Aspects of brown planthopper adaptation to resistant rice varieties with the Bph3 gene

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

Figure 10. Arctosa hainan Wang, Li in Review of the wolf spiders from Hainan Island, China (Araneae: Lycosidae)

Figure 10. Arctosa hainan Wang, Li & Zhang, sp. nov., holotype male (A, E–F), paratype male (C–D), paratype female (B, G–H). A. Male habitus, dorsal view; B. Female habitus, dorsal view; C. Left male palp, bulbus, ventral view; D. Same, retrolateral view; E. Left male palp, ventral view; F. Same, retrolateral view; G. Epigyne, ventral view; H. Vulva, dorsal view. Scale bars: A–B = 2.0 mm; C–D, G–H = 0.2 mm; E–F= 0.3 mm.Published as part of Wang, Luyu, Lu, Tian, Cai, Ducheng, Barrion, Alberto Tomas, Heong, Kong-Luen, Li, Shuqiang & Zhang, Zhisheng, 2021, Review of the wolf spiders from Hainan Island, China (Araneae: Lycosidae), pp. 16-74 in Zoological Systematics 46 (1) on page 26, DOI: 10.11865/zs.2021102, http://zenodo.org/record/536634

Figure 39 in Review of the wolf spiders from Hainan Island, China (Araneae: Lycosidae)

Figure 39. Ovia macritchie Lu, Koh, Zhang & Li, 2018, male and female from Hainan. A. Male habitus, dorsal view; B. Female habitus, dorsal view; C. Left male palp, ventral view; D. Same, retrolateral view; E. Left male palp, bulbus, ventral view; F. Same, retrolateral view; G. Epigyne, ventral view; H. Vulva, dorsal view. Scale bars: A–B = 1.0 mm; C–H = 0.2 mm.Published as part of Wang, Luyu, Lu, Tian, Cai, Ducheng, Barrion, Alberto Tomas, Heong, Kong-Luen, Li, Shuqiang & Zhang, Zhisheng, 2021, Review of the wolf spiders from Hainan Island, China (Araneae: Lycosidae), pp. 16-74 in Zoological Systematics 46 (1) on page 46, DOI: 10.11865/zs.2021102, http://zenodo.org/record/536634

Figure 47 in Review of the wolf spiders from Hainan Island, China (Araneae: Lycosidae)

Figure 47. Pardosa sumatrana (Thorell, 1890), male and female from Hainan. A. Male habitus, dorsal view; B. Female habitus, dorsal view; C. Left male palp, bulbus, ventral view; D. Same, retrolateral view; E. Left male palp, ventral view; F. Same, retrolateral view; G. Epigyne, ventral view; H. Same, cleaned the dorsal tissue; I. Vulva, dorsal view. Scale bars: A–B = 1.0 mm; C–D, G–I = 0.2 mm; E–F = 0.5 mm.Published as part of <i>Wang, Luyu, Lu, Tian, Cai, Ducheng, Barrion, Alberto Tomas, Heong, Kong-Luen, Li, Shuqiang & Zhang, Zhisheng, 2021, Review of the wolf spiders from Hainan Island, China (Araneae: Lycosidae), pp. 16-74 in Zoological Systematics 46 (1)</i> on page 53, DOI: 10.11865/zs.2021102, <a href="http://zenodo.org/record/5366340">http://zenodo.org/record/5366340</a&gt

Figure 58 in Review of the wolf spiders from Hainan Island, China (Araneae: Lycosidae)

Figure 58. Trochosa bannaensis Yin & Chen, 1995, male and female from Hainan. A. Male habitus, dorsal view; B. Female habitus, dorsal view; C. Left male palp, bulbus, ventral view; D. Same, retrolateral view; E. Left male palp, ventral view; F. Same, retrolateral view; G. Epigyne, ventral view; H. Vulva, dorsal view. Scale bars: A–B = 1.0 mm; C–H = 0.2 mm.Published as part of <i>Wang, Luyu, Lu, Tian, Cai, Ducheng, Barrion, Alberto Tomas, Heong, Kong-Luen, Li, Shuqiang & Zhang, Zhisheng, 2021, Review of the wolf spiders from Hainan Island, China (Araneae: Lycosidae), pp. 16-74 in Zoological Systematics 46 (1)</i> on page 61, DOI: 10.11865/zs.2021102, <a href="http://zenodo.org/record/5366340">http://zenodo.org/record/5366340</a&gt

