Pilophorus Hahn, 1826

Yasunaga, Tomohide, Duwal, Ram Keshari & Nakatani, Yukinobu, 2020, Reclassification of the plant bug genus Pilophorus in Japan and key to the genera and species of Japanese Pilophorini (Hemiptera: Heteroptera: Miridae Phylinae), Zootaxa 4942 (1), pp. 1-40 : 3-4

Pilophorus Hahn, 1826 (17 spp.)

P. choii Josifov, 1987 ( Fig. 1A )— Japan ( Hokkaido, Honshu, Kyushu); Korea, Russian Primorsky Territory.—Deciduous Quercus spp. ( Fagaceae ) ( Kerzhner, 1988; Yamamoto & Yasunaga, 2020).

P. erraticus Linnavuori, 1962 ( Figs 2 A–B)— Japan ( Hokkaido, Honshu, Shikoku, Kyushu, Koshiki Island); Korea, Russian Primorsky Territory.—Various deciduous broadleaf trees, Alnus spp. ( Betulaceae ), Maackia amurensis Rupr. & Maxim. ( Fabaceae ), Salix spp. ( Salicaceae ), Sambucus sieboldiana L. ( Adoxaceae ), Ulmus davidiana Planch. , Zelkova serrata (Thunb.) Makino ( Ulmaceae ) (Urayama et al. 2019; Yamamoto & Yasunaga 2020); both adults and immature forms were also found from a subtropical broadleaf, Ficus superba (Miq.) Miq. ( Moraceae ) ( Fukuda et al., 2020).

P. hyotan n. sp. ( Figs 3 A–F, 4A–H)— Japan (Honshu, Shikoku, Kyushu, Hachijo and Tsushima Islands, Ryukyus: Amami- Oshima, Okinawa, Ishigaki and Iriomote Islands), Korea, Nepal, Taiwan, Vietnam.—Various herbs, vegetables and flowers of dicot angiosperms (see below Biology section in description of new species).

P. lucidus Linnavuori, 1962 ( Fig. 2F )— Japan ( Hokkaido, Honshu, Shikoku, Kyushu, Tsushima Island); Korea, Russian Primorsky Territory.—Deciduous Quercus spp.; in some coastal zones of southwestern Japan, Castanopsis sieboldii (Makino) Hatus. ( Fagaceae ).

P. maeharai Yasunaga & Duwal, 2016 ( Fig. 1B )— Japan (Honshu: Tochigi Pref.).—Tree trunk of Prunus jamasakura Sieb. ( Rosaceae ).

P. miyamotoi Linnavuori, 1961 ( Figs 6A , 9O , 14 P–Q)— Japan ( Hokkaido, Honshu, Izu Islands, Shikoku, Kyushu, Tsushima Island); Korea, Russian Primorsky Territory.— Pinus densiflora Sieb. & Zucc. and P. koraiensis Sieb & Zucc. ( Pinaceae ) as main breeding hosts; sometimes found on P. thunbergii Parl. , introduced European or North American species of Pinus , or rarely Larix leptolepis (Sieb. et Zucc.) Gordon in northern Japan.

P. nakatanii Yasunaga & Duwal, 2016 ( Fig. 2E )— Japan (Ryukyus: Okinawa, Ishigaki & Iriomote Islands).— Pinus luchuensis Mayr ( Pinaceae ).

P. niger Poppius, 1914 ( Figs 1C , 20 A–C)— Japan ( Hokkaido, Honshu, Shikoku, Kyushu), China, Korea, Mongolia, Russian Primorsky Territory.— Unknown; adult individuals were collected from Cinnamomum yabunikkei H.Ohba ( Lauraceae ) or Lespedeza sp. ( Fabaceae ) ( Yamamoto & Yasunaga, 2020).

P. okamotoi Miyamoto & Lee, 1966 ( Figs 5C ̅D)— Japan (Tsushima Island); Korea ( Jeju Island), Russian Primorsky Territory.— Artemisia spp. ( Asteraceae ).

P. pseudoperplexus Josifov, 1987 ( Figs 2 C–D)— Japan ( Hokkaido, Honshu, Shikoku, Kyushu); Korea, Russian Primorsky Territory.— Quercus accutissima Carruth. , Q. serrata Murray ( Fagaceae ).

P. pullulus Poppius, 1914 sp. rev. ( Figs 1 D–G)— Japan (Ryukyus: Amami-Oshima Island); Taiwan—Unknown.

P. satoyamanus n. sp. ( Figs 5 A–B) — Japan (Honshu, Shikoku)— Artemisia spp.

P. setulosellus n. sp. ( Fig. 5E )— Japan (Honshu southwest of Kansai area, Shikoku)— Artemisia spp. ( Asteraceae ) but sometimes found on various dicot herbs or shrubs.

P. setulosus Horváth, 1905 ( Fig. 5F )— Japan ( Hokkaido, Honshu, Shikoku, Kyushu, and Rishiri, Rebun, Izu and Ogasawara Islands: records from SW Honshu, Shikoku, Kyushu and Izu and Ogasawara Islands need verification), Russian Far East (south Sakhalin); records from China (Inner Mongolian Prov.) ( Zou, 1989) and Korean Peninsula (cf. Duwal et al. 2014) need further verification.—Various deciduous broadleaf trees, herbs and shrubs; immature forms were found from Alnus spp., Artemisia sp., Salix spp. and Ulmus spp. ( Yasunaga 2001; Fukuda et al. 2020).

P. tagoi Yasunaga & Duwal, 2016 ( Figs 6C , 20D )— Japan (Honshu: Kanto area). — Cryptomeria japonica (Thunb. ex L.f.) D. Don. ( Cupressaceae ).

