Cycadaceae: Cycas revoluta Thunb. (Fig. 1B–F) (Kato 2001). The wood tissues of Toddalia asiatica (L.) and Aucuba japonica Thunb. are used as hosts (Coleopterological Society of Japan 1984).

Figure 1. 

Two cerambycid species associated with Cycas revouta: Mimectatina meridiana ohirai (A–F) and Sybra flavostriata flavostriata (G–I). A, G adult habitus B flowering cycad C flowering megasporophyll D cross section of mined megasporophyll E mine of megasporophyll F pupa in the mine H oviposition scars on leaf stalk I larva mining the leaf stalk.

Linear or linear-blotch mines in the megasporophyll of female cycad plants (Fig. 1E). The larva sometimes penetrates the testa of the cycad seed. Pupation occurs within the mine on the leaf stalk of the megasporophyll (Fig. 1D, F). Frass is granular and deposited within the mine.

• 25 adults, Hedo, Kunigami, Okinawa Is., Okinawa Pref., 1-II-1998 (collected as larva on Cycas revoluta), emerged on 3-III–20-VI-1998; • 7 adults, Hedo, Kunigami, Okinawa Is., Okinawa Pref., 10-III-1997 (as larva on C. revoluta), emerged on 25-IV–20-VI-1997; 2 adults, Hedo, Kunigami, Okinawa Is., Okinawa Pref., 19-IV-2000 (as larva on C. revoluta), emerged on ?-V-2000; • 8 adults, Angyaba, Kakeroma Is., Setouchi, Kagoshima Pref., 10-V-2001 (as larva on C. revoluta), emerged on 12–28-VI-2001.

Cycadaceae: Cycas revoluta (Fig. 1H, I) (Kato 2001). Wood of Ficus superba (Miq.), Ficus erecta Thunb., Pittosporum tobira (Thunb.), and Boehmeria biloba Wedd. are also used as larval hosts (Coleopterological Society of Japan 1984).

Linear mine in a leaf stalk. The adult female bites the lower surface of the leaf stalk and inserts eggs into the scar (Fig. 1H). The larva mines the woody leaf stalk linearly, but does not enter leaflets. The fully grown larva pupates in the mine (Fig. 1I).

• 5 adults, Haneji, Nago, Okinawa Is., Okinawa Pref., 1-II-1998 (as larva on Cycas revoluta), emerged on ?-V-1998; 2 adults, Hedo, Kunigami, Okinawa Is., Okinawa Pref., 1-II-1998 (as larva on Cycas revoluta), emerged on 20-III-1998.

Celastraceae: Celastrus orbiculatus Thunb., Euonymus alatus (Thunb.), E. fortunei (Turcz.), E. japonicus Thunb., E. macropterus Rupr., E. melananthus (Thunb.), E. oxyphyllus Miq., E. tricarpus Koidz., E. sieboldianus Blume. Tripterygium regelii is also recorded as a host by Takemoto (2019).

Full-depth linear-blotch mine on newly-opened leaf (Fig. 2O–Q). The mine often covers the whole area of the leaf causing deformation of the leaf, and the larva moves to another unmined leaf. Frass is thread-like and meandering, deposited along the center of the mine. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

Figure 2. 

Habitus and feeding scars of adult beetles and leaf mines of eight Zeugophora species A, B Z. nigricolis C–E Z. unifasciata F–I Z. chujoi J–M Z. flavonotata N–Q Z. annulata R, S Z. varipes T–V Z. hozumii W–Z Z. japonica. Host plants A, B Euonymus sieboldianus at Hirayu, Gifu Pref. C–E E. sieboldianus at Sugadaira, Nagano Pref. F–I E. fortunei at Nagao-toge, Shizuoka Pref. J–L E. tanakae at Yutsun, Iriomote Is., Okinawa Pref. M E. japonicus at Hoshidate, Iriomote Is., Okinawa Pref. N, O E. alatus at Kiyosato, Yamanashi Pref. P E. melananthus at Mt. Tara, Nagasaki Pref. Q E. fortunei at Mt. Myôjô, Niigata Pref. R, S Symplocos sawafutagi at Iyari, Nagano Pref. T–V Salix cardiophylla var. urbaniana at Kitazawa-toge, Nagano Pref. W–Z Populus suaveolens at Sôunkyô, Hokkaido.

• 4 adults, Iyari, Ômachi, Nagano Pref., 5-V-2016 (as larva on Euonymus alatus), emerged on 27-V-2016 (Fig. 2N, O); 5 adults, Saroma Lake, Yûbetsu, Hokkaidô, 5-VI-2016 (as larva on E. sieboldianus), emerged on 29–30-VI-2016; 3 adults, Mt. Tara, Isahaya, Nagasaki Pref., 19-IX-2017 (as larva on E. melananthus), emerged on 23-V-2017 (Fig. 2P); • 2 adults, Mt. Myôjô, Niigata Pref., Aoi-ku, Shizuoka Pref., 9-V-2018 (as larva on E. fortunei), emerged on 10-VI-2018 (Fig. 2Q); • 1 adult, Mt. Teine, Sapporo, Hokkaido, 6-VI-2016 (as larva on E. tricarpus), emerged on 25-VI-2016; • 2 adults, Ashiu, Nantan, Kyoto Pref., 5-V-1992 (as larva on E. oxyphyllus), emerged on ?-V-1992; • 1 adult, Hidakatsu, Tsushima Is., Nagasaki Pref., 19-IV-2009 (as larva on E. japonicus), emerged on 18-V-2009; • 1 adult and many leaf mines, Mt. Byôbu, Kizukuri, Aomori Pref., 14-V-1993 on E. macropterus; • 3 adults, Misakubo Dam, Hamamatsu, Shizuoka Pref., 10-V-2000 (as larva on Celastrus orbiculatus collected by T. Kato), emerged on 7-VI-2000.

Celastraceae: Euonymus fortunei (Turcz.).

Full-depth linear-blotch mine on the leaf blade and the midrib of a newly-opened leaf (Fig. 2G–I). The egg is laid in the midrib and the hatched larva enters the midrib and bores into the petiole, leading to leaf abscission from the shoot at the petiole base. Following the leaf fall, the mine diverges from the midrib and expands in the leaf blade, forming a blotch mine. Frass is granular and scattered throughout the mine. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

• 4 adults, Nagao-tôge, Gotenba, Shizuoka Pref., 15-V-2018 (as larva on Euonymus fortunei), emerged on 18-VI-2018 (Fig. 2F–I).

Celastraceae: Euonymus carnosus Maxim., E. japonicus Thunb.

Full-depth linear mine on the newly-opened leaf (Fig. 2K–M). The egg is laid near the leaf margin and the hatched larva mines along the leaf margin, slowly expanding the mine. Frass is thread-like, deposited somewhat to one side of the center. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

• 2 adults, Yutsun, Iriomote Is., Yaeyama, Okinawa Pref., 24-III-2022 (as larva on Euonymus carnosus), emerged on 16–18-IV-2022 (Fig. 2J–L); 3 adults, Todomari, Iriomote Is., Yaeyama, Okinawa Pref., 27-III-2018 (as larva on E. carnosus), emerged on 16–18-IV-2018; 14 adults, Hoshidate, Iriomote Is., Yaeyama, Okinawa Pref., 6-III-2019 (as larva on E. japonicus), emerged on 3–6-IV-2019 (Fig. 2M).

