Pochazia shantungensis (Chou & Lu, 1977)

Figs. 3, 4, 6–9, 11–15.

Ricania shantungensis Chou & Lu, 1977: 315, 317, fig. 11. Type locality: Shandong, Heze (China); Chou et al. 1985: 82; Shen et al. 2007: 116; Kwon et al. 2017: 901; Hizal et al. 2019: 9816–9818, figs. 1–3; Kang et al. 2020: 3813.

Pochazia shantungensis: Rahman et al. 2012: 243–246, figs. 3, 4; Duan & Chen 2020: 211, Pl. I-4, 9, 14, Pl. II-19, 24, Pl. III-29, 34, Pl. IV-39, Pl. V-47, 48, Pl. VI-54, 59, 64, 69; Bourgoin et al. 2020: 271–272, figs. 1–3; Ishida et al. 2020: Pl. 5, 22–23; Schrader 2021: 1–8; Hasegawa et al. 2022: 128, Table 7 [Catalogued]; Stroiński et al. 2022: 276–281, figs. 3–30; Stroiński & Bourgoin 2022: 273; Zhuravleva et al. 2023: 3–4, fig. 4.

Ricania sp.: Choi et al. 2011: 368–371, figs. 1, 2; Choi et al. 2012: 143–147, figs. 1, 2.

Ricania sublimata Jacobi, 1916 [Misidentification]: Park & Jung 2020: 318–322, figs. 1, 2; Park & Jung 2021: 1295–1297, figs. 1B, 2G, 3M, N, 4G, 11.

Diagnosis. The color of the general habitus of P. shantungensis is very similar to P. sublimata (Jacobi, 1916). However, the genitalia exhibit good diagnostic characters, especially the short dorsal process on phallic complex in male (Fig. 8D–F, J–L; Duan & Chen 2020: Pl. IV-39, Pl. V-48), shorter than 1/2 length of phallobase, 450–485 / 915–1235 µm in the present study (vs. longer than 1/2) (Fig. 8M, D, E, Duan & Chen 2020: Pl. IV-38, Pl. V-46); in female, anal segment is 1.0× length of gonoplac in lateral view (vs. 1.4×) (Fig. 9A; Duan & Chen 2020: Pl. VI-53, 54, 68, 69); 0.96–1.19× (1740 / 1460, 1330 / 1350, 1570 / 1620 µm) in the present study. The eastern Asian species P. albomaculata has a similarly white costal spot on brown tegmen (Fig. 16A–C, E–I), but differs from P. shantungensis by the lack of the ventral process in male (Fig. 16K–N) and anal segment being 0.5× length of gonoplac in lateral view in female (Fig. 16O).

Fifth instar nymph of P. shantungensis can be distinguished from that of P. sublimata by the following characters: white mesonotum and wing pads with dark brown patches (vs. uniformly yellowish brown); two pairs of spiny-shaped black spots and dark brown patches on metanotum (vs. no spot and patch) (Figs. 14, 15 A, G, I; Yang 1989: 197–199, fig. 14).

Description. Adult (Figs. 3, 4, 6, 7). See also Duan & Chen (2020, Pl. I-4, 9, 14, Pl. II-19, 24).