Figure 55 in Review of the wolf spiders from Hainan Island, China (Araneae: Lycosidae)

Figure 55. Piratula piratoides (Bösenberg & Strand, 1906), male and female from Hainan. A. Male habitus, dorsal view; B. Female habitus, dorsal view; C. Left male palp, bulbus, ventral view; D. Same, retrolateral view; E. Left male palp, ventral view; F. Same, retrolateral view; G. Epigyne, ventral view; H. Vulva, dorsal view. Scale bars: A–B = 0.5 mm; C–H = 0.1 mm.Published as part of Wang, Luyu, Lu, Tian, Cai, Ducheng, Barrion, Alberto Tomas, Heong, Kong-Luen, Li, Shuqiang & Zhang, Zhisheng, 2021, Review of the wolf spiders from Hainan Island, China (Araneae: Lycosidae), pp. 16-74 in Zoological Systematics 46 (1) on page 59, DOI: 10.11865/zs.2021102, http://zenodo.org/record/536634

Figure 72. Zoica hainan Wang, Li in Review of the wolf spiders from Hainan Island, China (Araneae: Lycosidae)

Figure 72. Zoica hainan Wang, Li & Zhang, sp. nov., holotype male (A, E–F), paratype male (C–D), paratype female (B, G–H). A. Male habitus, dorsal view; B. Female habitus, dorsal view; C. Left male palp, bulbus, ventral view; D. Same, retrolateral view; E. Left male palp, ventral view; F. Same, retrolateral view; G. Epigyne, ventral view; H. Vulva, dorsal view. Scale bars: A–B = 0.2 mm; C–H = 0.1 mm.Published as part of <i>Wang, Luyu, Lu, Tian, Cai, Ducheng, Barrion, Alberto Tomas, Heong, Kong-Luen, Li, Shuqiang & Zhang, Zhisheng, 2021, Review of the wolf spiders from Hainan Island, China (Araneae: Lycosidae), pp. 16-74 in Zoological Systematics 46 (1)</i> on page 70, DOI: 10.11865/zs.2021102, <a href="http://zenodo.org/record/5366340">http://zenodo.org/record/5366340</a&gt

Figure 23. Hippasa holmerae Thorell, 1895 in Review of the wolf spiders from Hainan Island, China (Araneae: Lycosidae)

Figure 23. Hippasa holmerae Thorell, 1895, male and female from Hainan. A. Male habitus, dorsal view; B. Female habitus, dorsal view; C. Left male palp, bulbus, ventral view; D. Same, retrolateral view; E. Left male palp, ventral view; F. Same, retrolateral view; G. Epigyne, ventral view; H. Vulva, dorsal view. Scale bars: A–B = 1.0 mm; C–H = 0.2 mm.Published as part of Wang, Luyu, Lu, Tian, Cai, Ducheng, Barrion, Alberto Tomas, Heong, Kong-Luen, Li, Shuqiang & Zhang, Zhisheng, 2021, Review of the wolf spiders from Hainan Island, China (Araneae: Lycosidae), pp. 16-74 in Zoological Systematics 46 (1) on page 34, DOI: 10.11865/zs.2021102, http://zenodo.org/record/536634

Figure 16. Arctosa tangguoi Wang, Li in Review of the wolf spiders from Hainan Island, China (Araneae: Lycosidae)

Figure 16. Arctosa tangguoi Wang, Li & Zhang, sp. nov., holotype male. A. Male habitus, dorsal view; B. Left male palp, ventral view; C. Same, retrolateral view; D. Right male palp, bulbus, ventral view (overturn); D. Same, retrolateral view (overturn). Scale bars: A = 1.0 mm; B–C = 0.2 mm; D–E = 0.1 mm.Published as part of <i>Wang, Luyu, Lu, Tian, Cai, Ducheng, Barrion, Alberto Tomas, Heong, Kong-Luen, Li, Shuqiang & Zhang, Zhisheng, 2021, Review of the wolf spiders from Hainan Island, China (Araneae: Lycosidae), pp. 16-74 in Zoological Systematics 46 (1)</i> on page 30, DOI: 10.11865/zs.2021102, <a href="http://zenodo.org/record/5366340">http://zenodo.org/record/5366340</a&gt