Celastraceae: Euonymus sieboldianus Blume.

Upper-layer ophiogenous blotch mine on the mature leaf (Fig. 2D, E). Within the blotch, the larval trajectory itself is linear and meandering compactly. The egg is laid along the basal margin of the leaf and the hatched larva mines along the leaf margin, abruptly expanding the mine. Within the blotch, the larval trajectory itself is linear and meandering compactly. Frass is granular and deposited along the meandering trajectory. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

• 2 adults, Sugadaira, Ueda, Nagano Pref., 18-VI-2017 (as larva on Euonymus sieboldianus), emerged on 20–28-VII-2017 (Fig. 2C–E).

Celastraceae: Euonymus sieboldianus.

Upper-layer linear mine on mature leaf (Fig. 2B). The egg is laid in the leaf blade and the hatched larva construct slightly meandering mine. Width of the mine is slightly wider than the width of the larva. Frass is thread-like, deposited along the meandering trajectory. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

• 5 adults, Hirayu-tôge, Takayama, Gifu Pref., 6-IX-2019 (as larva on Euonymus sieboldianus), emerged on 6–7-X-2019 (Fig. 2A, B); • 1 adult, Toyohara, Nasu, Tochigi Pref., 2-VII-2022 (as larva on E. sieboldianus), emerged on 3-VIII-2022.

Symplocaceae: Symplocos coreana (H. Lev.), S. sawafutagi Nagam.

Full-depth linear-blotch mine on young leaf (Fig. 2S). The egg is laid in the leaf blade, and the hatched larva constructs linear mine along the leaf margin, then gradually expands the mine. The mine is wider than the width of the larva. Frass is granular, scattered throughout the mine. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

• 1 adult and 5 leaf mines, Iyari, Ômachi, Nagano Pref., 5-V-2016 on Symplocos sawafutagi (Fig. 2R, S); 6 adults, Omotsubo, Okayama Pref., 27-V-1993 (as larva on Symplocos coreana), emerged on 20–24-VI-1993.

Salicaceae: Salix cardiophylla Trautv. et C. A. Mey. Although Takemoto (2019) lists Salix caprea L. as a host plant, this record is not confirmed.

Black upper-layer linear-blotch mine on mature leaf (Fig. 2V). The egg is laid in the leaf blade, and the hatched larva constructs a linear-blotch mine, the upper surface of which turns black. Frass is granular, scattered throughout the mine. The fully grown larva exits the mined leaf, falls to the ground, pupates underground, and hibernates as a pupa.

• 8 adults and many leaf mines, Kitazawa Tôge, Ina, Nagano Pref., 30-VII-2016, feeding lower surface of leaf of Salix cardiophylla (Fig. 2T–V); • 1 adult, Azusa-gawa, Matsumoto, Nagano Pref., 24-X-2020 (as larva on Salix cardiophylla), emerged on 22-III-2021.

Salicaceae: Populus suaveolens Fisch.

Black upper-layer blotch mine on mature leaf, sometimes gregarious (Fig. 2X–Z). The egg is laid along the leaf margin, and the hatched larva constructs a blotch mine, the upper surface of which turns black. Frass is granular, scattered throughout the mine. The fully grown larva exits the mined leaf, falls to the ground, pupates underground.

• 1 adult, Rekifune Nakanokawa, Taiki, Tokachi, Hokkaidô, 23-VI-2017 (as larva on Populus suaveolens), emerged on 21-VIII-2017 (Fig. 2W–Z); • 2 adults, Rubeshibe, Kitami, Hokkaidô, 25-VI-2017 (as larva on P. suaveolens), emerged on 21-VIII-2017.

Salicaceae: Populus suaveolens (Takemoto 2019). Leaf mine has not yet been reported.

There is no recent collection of this species, and its host plant and mine are not known (Takemoto 2019).

Brassicaceae: Draba nemorosa L. The host plant grows on levee of traditional rice fields and upland fields in Central Honshu (Fig. 3C).

Figure 3. 

Habitus of adults and leaf mines of two Phyllotreta species A–I P. ezoensis J–M P. shirahatai. Host plants A–I Draba nemorosa at Kiyosato, Yamanashi Pref. J–M Cardamine appendiculata at Eniwa (J, K) and Tobai (L, M), Hokkaido.

Full-depth linear mine of leaf blade, midrib, petiole, and shoot (Fig. 3D–I). The egg is laid on the leaf, and the hatched larva mines toward the midrib, and reenters midrib/leaf blade. Larvae repeats mining leaf and mining midrib, petiole, and/or shoot, often by exiting its mine and establishing a new mine. Frass is granular, deposited linearly along midline of the mine. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground. The adult emerges ~ 2 weeks after pupation, and varies in elytral pattern (Fig. 3A, B).

• 6 adults, Matsubara-Lake, Koumi, Nagano Pref. 21-V-2021 (as larva on Draba nemorosa), emerged on 4-VI-2021 (Fig. 3A–E); • 15 adults, Kiyosato, Hokuto, Yamanashi Pref., 4-V-2022 (as larva on D. nemorosa), emerged on 17–20-V-2022 (Fig. 3F–I).

Brassicaceae: Cardamine leucantha (Tausch.).

Full-depth linear-blotch mine on mature leaf (Fig. 3K–M). Frass is granular, scattered throughout the mine. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

• 1 adult and several leaf mines on Cardamine leucantha, Eniwa Park, Eniwa, Hokkaidô, 21-VII-2020 (Fig. 3J, K); several leaf mines, Tôbai, Nemuro, Hokkaidô, 2-VIII-2023 on C. leucantha (Fig. 3L, M).

Whereas six collected specimens were tentatively identified as this species, more specimens and further taxonomic studies are necessary.

Plantaginaceae: Pennellianthus frutescens (Lamb.). Host plants of Longitarsus holsaticus are reported to be Veronica spp. by Kimoto and Takizawa (1993).

Full-depth linear or radiate mine in young leaf (Fig. 4F–G). Frass is deposited compactly in a few parts of the mine, and sometimes discharged from the mine through holes perforated along the mine edge. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

Figure 4. 

Habitus of adults and leaf mines of two species belonging to two Alticini genera, Longitausus and Dibolia A–G Longitarsus holsaticus H–L Dibolia japonica. Host plants A–G Pennellianthus frutescens at Mt. Tarumae, Hokkaido H–L Stachys aspera var. hispidula at Lake Shoji-ko, Yamanashi Pref.

• 6 adults, Mt. Tarumai, Chitose, Hokkaidô, 4-VII-2010 (as larva on Pennellianthus frutescens), emerged on 17-VIII-2010 (Fig. 4A–G).

Lamiaceae: Stachys aspera Michx.

Full-depth linear-blotch mine on mature leaf (Fig. 4J–L). Frass is thread-like, deposited along trajectory in the mine. The fully grown larva exits the mined leaf, falls to the ground, pupates underground.