Measurements (Japanese specimens): body length (including tegmen): male 11.2–13.0 mm (mean 11.9 mm, n = 11), female 12.4–14.1 mm (mean 13.5 mm, n = 13); frons to distal abdomen length: male 5.6–7.8 mm (mean 6.5 mm, n = 15), female 6.0– 8.1 mm (mean 6.9 mm, n = 8); tegmen length: male 10.2–12.0 mm (mean 11.1 mm, n = 10), female 11.0– 13.5 mm (mean 12.7 mm, n = 15). Male genitalia (n = 8): length and width: anal segment 775 × 415 µm (Fig. 8N); pygofer 925 × 790 µm (Fig. 8M); genital style 920–945 µm × 370 µm (Fig. 7M). Phallic complex (Fig. 8D–F): phallobase 915–1235 µm in length, 350–410 µm in diameter, with three pairs of spinose processes (Fig. 8D–I): dorsal one 450–485 µm in length, lateral view (Fig. 8M, D, E), lateral one 125–290 µm in length, 130–190 µm in width at base (Fig. 8D), ventral one 275–500 µm in length (Fig. 8D, F), aedeagus 840–1165 µm in length (Fig. 8J–L). Female genitalia (n = 5): length and width: anal segment 1570–1780 µm × 1470–1480 µm (Fig. 9F); gonoplac (third valve) 1450–1550 µm × 1300–1580 µm (Fig. 9B, C); gonapophysis VIII 1200 –1700 µm × 1700–1900 µm (process) (Fig. 9H, I); gonapophyses IX 960–1000 µm × 270–290 µm (Fig. 9G); pregenital tergite 1940, 2300 µm × 100–180, 270–400 µm (Fig. 9D). Pregenital sternite 3000, 3700 µm × 250–810, 430–950 µm (Fig. 9E). Egg (Fig. 13A): oval shape, 0.99–1.15 mm in diameter (mean 1.04 mm, n = 4). Nymphs (Figs. 12H–K, 13B–F): white in coloration (Figs. 12H–K, 13B–F, 14) with the following body length: first instar 1.19, 1.28 mm (Fig. 13B, C); second instar 1.32, 1.34 mm (Fig. 13D); third instar 1.64, 1.66 mm (Fig. 13E); fourth instar 2.23, 2.30 mm (Fig. 13F); fifth instar 3.18, 3.28 mm. Fifth instar: head width (including eyes) 1.32, 1.33 mm; anterior wing pad length: 1.74, 1.84 mm; antenna length: 0.82, 0.88 mm.

Male genitalia (Fig. 8). Rahman et al. (2012: 243–245, figs. 3) gave description based on specimens from Korea. See also Duan & Chen (2020, Pl. III-29, 34, Pl. IV-39, Pl. V-47, 48), Stroiński et al. (2022, figs. 13–23), Shichiri & Sakuma (2022, fig. 1C, D), Nakada (2022, fig. 2A) and Harusawa & Miyatake (2023, fig. 1) for specimens from China, Italy and Japan.

Female genitalia (Fig.9). Park &Jung(2020:321,fig.2H–N)described as ‘ Ricania sublimata ’[misidentification] based on a Korean specimen. See also Duan & Chen (2020, Pl. VI-54, 59, 64, 69) and Stroiński et al. (2022, figs. 24–30) for specimens from China and Italy.

Fifth instar nymph (Figs. 14, 15) (n = 6). General color white. Frons and vertex white; laterofrons with 25–26 sensory pits (Fig. 15B). Pronotum with about 28–31 pits on each side (Fig. 15D, E) and two pairs of black spots, smaller ones near body midline, larger oblique ones behind eyes (Figs. 14A, C–E, 15A, D, E). Mesonotum dark brown at distal 3/4 and from front area to 3/4 with a pair of large black spots (Figs. 14B, D, 15F, I). Metanotum dark brown from distal area to the middle (Figs. 14B, C, 15I) with two pairs of spiny shaped black spots at the middle (Fig. 15G). Wing pads brown to dark brown, except base and costal margin: anterior wing pad with a pair of large black spots (Fig. 15F); posterior wing pad without black spot (Fig. 15G), a pair of small black spots on dorsal area of metathorax (Fig. 15I). Legs white, tarsus brown (Fig. 14); fore femur with five or six small teeth laterally at apical half (Fig. 15J); mid femur with seven teeth laterally (Fig. 15K); hind tibia with three teeth laterally, six or seven apically (Fig. 15L, M), first tarsal segment with six apical teeth, second without tooth (Fig. 15P–S). Abdominal tergites white, dark brown at lateral side; tergites IV–V on each lateral side with 4–5 and 5 pits, respectively, one pit of V laterodorsally; tergite VI on each side with 5 wax-pore plates, 2 ventrally, 3 dorsally, each with a sensory pit laterally embedded in plate (Fig. 15T, U). Tergite VII distinctly narrower than preceding ones, incised at posterior margin in dorsal view (Fig. 15A). Wax-pore plates of seventh and eighth segments elongate, of seventh each with 2 pits embedded in ventral half of plate, another one associated with a separate plate (Fig. 15X), of eighth each with 9 pits embedded in lateral margin of plate, single in top, double in remains (Fig. 15W). Ninth abdominal segment with opening caudad, elongate triangular, anal combs large, two pairs of processes at dorsal half, medial one about 3× length of dorsal one (Fig. 15Z).

