetic mechanisms, some of which are determined only by the female parent.