• 1 adult, Shôji-ko Lake, Fuji-kawaguchiko, Yamanashi Pref., 4-VII-2010 (as larva on Stachys aspera), emerged on 3-VIII-2010 (Fig. 4H–L).

Polygonaceae: Rumex japonicus Houtt., Polygonum aviculare L. Rumex acetosa L. is also recorded as a host plant (Kimoto and Takizawa 1993).

Full-depth linear mine occurs on mature leaf (Fig. 5D–F, I). The larva alternates mining the leaf blade and mining the midrib, the petiole, and the shoot, and often relocates its mine. Frass is granular, deposited linearly along middle line of the mine. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

Figure 5. 

Habitus of adults, habitat, and leaf mines of two Mantura species A–I M. clavareaui J–L M. fulvipes. Host plants A–F Rumex japonicus at Notojima Island, Ishikawa Pref. G–I Polygonum aviculare at Mihogaseki, Shimane Pref. J–L Oxalis corniculata at Tamanoura, Fukue Is., Nagasaki Pref.

• 3 adults, Kôda, Notojima Is., Nanao, Ishikawa Pref. 2-V-2023 (as larva on Rumex japonicus), emerged on 22-V-2023 (Fig. 5A–F); • 21 adults, Kamematsu, Shivani, Kumamoto Pref., 28-IV-2017 (as larva on R. japonicus), emerged on 13–14-V-2017; • 14 adults, Mihonoseki, Matsue, Shimane Pref., 15-V-2006 (as larva on Polygonum aviculare), emerged on 31-V–3-VI-2006 (Fig. 5G–I).

Oxalidaceae: Oxalis corniculata L.

Full-depth linear mine on mature leaf (Fig. 5L). The egg is laid along the leaf margin, and hatched larva mines along the leaf margin, gradually expanding the mine. Width of the mine is wider than that of larva. Frass is thread-like, deposited along middle line of the mine. The fully grown larva exits the mined leaf, falls to the ground, pupates underground.

• 19 adults, Tamanoura, Fukue Is. Goto, Nagasaki Pref. 9-X-1998 (as larva on Oxalis corniculata), emerged on 22–26-X-1998 (Fig. 5J–L).

Host plant has not been reported (Hayashi et al. 1984; Kimoto and Takizawa 1993).

Equisetaceae: Equisetum arvense L., E. fluviatile L.

Full-depth or internal linear mine on leaf-like branch and the green stem (Fig. 6B–E). The larva is cylindrical (Fig. 6F) and sometimes exits its mine and moves to the other shoot. Frass is thread-like, deposited in the mine.

Figure 6. 

Habitus of adults, leaf mine, and larva of Hippuriphila babai on Equisetum arvense A habitus of adults B, C leaf-mine at Mt. Teine D–F larva at Yubari, Hokkaido.

• 2 adults, Mt. Teine, Sapporo, Hokkaidô, 9-X-1998 (as larva on Equisetum arvense), emerged on 25-VII–3-VIII-1998 (Fig. 6A–C); • 2 adults, Rebun Is., Hokkaidô, 9-VII-1995 (as larva on E. arvense), emerged on 25–27-VII-1995; • 5 adults, Bibi, Chitose, Hokkaidô, 30-VI-2023 (as larva on E. fluviatile), emerged on 17–20-VII-2023.

Brassicaceae: Brassica juncea (L.), Raphanus sativus L. var. raphanistroides (Makino), Rorippa palustris (L.), Eutrema japonicum (Miq.). Cardamine anemonoides, Brassica chinensis and B. napus are also recorded as host plants (Takizawa 2005).

Full-depth linear mine occurs on leaf blade, midrib, petiole, and shoot (Fig. 7C–E, H, I, L–P). The egg is laid on the leaf, and the hatched larva mines toward midrib, and reenters leaf blade. Larvae alternates mining the leaf and mining the midrib/petiole/shoot, and often relocates its mine. Frass is deposited compactly in a few parts of the mine, and sometimes discharged from the mine through the perforated holes. The fully grown larva exits the mined leaf, falls to the ground, pupates underground. Adult emerges ~ 1 month after pupation. On Eutrema japonicum with long petiole, linear mine of petiole and midrib of mature leaf (Fig. 7L–P). The mined petiole turns blackish. Frass is deposited compactly in a few parts of the mine, and sometimes discharged from the mine through perforated holes.

Figure 7. 

Habitus of adults and larvae, habitat, and leaf mines of three Psylliodes species A–J P. punctifrons K–P P. sasakii Q–T P. subrugosa. Host plants A–F Brassica juncea at Kizu-gawa, Kyôto G–I Raphanus sativus var. hortensis at Notojima Is., Ishikawa Pref. K–M Eutrema japonicum at Futamata, Oshamanbe, Hokkaido N–P Eutrema japonicum at Nyûkawa, Takayama, Gifu Pref. Q–T Arabis hirsuta at Shimo-suwa, Nagano Pref.

• 23 adults, Kusauchi, Kizu-gawa, Kyôtanabe, Kyoto Pref. 11-III-2020 (as larva on Brassica juncea), emerged on 1–15-V-2020 (Fig. 7A–E); • 1 adult, Kôda, Notojima Is., Nanao, Ishikawa Pref., 2-V-2023 (as larva on Raphanus sativus var. raphanistroides), emerged on 4-VI-2023 (Fig. 7G–J); • 2 adults, Watarase, Fujioka, Tochigi Pref., 25-V-2020 (as larva on Rorippa palustris), emerged on 8–14-VI-2020; • 19 adults, Futamata, Oshamanbe, Hokkaidô, 20-V-2023 (as larva on Eutrema japonicum), emerged on 14–25-VI-2023 (Fig. 7K–M); • 2 adults, Nyû-kawa, Takayama, Gifu Pref., 1-V-2023 (as larva on Eutrema japonicum), emerged on 29-V–5-VI-2023 (Fig. 7N–P).

Two female specimens were reared and tentatively identified as related to this species, but there is a possibility that this is an undescribed species.

Brassicaceae: Arabis hirsuta (L.). Crucifers are known as host plants (Kimoto and Takizawa 1993), while its biology has not been reported.

Linear mine in the midrib and the petiole of mature leaf (Fig. 7R–T). The egg is laid on the petiole, and the hatched larva mines the petiole, and sometimes enter the midrib. The mined petiole becomes blackish. Frass is deposited compactly in a few parts of the mine, and sometimes discharged from the mine through perforated holes. The fully grown larva (Fig. 7N) exits the mined leaf, falls to the ground, and pupates underground. The adult emerges ~ 1 month after pupation.

• 2 adults, Higashimata, Shimosuwa, Suwa-gun, Nagano Pref., 28-VI-2020 (as larva on Arabis hirsuta), emerged on 9-VIII-2020 (Fig. 7Q, S, T): several leaf mines on Arabis hirsuta, Sanjiro, Matsumoto, Nagano Pref., 28-VII-1995 (Fig. 7R).