Material examined. 162 exx. (54 ♂, 108 ♀). Adults: Kyoto Pref.: 1 ♀, Keihanna Commemorative Park, Seika, 6.VII.2022, Y. Inamoto leg., on Morus indica, OMU 114. Nara Pref.: 1 ♂, Takama-cho, Nara City, 2.XI.2022, M. Suzuki leg., OMU 090; 5 ♀, Nara-shi, Saki-chô, Heijô-kyô Palace Site, 34.6930˚N, 135.7905˚E, 21.X.2019, R. Matsumoto (MNHAH); 2 ♀, Ikaruga-cho, 24. VI.2023, W. Nitta leg., OMU 135, 136; 1 ♂, Gojo-shi, Gojô, 34.3488˚N, 135.7052˚E, 22.X.2019, R. Matsumoto (MNHAH). Osaka Pref.: 1 ♀, Furuedai, Suita, 26.X.2022, K. Nakatani leg., on Pueraria montana var. lobata, OMU 088; 1 ♀, Oike, Ibaraki, 29.IX.2022, M. Tanida leg., OMU 089; 2 ♀, Osaka-shi, Sumiyoshi-ku, Nagai Park, 18. VI.2022, R. Matsumoto (MNHAH); 1 ♀, same locality, 14.VII.2022, K. Yamada (MNHAH); 1 ♀, Near Nisanzai-Kofun, Sakai, 3.VII.2023, S. Kobayashi, on Triadica sebifera, 34.548314 ˚N, 135.498961˚E, el. 23.4 m, OMU 146; 2 ♂ 6 ♀, Osaka Metro. Univ., Sakai, 20& 29. VI.2023, S. Kobayashi, on Quercus acutissima, OMU 138–145; 1 ♀, Kanan, Rv. Ishi [Ishi River], 1.X.2015, S. Shiyake leg., SK1032 (OMNH); 1 ♂, Hatsugano, Izumi, IX.2021, Y. Kumura leg., OMU 130; 1 ♀, Okayama-cho, Kishiwada, 22.X.2022, M. Kawai leg. (NHMKCO); 1 ♀, Hashimoto, Kaizuka, 20.IX.2021, K. Temma leg., on Mallotus japonicus, OMU 116. [M. Suzuki leg.]: 1 ♂, Botanical Gardens of Osaka Metro. Univ., Katano, 6.VII.2023, on Lagerstroemia indica, OMU 137; 5 ♂ 17 ♀, Fukakita Ryokuchi, Daito, 19.X.2022, on Ulmus parvifolia, Triadica sebifera and Celtis sinensis, OMU 061–082 (SK 1022 ♂ | OMU 062, SK1023 ♀ | OMU 082); 2 ♂ 1 ♀, Osaka Sangyo Univ., Daito, 22&29.III, 5.IV.2023 em., egg on Cinnamomum camphora, 21.I.2023 (coll.), OUM 127–129; 1 ♀, Chidoribashi Park, Osaka, 11.VII.2023, on Cinnamomum camphora, OMU 174; 1 ♂, Kire, Osaka, 16. VI.2023, on Quercus glauca, OMU 131; 1 ♂ 2 ♀, Houzen-ji, Kashiwara, 15.X.2022, on Olea europaea and Acer palmatum, OMU 084–086; 1 ♀, Ogata, Kashiwara, 7.XII.2022, on Cinnamomum camphora, OMU 087; 1 ♀, Mozu-cho, Sakai, 15.X.2022, on Quercus glateragauca, OMU 039; 12 ♂ 26 ♀, Osaka Metro. Univ., Sakai, 14& 18.X.2022, Adult on Acca sellowiana, Camellia sasanqua and Ligustrum vulgare, OMU 001–038 (SK 1020 ♂ | OMU 001, SK1021 ♀ | OMU 006, SK1029 ♀ | OMU 0034, OMU 010, 011 are deposited in MNHAH); 1 ♂ 1 ♀, Hata, Sakai, 18.X.2022, on Pueraria montana var. lobata, OMU 040, 041; 6 ♂ 13 ♀, Sakai Nature Forest, Sakai, 16& 28.X.2022, on Ailanthus altissima and Ilex serrata, OMU 042–060 (SK 1024 ♂ (wing)| OMU 048, SK1025 ♂ (wing) | OMU 049, SK1026 ♀ (wing) | OMU 052, SK1027 (wing)/SK1030 ♀ | OMU 058, OMU 043, 050, 059, 060 deposited in MNHAH); 2 ♂ 3 ♀, Hachigamineji, Sakai, 17.VII.2023, on Ligustrum lucidum, Quercus glauca and Q. serrata, OMU 182–186; 1 ♂, Fuseya, Izumi, 8.XI.2022, on Ilex rotunda, OMU 083. [Y. Yamuchi leg.]: 1 ♀, Ote-cho, Ibaraki, 31.X.2022, adult on Citreae sp., OMU 109; 2 ♀, Hattori Ryokuchi, Toyonaka, 29.X.2022, on Camellia japonica and Ligustrum lucidum, OMU 095, 096; 3 ♂ 4 ♀, Fujisaka-minami-machi, Hirakata, 30.X.2022, on Solidago altissima, Camellia sasanqua, Quercus glauca and Celtis sinensis, OMU 097–103; 1 ♂ 2 ♀, Neyagawa Park, Neyagawa, 30.X.2022, on Osmanthus fragrans var. aurantiacus and Camellia sasanqua, OMU 104–106; 1 ♀, Minamino, Shijonawate, 30.X.2022, on Camellia sasanqua, OMU 107; 1 ♀, Nozaki, Daito, 30.X.2022, on Ligustrum lucidum, OMU 108; 1 ♂ 3 ♀, Shirokita Wando, Asahi, Osaka, 23.X.2022, on Solidago altissima, Triadica sebifera and Ligustrum lucidum, OMU 091– 094. Wakayama Pref.: 2 ♂, Nakado, Hashimoto, 24. VI.2023, N. Hiraiwa leg., on Rubus fruticosus, OMU 132, 133; 2 ♀, Mouko, Kainan, 2& 15.VII.2023, M. Suzuki leg., on Solidago altissima, OMU 134, 175. Hyogo Pref.: 2 ♂, Akashi-koen Park, south of Fujimiike, Akashi, 14.IX.2022, A. Ichikawa (MNHAH). Hiroshima Pref.: 2 ♀, Sannomaru-cho, Fukuyama, 27.X.2022, K. Iida & T. Ueda leg., on “Sunflower” [= Helianthus sp.], OMU 112–113.