P. typicus (Distant, 1909) ( Figs 3 G–I, 4I–J)— Japan † (Honshu, Shikoku, Kyushu, and Koshiki and Tsushima Islands, Ryukyus); SE China, India, whole Indochina, Indonesia ( Java, Sumatra, Irian Jaya), S Korea †, E & W Malaysia, Nepal †, Philippines, Singapore, Sri Lanka, Taiwan; † previous records in need of critical verification as P. hyotan n. sp. was found to co-occur or regionally predominate—Associated with various herbs and vegetables (cucumber, eggplant, red pepper, pimento, pumpkin, including those in greenhouses); immature forms found from Acanthaceae , Asteraceae , Convolvulaceae , Ericaceae , Euphorbiaceae , Fabaceae , Laminaceae, Rutaceae , Solanaceae , Urticaceae , etc. ( Fukuda et al. 2020); often observed to prey on thrips, whiteflies, leafhoppers and spider mites ( Ito et al., 2011; present observation, Fig. 4 ); all developmental stages can be bred with a kalanchoe, Kalanchoe daigremontiana (cf. Fukuda et al., 2020).

P. varidicornis Kerzhner, 1977 ( Fig. 6B )— Japan ( Hokkaido incl. Kunashiri Island); Russia ( Sakhalin).— Picea spp. ( Pinaceae ); expanding the habitat to urbanized and residential zones and sometimes abundant on introduced European spruces for landscaping ( Yasunaga, 2001).

Figures

FIGURE 1. Habitus images of Pilophorus spp. A. P. choii ³. B. P. maeharai, holotype ³. C. P. niger ♀. D–E. P. pullulus, holotype ♀ (Taiwan). F–G. P. pullulus ³ (Amami-Oshima Is., Japan).

FIGURE 2. Habitus images of Pilophorus spp., live individuals. A. P. erraticus ³. B. P. erraticus ♀. C–D. P. pseudoperplexus ³. E. P. nakatanii, holotype ³. F. P. lucidus ♀.

FIGURE 3. Habitus images of Pilophorus hyotan n. sp. (A–F) and P. typicus (G–I), live adult individuals. A. ³. B. ♀. C. ♀, ventral view. D. ♀, ventral view. E. ♀, praying on an aphid. F. ♀, praying on a leafhopper nymph. G. ³, from Kathmandu, Nepal. H. ♀, Surin, Thailand. I. ♀, Nagasaki, Japan, ventral view.

FIGURE 4. Life cycle and immature stages of Pilophorus hyotan n. sp. (A–H) and P. typicus (I–J), laboratory observations in Nagasaki, Japan. A. Mating pair (³ left). B. Eggs (exposed opercula) oviposited into border part between stem and root of Artemisia sp. C. First instar nymph, just after hatching. D. Same, before molting to 2nd instar. E. Second instar nymphs. F. Third instar nymph. G. Fourth instar nymph with 5th (left). H. Fifth instar nymph with its exuvia. I. Fourth instar nymph. J. Fifth instar nymph (feeding on brine-shrimp eggs) [MTNR = metanotal ridge].

FIGURE 5. Habitus images of Pilophorus spp., live adult individuals. A–B. P. satoyamanus n. sp. ³ (images courtesy of M. Takai). C–D. P. okamotoi ³. E. P. setulosellus n. sp. ♀ (courtesy of M. Takai). F. P. setulosus ♀.

FIGURE 6. Habitus images of conifer-inhabiting Pilophorus spp. (A–C) and Pherolepis spp. (D–H), live adults and 5th instar nymph (G). A. Pilophorus miyamotoi ³. B. P. validicornis ♀ (image courtesy of M. Nozawa). C. P. tagoi ♀ (courtesy of M. Nozawa). D. Pherolepis hizenicus n. sp., ³. E. Same, lateral view. F. Same, ♀. G. Same, 5th instar nymph, ³. H. Pherolepis fasciatus ♀.

FIGURE 9. Male genitalia of Pilophorus spp. A–E, I–K. P. hyotan n. sp. (from Nagasaki, Japan: A–C, K; Solkhumbu, Nepal: D–E, I–J). F–H, L–N. P. typicus (Nakhon Nayok, Thailand: F–G; India: H, L–M; Nagasaki: N). O. P. miyamotoi (Kochi Pref., Japan). P–S. P. pullulus (Amami-Oshima Is.). A– B, D, F–G, O–P. Endosoma (vesica). C, E, H, S. Phallotheca. I–N, Q–R. Parameres.

FIGURE 14. Scanning electron micrographs for Japanese Pilophorus spp., males (except for E, L, P: ♀). A–D. P. choii. E–K. P. erraticus. L–N. P. pseudoperplexus. O–Q. P. miyamotoi. A, E. Left-dorsolateral habitus. B, I, N. Apex of pygophore, left lateral view. C–D, J–K, Q. Endosoma (vesica).A, E. Dorsal habitus. F. Thoracic pleura, left lateral view. H, L, M, O, P. Lateral habitus. G. Pygophore, dorsal view.

FIGURE 20. Scanning electron micrographs for Pilophorus niger (A–C), P. tagoi (D), and Pherolepis hizenicus n. sp. (E–O). A, D, G. Left lateral habitus. B, I. Metatarsus. Right lateral habitus. C, L. Endosoma. E. Dorsal habitus. F. Left-dorsolateral habitus. H. Thoracic pleura, left lateral view. J. Pygophore, dorsal view. K. Left paramere. M. Bursa copulatrix, dorsal view. N–O. Posterior wall of bursae.