Polypodiaceae: Lepisorus miyoshianus (Makino)., L. onoei (Franch. et Sav.), L. thunbergianus (Kaulf.), Pyrrosia linearifotia (Hook.) and Lemmaphyllum microphyllum C. Presl. are reported as adult’s host plants by Suzuki et al. (2008).

Upper-layer linear-blotch mine on mature leaf (Fig. 8B, F, G). The fully grown larva (Fig. 8H) exits the mined leaf, falls to the ground, and pupates underground.

Figure 8. 

Habitus of adults and leaf mines of four Halticorcus species A–H H. kasuga I–L H. sauteri M–T H. hiranoi U–X H. duodecimmaculata. Host plants A, B Lepisorus thunbergianus at Mt. Shizuhata, Shizuoka Pref. C Lepisorus onoei at Shôjiko Lake, Yamanashi Pref. D–H L. miyoshianus at Kiso-fukushima, Nagano Pref. I–J Leptochilus neopothifolius at Kanyû, Kakeroma Is., Kagoshima Pref. K–L Cyrtomium falcatum at Imasato, Amami-ôshima Is., Kagoshima Pref. M–O Phymatosorus scolopendria at Tonaki Is., Okinawa Pref. P, Q Loxogramme salicifolia at Inohae, Kitago, Miyazaki Pref. R–T Pyrrosia lingua at Mt. Yuwan, Amami-ôshima Is., Kagoshima Pref. U, V Phymatosorus scolopendria at Iriomote Is., Okinawa Pref. W, X Crypsinus yakushimensis at Iriomote Is., Okinawa Pref.

• 3 adults, Mt. Shizuhata, Aoi-ku, Shizuoka Pref., 27-VII-2017 (as larva on Lepisorus thunbergianus), emerged on 12–18-IX-2017 (Fig. 8A, B); • 1 adult on L. onoei, Shôji-ko Lake, Fuji-kawaguchi-ko, Yamanashi Pref., 8-IX-2019 (Fig. 7C); • 2 adults, Shimo-shimizu, Kiso-fukushima, Nagano Pref., 7-VIII-2011 (as larva on L. miyoshianus), emerged on 4-IX-2011 (Fig. 7D–H).

Dryopteridaceae: Cyrtomium falcatum (L. f.); Oleandraceae: Nephrolepis cordifolia (L.); Polypodiaceae: Colysis elliptica (Thunb.), Loxogramme salicifolia (Makino), Leptochilus neopothifolius Nakaike, Lemmaphyllum microphyllum Presl., Microsorum buergerianum (Miq.), Phymatosorus scolopendria (Burm. f.). Thelypteris acuminata (Houtt.) is also reported as adult’s host plants by Suzuki et al. (2008).

Upper-layer linear-blotch mine on mature leaf (Fig. 8J–L, N, O). Frass is thread-like, thin, and often intermittent, and deposited along the meandering larval trajectory. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

• 1 adult and several leaf mines, Imasato, Uken, Amami-ôshima Is. Kagoshima Pref., 4-IX-2011 on Cyrtomium falcatum (Fig. 8K, L); • 2 adults, Angyaba, Tatsugô, Amami-ôshima Is. Kagoshima Pref., 8-V-1997 (as larva on Cyrtomium falcatum), emerged on 22-VI-1997; • 22 adults, Kinsakubaru, Naze, Amami-ôshima Is. Kagoshima Pref., 3-VI-1996 (as larva on Nephrolepis cordifolia), emerged on 2-VII-1996; • 1 adult, Naon, Yamato, Amami-ôshima Is. Kagoshima Pref., 11-V-2002 (as larva on Colysis elliptica), emerged on 11-VI-2002; • 10 adults, Segiri, Nagata, Yaku Is., Kumage-gun, Kagoshima Pref., 27-V-1990 (as larva on Leptochilus neopothifolius), emerged on ?-VII-1990; • 3 adults, Higashinamaka, Amami-ôshima Is. Kagoshima Pref., 20-V-2015 (as larva on Leptochilus neopothifolius), emerged on 17–18-VI-2015 (Fig. 8I–J); • 2 adults, Mt. Yuwan, Uken, Amami-ôshima Is., Kagoshima Pref., 30-VI-1992 (as larva on Microsorum buergerianum), emerged on 25-VII-1992; • 1 adult and several leaf mines, Tonaki Is., Shimajiri-gun, Okinawa Pref., 18-III-2020 (as larva on on Phymatosorus scolopendria), emerged on 20-V-2020 (Fig. 8M–O).

Aspleniaceae: Asplenium antiquum Makino; Polypodiaceae: Pyrrosia lingua, Lemmaphyllum microphyllum, Lepisorus thunbergianus, Leptochilus neopothifolius, Loxogramme salicifolia; Vittariaceae: Vittaria flexuosa Fee.

Upper-layer linear-blotch mine on mature leaf (Fig. 8Q, S, T). Frass is thread-like, thin, and often intermittent, and deposited along meandering larval trajectory. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

• 1 adult and several leaf mines, Inohae, Nichinan, Miyazaki Pref., 20-V-2015 on Loxogramme salicifolia (Fig. 8P, Q); • 3 adults, Mt. Yuwan, Uken, Amami-ôshima Is., Kagoshima Pref., 8-III-2000 (as larva on Pyrrosia lingua), emerged on 14–18-V-2000 (Fig. 8R–T); • 1 adults, Mt. Yuwan, Uken, Amami-ôshima Is., Kagoshima Pref., 16-II-1999 (as larva on Lepisorus thunbergianus), emerged on 22-IV-1999; • 4 adult, Mt. Yuwan, Uken, Amami-ôshima Is., Kagoshima Pref., 7-III-2004 (as larva on Asplenium antiquum), emerged on 25-V–5-VI-2004; • 2 adults, Mt. Katsuu-dake, Nago, Okinawa Pref., 23-XII-1989 (as larva on Lemmaphyllum microphyllum), emerged on ?-III-1990; • 1 adult, Inokawa-dake, Tokunoshima Is., Kagoshima Pref., 12-III-2001 (as larva on Vittaria flexuosa), emerged on 25-VI-2001; 8 adults, Kuroshima Is., Mishima-mura, Kagoshima Pref., 11-III-1995 (as larva on Vittaria flexuosa), emerged on 21–25-V-1995.

Polypodiaceae: Phymatosorus scolopendria, Selliguea yakushimensis (Makino).

Full-depth linear-blotch mine on mature leaf (Fig. 8V–X). Frass is thread-like, thin, and often intermittent, and deposited along meandering larval trajectory. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

• 5 adults, Ômija, Iriomote Is., Yaeyama, Okinawa Pref., 27-III-2018 (as larva on Phymatosorus scolopendria), emerged on 25-V–7-VI-2018 (Fig. 8U, V); • 9 adults, Urauchi, Iriomote Is., Yaeyama, Okinawa Pref., 11-V-1999 (as larva on Selliguea yakushimensis collected by Shirô Kobayashi), emerged on ?-VI-1999 (Fig. 8W, X).