Kumamoto Pref.: 8 ♂ 1 ♀, Kurokami, Kumamoto, 32.8176934˚N, 130.7211841˚E, 20& 29–30.IX.2021 (Light), R. Kuwahara leg., preserved in 70 % ethanol, specimen vial ID CS024-51 (2 ♂), 52 (1 ♂), 53 (5 ♂), CS037-35 (1 ♀) (MNHAH).

Fifth instar nymphs: 17 exx. Osaka Pref.: [M. Suzuki leg.]: 1 exuvum, Fukakita Ryokuchi, Daito, 27.X– 1.XI.2022 em., on Akebia quinate, 19.X.2022, OMU 115 (exuvum)/ OMU 061(emerged adult); 3 exx., Osaka Sangyo University, Daito, 21.I.2023, from eggs on Cinnamomum camphora, Zelkova serrata, and Cornus florida, OMU 117–119; 6 exx., Sakai Nature Forest, Sakai, 25.II.2023, from eggs on Deutzia crenata, OMU 120–125; 1 ex., Hachigamine-ji, Sakai, 25.II.2023, from eggs on Castanea crenata, OMU 126. Wakayama Pref.: 6 exx., Nakado, Hashimoto, 24. VI.2023, N. Hiraiwa leg., on Rubus fruticosus, OMU 155.