Oleaceae: Ligustrum micranthum Zucc., L. ovalifolium Hassk. Ligustrum japonicum Thunb., Osmanthus heterophyllus (G. Don), O. insularis Koidz., O. × fortunei Carr. are also recorded as adult hosts (Kimoto and Takizawa 1993). In Ogasawara Islands, this is the only leaf-mining chrysomelid species.

Full-depth linear mine on young leaf (Fig. 9B). The egg is laid on leaf margin, and the hatched larva mines adjoining along the leaf margin or its mine. Frass is thread-like, deposited along the middle line of the mine. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

Figure 9. 

Habitus of adults and leaf mines of four Argopistes species A, B A. coccinelliformis C–H A. biplagiata I–N A. tsekooni O–Q A. ryukyuensis. Host plants A, B Ligustrum ovalifolium at Shirahama, Wakayama Pref. C, D Osmanthus × fortunei at Mizorogaike Lake, Kyôto Pref. E–H Osmanthus heterophyllus at Iwakura, Kyôto Pref. I–K Ligustrum obtusifolium at Mt Nabejiri, Shiga Pref. L Fraxinus sieboldiana at Uri-tôge, Hamamatsu, Shizuoka Pref. M, N Syringa reticulata at Tôro Lake, Kushiro, Hokkaido O–Q Ligustrum japonicum at Abu, Okinawa Is., Okinawa Pref.

• 3 adults, Rinkai, Shirahama, Nishimuro-gun, Wakayama Pref., 13-V-1998 (as larva on Ligustrum ovalifolium), emerged on 2-VI-1998 (Fig. 9A, B); • 1 adult, Mukô-jima Is. Ogasawara, Tokyo Pref. on Ligustrum micranthum.

Oleaceae: Ligustrum japonicum, Osmanthus heterophyllus, O. insularis, O. × fortunei.

Full-depth linear mine on young leaf (Fig. 9D, G, H). The egg is laid on the leaf margin, and the hatched larva mines adjoining along the leaf margin or its mine. Frass is thread-like, deposited along the middle line of the mine in young instars, but in old instars along meandering larval trajectory, scattered throughout the mine. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

• 15 adults and many leaf mines, Mizorogaike Lake, Sakyo, Kyoto Pref., 13-V-1998 on Osmanthus × fortunei (Fig. 9C, D); • 7 adults, Iwakura, Sakyo, Kyoto Pref., 13-V-2016 (as larva on Osmanthus heterophyllus), emerged on 4–8-VI-2016 (Fig. 9E–H).

Oleaceae: Ligustrum obtusifolium Sieb. et Zucc., Fraxinus sieboldiana Blume, Syringa reticulata (Blume).

Full-depth wide linear mine on mature leaf in young instars and full-depth ophiogenous blotch mine in old instars (Fig. 9J–N). The mine is wider than the larval width, and frass is thick thread-like, deposited linearly along the middle line of the mine. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

• 3 adults, Mt. Nabejiri, Taga, Shiga Pref., 23-V-2015 (as larva on Ligustrum obtusifolium), emerged on ?-VII-2015 (Fig. 9I–K); • 7 adults, Iwakura, Sakyo, Kyoto Pref., 22-IV-2013 (as larva on Fraxinus sieboldiana), emerged on 3-VI-2013 (Fig. 9L); • 7 adults, Tôro Lake, Shibecha, Kawakami-gun, Kushiro, Hokkaido, 25-VI-2017 (as larva on Syringa reticulata), emerged on 8-VIII-2017 (Fig. 9M, N).

Oleaceae: Ligustrum japonicum.

Full-depth wide linear mine on mature leaf in young instars and full-depth ophiogenous blotch mine in old instars (Fig. 9O–Q). Frass is thread-like, deposited along the middle line of the mine in young instars, but in old instars along meandering larval trajectory, scattered throughout the mine. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

Several leaf mines, Abu, Nago, Okinawa Is., Okinawa, Pref., 18-V-2017 (Fig. 9O–Q).

Oleaceae: Osmanthus heterophyllus (Hayashi et al. 1984; Kimoto and Takizawa 1993).

Unknown.

Ranunculaceae: Clematis stans Sieb. et Zucc. C. apiifolia DC. and Clematis terniflora DC. are recorded as adult hosts (Hayashi et al. 1984; Kimoto and Takizawa 1993).

Upper-layer radiate mine along primary leaf vein in young instars (Fig. 10C) and full-depth blotch mine in old instars (Fig. 10D) occur on the mature leaves. Frass is thin threadlike, deposited along meandering larval trajectory in the mine. The fully grown larva exits the mine, falls to the ground, and pupates underground.

Figure 10. 

Habitus of adults and leaf mines of four Argopus species A–D A. balyi E–H A. clypeatus. Host plants A–D Clematis stans at Fukuji-onsen, Gifu Pref. E–H Clematis terniflora (E–G at Tsushima Nagasaki Pref., H at Uri-tôge, Shizuoka Pref.).

• 1 adult, Fukuji-onsen, Takayama, Gifu Pref., 3-X-2018 (as larva on Clematis stans), emerged on 9-V-2018 (Fig. 10A–D); • 1 adult, Iwakura, Sakyo, Kyoto Pref., 12-X-2009 (as larva on C. stans), emerged on 10-V-2009.

Ranunculaceae: Clematis terniflora is recorded as larval and adult hosts (Kimoto and Takizawa 1993).

Ranunculaceae: Clematis terniflora, C. apiifolia (Hayashi et al. 1984; Kimoto and Takizawa 1993).

Whitish full-depth blotch mine on mature leaf (Fig. 10G, H). Frass is granular, deposited along the larval trajectory in the mine. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

• 3 adults, Mine, Kami-tsusima Is., Nagasaki Pref., 19-VI-2016 (as larva on Clematis terniflora), emerged on 18–29-VIII-2016 (Fig. 10E–G); • 6 adults, Uri-tôge, Mikkabi, Hamamatsu, Shizuoka Pref., 22-IV-2013 (as larva on C. terniflora), emerged on 28-V–7-VI-2013 (Fig. 10H); • 5 adults, Nekata, Hamakita, Shizuoka Pref.

Leaf-mining larvae of this species have been found from 26 plant species belonging to three taxonomically isolated plant families: Aristolochiaceae (Fig. 11A–G), Ranunculaceae (Fig. 11H–M) and Asteraceae (Fig. 11N–T). The morphology of the adult beetle (Fig. 11A, B, G–H, N–P) and its male genitalia (Fig. 11C, D, J, K, P, Q) suggest that all these specimens belong to this species.

Figure 11. 

Adult morphology and leaf mines of Argopus punctipennis A–H on Asarum I–N on Aconitum O–S on Cirsium. Host plants A–F Asarum heterotropoides at Samani, Hidaka, Hokkaido G–H Asarum megacalyx at Mt. Haguro, Tsuruoka, Yamagata Pref. I–M Aconitum japonicum subsp. subcuneatum at Shiriya, Aomori Pref. N Aconitum sachalinense at Hinoemata, Fukushima Pref. O–R Cirsium kamtschaticum at Aibetsu, Kamikawa, Hokkaido, S Cirsium japonicum at Mt. Mikusa, Hyogo Pref. A, I, O Habitus B, J, P frontal head C, D, K, L, Q, R male genitalia in ventral and lateral views E–G, L, M, R, S leaf mines.