Distribution. Japan: Honshu: Chiba (Ban 2023), Saitama (Osada 2023, Nozawa et al. 2023), Tokyo (Han 2023, Miyazaki 2023), Gunma (Kanasugi 2024), Kanagawa (Shichiri & Sakuma 2022), Shizuoka (Nakada 2022), Gifu (Kawakami 2022), Aichi (Yoshitsuru 2023), Mie (Nishikawa 2024), Kyoto (Kawai 2024), Nara (Ito 2023, Tominaga 2023), Osaka, Hyogo, Wakayama (Ishida et al. 2020), Okayama (Suenaga 2023), Hiroshima (new record) Prefectures, Kyushu: Kumamoto Prefecture (Takeuchi 2023); China: Shandong (Chou & Lu, 1977), Zhejiang (Chou et al. 1985); Korea (Rahman et al. 2012); Russia (Zhuravleva et al. 2023); Turkey (Hizal et al. 2019); the Netherlands (EPPO RS 2023); Germany (Schrader 2021); Italy (Stroiński et al. 2022); France (Bourgoin et al. 2020).

Countries underlined denote type-locality countries.

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Host plants in Japan. (Table 3, fig. 2). Aquifoliaceae: Ilex rotunda Thunb.; Lauraceae: Cinnamomum camphora (L.) J.Presl; Oleaceae: Ligustrum lucidum W.T.Aiton and L. vulgare L., Osmanthus fragrans var. aurantiacus Makino; Sapindaceae: Acer palmatum Thunb.; Simaroubaceae: Ailanthus altissima (Mill.) Swingle; Theaceae: Camellia japonica L., C. sasanqua Thunb.; Ulmaceae: Ulmus parvifolia Jacq., Zelkova serrata (Thunb.) Makino etc.

Biology (Figs. 11–14). This species has two generations per year in Zhejiang, China (Li et al. 2006; Shen et al. 2007) and one generation per year in Korea (Choi et al. 2012, fig. 3 as Ricania sp.; Baek et al. 2019); overwintering takes place in the egg stage on tree branches, nymphs emerged in April and July, adults in June and September (Shen et al. 2007). In Europe, the adults were collected in June and around October (Bourgoin et al. 2020; Stroiński et al. 2022). In Japan, this species has two generations per year and overwinters in the egg stage on tree branches in Osaka Prefecture. The adults were observed in June, July and between September to December. In Sakai, the adults have been observed around agricultural fields and at forest edge in July, September and October from 2015 to 2022 (Fig. 11A–C). In Kumamoto, many adults were observed to attract around lights at night. The females laid white eggs in the branches and stems of host plants; the tip of egg masses was covered with white waxy filaments (Fig. 12A–E); ca. 2–5 cm in the length; old egg masses lacked the filaments in winter (Fig. 12F, G).

Remarks. This species has black tegmina in ground color, covered with a thin white and distinct brown wax layer, but old and/or dried pinned specimen often lack wax layer and look dark (Fig. 3C, F, H, K). Emerged adults were at first general white (Fig. 11H–I), and become general black which red eyes in coloration (Fig. 11J, K), and covered with brown wax layer in three days (Fig. 11L). We identified Ricaniidae gen. sp. 1–3 of Yoshitsuru (2023) from Aichi Prefecture as P. shantungensis by the rounded triangular-shaped costal spot on forewing.

In the present study, this species was collected on more than 32 plant families, 61 species, particularly on Ligustrum and Camellia species, and Cinnamomum camphora (Table 3). In Osaka Prefecture, the roadside tree, Cinnamomum camphora and the hedge plant, Ligustrum vulgare had the highest density of this species (> 50 adults and egg masses per one collecting); the egg masses were found in particular on Camellia sasanqua and C. japonica trees in winter. Several authors observed egg-laying females or on white waxy filaments of egg mass on several trees, e.g. Fagaceae, Lauraceae, Magnoliaceae, Moraceae (Shichiri & Sakuma 2022, 94; Miyazaki 2023, 15; Nozawa et al. 2023, 67). In private and public gardens, the species was reported on several host plants (information from the internet), e.g., Celastraceae: Euonymus japonicus; Fagaceae: Quercus myrsinaefolia; Rosaceae: Spiraea thunbergii; Rubiaceae: Gardenia jasminoides; Sapindaceae: Maple (Acer); sometimes with> 10 adults and/or nymphs per shoot or tree.

One female adult was attended by three arboreal ants, Camponotus vitiosus Smith, 1874 (Formicidae) in Keihanna Commemorative Park, Kyoto Prefecture (Fig. 11E); One of the ants fed on the planthopper excreted honeydew. Another female adult was also attended by a Camponotus ant in Sakai, Osaka Prefecture.