Aristolochiaceae: Asarum asperum F. Maek., A. blumei Duch., A. caulescens Maxim., A. curvistigma F. Maekawa, A. heterotropoides Miq., A. megacalyx (F. Maek.), A. nipponicum F. Maek., A. sieboldii Miq., A. tohokuense Yamaji et Ter. Nakam; Ranunculaceae: Aconitum gigas H. Lév. et Vaniot, A. japonicum Thunb., A. okuyamae Nakai, A. pterocaule Koidz., A. sachalinense F. Schmidt; Asteraceae: Cirsium austrohidakaense Kadota, C. iito-kojianum Kadota, C. japonicum Fisch. ex DC., C. kamtschaticum Ledeb, C. kiotoense (Kitam.), C. makinoi Kadota, C. microspicatum Nakai, C. oligophyllum (Franch. et Sav.), C. otayae Kitam., C. taishakuense Kadota, C. ugoense Nakai, C. yoshinoi Nakai.

Upper-layer linear-blotch mine in young instars, and full-depth blotch mine in old instars occur on mature leaves (Fig. 11E–G, L, M, R, S). Frass is thin thread-like, deposited along meandering larval trajectory in the mine. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground. The mining pattern is largely similar among leaves of the different plant species/families.

Aristolochiaceae: • 1 adult, Samani, Hidaka, Hokkaidô, 31-V-2020 on Asarum heterotropoides (Fig. 11A, B); • 4 adults, Samani, Hidaka, Hokkaidô, 31-V-2020 (as larva on Asarum heterotropoides), emerged on 13–14-VII-2020 (Fig. 11C–E); • 1 adult, Hachimori, Happô, Yamamoto-gun, Akita Pref., 15-VI-2015 (as larva on Asarum sieboldii), emerged on ?-VII-2015; • 4 adults, Mt. Haguro, Tsuruoka, Yamagata Pref., 9-VII-2018 (as larva on Asarum megacalyx), emerged on 19-VIII–4-IX-2018 (Fig. 11G, H); • 6 adults, Mt. Yakeishi, Ôshû, Iwate Pref., 15-VII-2019 (as larva on Asarum tohokuense), emerged on 30-VIII-2018; • 1 adult, Mt. Kiyosumi, Kamogawa, Chiba Pref., 14-V-2008 (as larva on Asarum nipponicum), emerged on 20-VII-2008; • 1 adult, Warabino, Aoi-ku, Shizuoka, Shizuoka Pref., 11-V-2004 (as larva on Asarum curvistigma), emerged on 21-VII-2004; • 1 adult, Mt. Gassan, Tsuruoka, Yamagata Pref., 12-VI-2019 (as larva on Asarum sieboldii), emerged on 16-V-2019; • 1 adult, Inogashira, Fujinomiya, Shizuoka Pref., 26-V-2002 (as larva on Asarum caulescens), emerged on 4-VII-2002; • 1 adult, Iwakura, Sakyo, Kyoto Pref., 24-V-1991 (as larva on Asarum asperum), emerged on ?-VII-1991.

Ranunculaceae: • 2 adults, Shiriya, Higashidôri, Shimokita, Aomori Pref., 16-VI-1995 (as larva on Aconitum japonicum), emerged on 13–14-VII-1995 (Fig. 11I–M); • 6 adults, Mt. Teine, Sapporo, Hokkaidô, 10-VII-1995 (as larva on Aconitum yezoense), emerged on 5-VIII-1995; • 4 adults, Monbetsu, Hidaka, Saru-gun, Hokkaidô, 5-VI-1993 on Aconitum yezoense; 3 adults, Mt. Obira, Shimamaki, Hokkaidô, 6-VII-2011 (as larva on Aconitum pterocaule, emerged on 28-VII-2011; • 5 adults, Hinoemata, Aizu-gun, Fukushima Pref., 16-VII-2023 (as larva on Aconitum japonicum, emerged on 17–21-VIII-2023; 5 adults, Mt. Hiuchi, Myôkô, Niigata Pref., 16-VII-2023 (as larva on Aconitum japonicum), emerged on 17–21-VIII-2023 (Fig. 11N); 12 adults, Funakawa, Oga, Akita Pref., 2-VI-2017 (as larva on Aconitum japonicum), emerged on 24–28-VII-2017; • 3 adults, Atsumi, Tsuruoka, Yamagata Pref., 2-VI-2017 (as larva on Aconitum okuyamae), emerged on 20-VII-2017; • 4 adults, Tokoro, Abashiri, Hokkaidô, 24-VII-2017 (as larva on Aconitum gigas), emerged on 3–15-IX-2017; • 1 adult, Mt. Yudono, Tsuruoka, Yamagata Pref., 28-VIII-2017 (as larva on Aconitum pterocaule), emerged on 15-X-2017; • 1 adult, Donden, Sado Is., Niigata Pref., 13-VII-2019 (as larva on Aconitum japonicum), emerged on ?-XI-2019; • 2 adults, Mt. Fuji, Fujinomiya, Shizuoka Pref., 27-VI-2001 (as larva on Aconitum japonicum), emerged on 9–12-VIII-2001.

Asteraceae: 14 adults, Aibetsu, Kamikawa, Hokkaidô, 26-VI-2016 (as larva on Cirsium kamtschaticum), emerged on 5–21-VIII-1995 (Fig. 11O–S); 4 adults, Kashiwadai, Chitose, Hokkaidô, 26-VI-2017 (as larva on C. kamtschaticum), emerged on 8–15-VIII-2017; 3 adults, Rebun Is., Hokkaidô, 9-VII-1995 (as larva on C. kamtschaticum), emerged on 10-VIII-1995; 1 adult, Tôbai, Nemuro, Hokkaidô, 23-VII-2018 (as larva on C. iito-kojianum), emerged on 28-VIII-2018; 2 adults, Mt. Yûbari, Yubari, Hokkaidô, 20-VII-2020 (as larva on C. austrohidakaense), emerged on 5–9-IX-2020; 1 adult, Shiriya, Higashidôri, Shimokita, Aomori Pref., 16-VI-2017 (as larva on C. aomorense), emerged on 13-VII-2017; 1 adult, Funakawa, Oga, Akita Pref., 2-VI-2017 (as larva on C. makinoi), emerged on ?-VII-2017; 4 adults, Mt. Haguro, Tsuruoka, Yamagata Pref., 9-VII-2018 (as larva on C. ugoense), emerged on 10-VIII--2018; 1 adult, Mt. Hiuchi, Myôkô, Niigata Pref., 14-VIII-2023 (as larva on C. otayae), emerged on 10–11-IX-2023; 1 adult, Mt. Hiuchi, Myoko, Niigata Pref., 4-V-2002 (as larva on C. oligophyllum), emerged on 10-VI-2002; 2 adults, Iwakura, Sakyo, Kyoto Pref., 1-V-1999 (as larva on C. kiotoense), emerged on 4-VI-1999; 1 adult, Mt. Kiyosumi, Chiba Pref., 14-V-2008 (as larva on C. japonicum), emerged on 20-VII-2008; 2 adults, Tochiu, Takashima, Shiga Pref., 6-V-1998 (as larva on C. yoshinoi), emerged on 7-VI-2017; 1 adult, Ippekiko Lake, Ito, Shizuoka Pref., 15-V-1999 (as larva on C. microspicatum), emerged on 14-VI-1999; 2 adults, Mt. Mikusa, Katô, Hyôgo Pref., 20-V-2018 (as larva on C. japonicum), emerged on 27–28-VI-2018; 1 adult, Yasukawa-keikoku, Tanabe, Wakayama Pref., 15-IV-2007 (as larva on C. yoshinoi), emerged on 16-V-2007; 8 adults, Hashirajima Is. Iwakuni, Yamaguchi Pref., 8-V-1993 (as larva on C. japonicum), emerged on 14–17-VI-1993.

A host plant has not been reported (Hayashi et al. 1984; Kimoto and Takizawa 1993).

A host plant has not been reported (Hayashi et al. 1984; Kimoto and Takizawa 1993).

Liliaceae: Cardiocrinum cordatum (Thunb.), Lilium auratum Lindley, Tricyrtis flava Maxim, T. macropoda Miq.; Smilacaceae: Smilax china L., S. nipponica Miq., S. riparia A. DC., S. stans Maxim., Heterosmilax japonica Kunth; Stemonaceae: Croomia heterosepala (Baker), C. japonica Miq. Note: this species was collected on 11 host plants (six genera and three families of monocots, and the emerged adults varied in color and size among their host plants).

Full-depth long linear mine on mature leaf, often adjoining its own previous trajectory (Fig. 12B, D, G, H, J, N. O). Frass is thread-like, deposited along the middle line of the mine. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground. The mine on Tricyrtis is unique in that the mine in young instars is not linear but radiate (Fig. 12D).

Figure 12. 

Habitus of adults and leaf mines of host races of Sphaeroderma nigricolle. Host plants A, B Lilium auratum at Mt. Haguro-san, Yamagata Pref. C, D Tricyrtis macropoda at Tengu-kogen, Kôchi Pref. E–G Smilax stans at Mt. Torigata, Kôchi Pref. H Smilax stans at Mt. Ôtaki, Kagawa Pref. I–K Croomia heterosepala at Kikuchi-keikoku, Kumamoto Pref. L–O Croomia japonica at Naon, Amami-ôshima Is., Kagoshima Pref.

• 5 adults, Mt. Haguro, Tsuruoka, Yamagata Pref., 8-VII-2018 (as larva on Lilium auratum), emerged on 31-VIII–7-IX-2018 (Fig. 12A, B); • 1 adult, Iwakura, Sakyo, Kyoto Pref., 8-IX-2008 (as larva on Tricyrtis macropoda), emerged on 16-XI-2008 (Fig. 12C, D); • 1 adult, Mt. Torigata, Niyodogawa, Agawa-gun, Kôchi Pref., 6-X-2020 (as larva on Smilax stans), emerged on 4-III-2021 (Fig. 9E–G); • 2 adults, Mt. Haguro, Tsuruoka, Yamagata Pref., 12-VI-2019 (as larva on Smilax nipponica), emerged on 16-VIII-2019; several leaf mines, Mt. Ôtaki, Kagawa, Pref., 7-IX-1998 on Smilax stans (Fig. 12H); • 1 adult, Kikuchi-keikoku, Kumamoto Pref., 2-V-2018 on Croomia japonica (Fig. 12I–K); • 1 adult, Naon, Amami-ôshima Is., Kagoshima Pref., 12-XII-2014 (as larva on Croomia heterosepala), emerged on 29-VI-2015 (Fig. 12L–O).

Commelinaceae: Commelina communis L.

Full-depth long linear mine on mature leaf (Fig. 13B, C). Frass is thread-like, deposited along the middle line of the mine. The fully grown larva exits the mined leaf, falls to the ground, and pupates underground.

Figure 13. 

Habitus of adults and leaf mines of monocot-associated Sphaeroderma species A–C S. japanum D–L S. tarsatum M–P S. seriatum. Host plants A–C Commelina communis at Makidô, Niimi, Okayama Pref. E–G Sasa nipponica at Akiu, Sendai, Miyagi Pref. H, I Sasa senanensis at Kamikôchi, Nagano Pref. J–L Stipa coreana var. japonica at Mt. Fuji, Shizuoka M–P Panicum bisulcatum at Yogo Lake, Shiga Pref. (M–O) and at Seryo, Kyôto Pref. (P).

• 5 adults, Maki-dô, Niimi, Okayama Pref., 22-VI-2020 (as larva on Commelina communis), emerged on 16-III-2021 (Fig. 13A–C).

Poaceae: Phyllostachys bambusoides Sieb. et Zucc., Pleioblastus chino (Franch. et Savat.) var. viridis (Makino), Sasa kurilensis (Ruprecht), S. nipponica M. et S., S. senanensis (Franch et Savat.), Sasamorpha borealis (Hack.), Shibataea kumasaca Makino, Stipa coreana Honda var. kengii Ohwi).

Upper-layer long linear mine on mature leaf, often adjoining its own trajectory (Fig. 13E–G, I, L). The egg is laid in leaf blade, hatched larva mines usually toward leaf apex along leaf vein, and sometimes turns adjoining its past mine. Frass is granular, deposited linearly along the middle line of the mine. The mining larva is found in autumn from October to December. The fully grown larva exits the mined leaf from late autumn to early winter, falls to the ground, and pupates underground. Pupa hibernates under the ground, and the adult emerges the next spring.

• 2 adults, Akiu, Taihaku-ku, Sendai, Miyagi Pref., 14-XI-2014 (as larva on Sasa nipponica), emerged on 7-V-2015 (Fig. 13D–G); • 1 adult, Azusagawa, Matsumoto, Nagano Pref., 24-X-2020 (as larva on Sasa senanensis), emerged on 2-III-2021 (Fig. 13H, I); • 1 adult, Ashiu, Nantan, Kyoto Pref., 9-XI-1999 (as larva on Sasa kurilensis), emerged on 28-III-2000; 1 adult, Kuchisakamoto, Aoi-ku, Shizuoka, Shizuoka Pref., 27-XII-2005 (as larva on Sasamorpha borealis), emerged on 1-III-2006; • 1 adult, Inogashira, Fujinomiya, Shizuoka Pref., 11-XI-2001 (as larva on Phyllostachys bambusoides), emerged on 14-V-2002; • 3 adult, Usuzuka, Mt. Fuji, Fujinomiya, Shizuoka Pref., 24-X-2019 (as larva on Stipa coreana var. kengii), emerged on 7–10-IV-2020 (Fig. 13J–L).

Poaceae: Panicum bisulcatum Thunb.

Full-depth linear mine on mature leaf (Fig. 13N–P). The egg is laid along the leaf margin, and the hatched larva mines usually toward leaf apex along leaf margin, and sometimes turns adjoining its mine. Frass is intermittent thread-like, deposited linearly in two rows along both sides of the mine. Mining larva is found in October. The fully grown larva exits the mined leaf from late autumn to early winter, falls to the ground, pupates underground, and the adult emerges in late autumn.

• 3 adults, Yogo Lake, Nagahama, Shiga Pref., 12-IX-2015 (as larva on Panicum bisulcatum), emerged on 7-X-2015 (Fig. 13M–O); • 4 adults, Seryô, Sakyô, Kyoto, Kyoto Pref., 11-IX-1998 (as larva on Panicum bisulcatum), emerged on 28-IX–1-X-1998 (Fig. 13P).

Poaceae: Miscanthus sinensis (Hayashi et al. 1984; Kimoto and Takizawa 1993).

Unknown.

Lardizabalaceae: Akebia trifoliata (Thunb.), A. quinata Decne.

Upper-layer linear mine on mature leaf (Fig. 14B, C). The mine is wider than the larval width, having dead ends and branches. Frass is granular and minute, deposited linearly in two rows along both sides of the mine, and inner space between the frass lines is colored darker against the outer space. The mining larva is found from late autumn to early winter. The fully grown larva exits the mined leaf in early winter, falls to the ground, and pupates underground. The pupa hibernates underground, and the adult emerges the next spring.

Figure 14. 

Habitus of adults, habitats, and leaf mines of Akebia-associated and Clematis-associated Sphaeroderma species A–C S. akebia D S. inaizumii E–L S. quadrimaculatum M–Q S. flavonotatum R–V S. separatum. Host plants A–C Akebia trifoliata at Tsuruoka, Yamagata Pref. D Akebia trifoliata at Shôji Lake, Yamanashi Pref. E–G Clematis taiwaniana var. ryukiuensis at Hedo, Okinawa Is., Okinawa Pref. J–L Clematis taiwaniana var. ryukiuensis at Tonaki Is., Okinawa Pref. M–P Clematis tashiroi at Iriomote Is. Okinawa Pref. S–V Clematis japonica at Shirakawa-gô, Gifu Pref.

• 3 adults, Shimoike Lake, Tsuruoka, Yamagata Pref., 15-XI-2014 (as larva on Akebia trifoliata), emerged on 7-V-2015 (Fig. 14A–C); • 2 adults, Aibano, Takashima, Shiga Pref., 13-XII-1998 (as larva on Akebia trifoliata), emerged on 8-IV-1999.

Lardizabalaceae: Akebia trifoliata, A. quinata.

Upper-layer blotch mine on mature leaf (Fig. 14D). The mine is orbicular, and the larva often relocates its mine. Frass is minute and often liquid, deposited as dark band in the mine. The mining larva is found from late autumn to early winter. The fully grown larva exits the mined leaf in early winter, falls to the ground, and pupates underground.

Many leaf mines, Shoji Lake, Kawaguchiko, Yamanashi Pref., 24-XI-2018 on Akebia trifoliata (Fig. 14D).

Ranunculaceae: Clematis taiwaniana Hayata var. ryukiuensis Tamura.

Full-depth linear mine on mature leaf (Fig. 14F–H, L). Frass is granular and minute, deposited linearly in two rows along both sides of the mine. The fully grown larva exits the mined leaf in early winter, falls to the ground, and pupates underground.

• 2 adults, Hedo, Kunigami, Okinawa Pref., 10-XI-2021 on Clematis taiwaniana var. ryukiuensis (Fig. 14E); • 3 adults, Hedo, Kunigami, Okinawa Pref., 14-X-2000 (as larva on C. taiwaniana var. ryukiuensis), emerged on 21-XI–10-XII-2000 (Fig. 14F–H); • 2 adults, Tonaki Is., Shimajiri-gun, Okinawa Pref., 18-III-2020 (as larva on C. taiwaniana var. ryukiuensis), emerged on 10-V-2020 (Fig. 14I–L).

Ranunculaceae: Clematis tashiroi Maxim. The host plant record of Smilax spp. by Takizawa (2021) is uncertain.

Full-depth linear mine on mature leaf (Fig. 14N–P). Frass is granular, deposited as a band along middle line of the mine. The fully grown larva (Fig. 14Q) exits the mined leaf in early winter, falls to the ground, and pupates underground.

• 6 adults, Funaura, Iriomote Is., Yaeyama, Okinawa Pref., 6-III-2019 (as larva on Clematis tashiroi), emerged on 20-IV-2019 (Fig. 14M–Q).

Ranunculaceae: Clematis japonica. Takizawa (2021) lists up three host plants: C. apiifolia DC., C. pierotti Miq. and Chelidonium majus L. (Papaveraceae).

Full-depth radiate mine along leaf vein on mature leaf (Fig. 14T–V). Frass is liquefied, deposited near the center of the mine.

• 2 adults copulating on a leaf of Clematis japonica at Shirakawa-gô, Shirakawa, Ôno-gun, Gifu Pref., 12-VIII-2024 (Fig. 14R, S), and many leaf mines on the plant species at the same locality 29-IX-2023 (Fig. 14T–V).

Ranunculaceae: Clematis apiifolia.

Black upper-layer blotch mine on mature leaf. Frass is granular scattered along larval trajectory at young instars, and accumulated in a discoid area in a center of the mine in old instars. The mining larva is found from late autumn to early winter. The fully grown larva exits the mined leaf in early winter, falls to the ground, and pupates underground. The pupa hibernates under the ground, and the adult emerges the next spring.

• 3 adults, Shoji Lake, Kawaguchiko, Yamanashi Pref., 24-XI-2018 (as larva on Clematis apiifolia), emerged on 10-V-2019 (Fig. 15A–D); • 2 adults, Kisofukushima, Kiso, Nagano Pref., 24-X-1999 (as larva on Clematis apiifolia), emerged on 22-V-2000; 11 adults and several leaf mines, Fukasawa, Gotenba, Shizuoka Pref., 29-VI-2014 on Clematis apiifolia; • 1 adult and several leaf mines on Clematis apiifolia, Ashiu, Nantan, Kyôto Pref., 13-IX-1993 (Fig. 15E–G).

Figure 15. 

Adult morphology, behavior, and leaf mines of Clematis-associated and Cimicifuga-associated Sphaeroderma species A–G S. placidum H–L S. unicolor M–R S. uenoi S–Z S. ohkuboi A, E, H, J, M, S, Y adult habitus in dorsal view F, N ventral view B, F, I, O Frontal view of head K, P, T adult feeding behavior C, D, G, L, Q, R, U–X, Z leaf mines. Host plants A–D Clematis apiifolia at Shyôji Lake, Yamanashi Pref. E–G Clematis apiifolia at Ashiu, Kyôto Pref. H, I Clematis terniflora at Obama, Fukui Prefecture J–L Clematis terniflora at Iya, Tokushima Prefecture M–Q Clematis terniflora at Kizu-gawa, Kyôto Pref. R Clematis terniflora at Iwakura, Kyôto Pref. S–X Cimicifuga japonica at Mt. Nakatsu-myôjin, Kôchi Pref. Y, Z Cimicifuga simplex at Shirahone-onsen, Nagano Pref.