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. 2018 Feb 8:(736):79-118.
doi: 10.3897/zookeys.736.20739. eCollection 2018.

Systematics of Phyllocnistis leaf-mining moths (Lepidoptera, Gracillariidae) feeding on dogwood (Cornus spp.) in Northeast Asia, with the description of three new species

Natalia Kirichenko  1   2   3 , Paolo Triberti  4 , Shigeki Kobayashi  5 , Toshiya Hirowatari  6 , Camiel Doorenweerd  7   8 , Issei Ohshima  9 , Guo-Hua Huang  10 , Min Wang  11 , Emmanuelle Magnoux  3 , Carlos Lopez-Vaamonde  3   12
Affiliations

Systematics of Phyllocnistis leaf-mining moths (Lepidoptera, Gracillariidae) feeding on dogwood (Cornus spp.) in Northeast Asia, with the description of three new species

Natalia Kirichenko et al. Zookeys. .

Abstract

During an ongoing DNA-barcoding campaign of the leaf-mining moths that feed on woody plants in Northeast Asia, four lineages of the genus Phyllocnistis (Gracillariidae, Phyllocnistinae) were discovered on dogwood (Cornus spp): P. cornella Ermolaev, 1987 on C. controversa Hemsl. (Japan: Hokkaido) and three new species - one feeding on C. controversa, C. florida L. and C. macrophylla Wall. in Japan (Honshu, Shikoku, Kyushu), a second species on C. macrophylla in China (Yunnan) and a third on Siberian dogwood Cornus alba L. in Russia (Siberia). All these species showed differences in morphology, in the barcode region of the cytochrome c oxidase I gene and in two nuclear genes (histone H3 and 28S ribosomal RNA). No correlation was found between the deep mitochondrial splits observed and the Wolbachia infection pattern. Based on both morphological and molecular evidence, the three recently discovered lineages are described here as new species: P. indistincta Kobayashi & Triberti, sp. n. (Japan), P. saepta Kirichenko, Ohshima & Huang, sp. n. (China) and P. verae Kirichenko, Triberti & Lopez-Vaamonde, sp. n. (Russia). In addition, the authors re-describe the adult morphology of P. cornella, provide the first record of this species from Japan and highlight the diagnostic characters that allow these Cornus-feeding Phyllocnistis species to be distinguished.

Keywords: China; Cornus; DNA barcoding; Japan; Phyllocnistis; Russia; Wolbachia; new species; nuclear genes.

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Figures

Figure 1.
Figure 1.
A neighbour-joining tree based on the COI barcode fragment of the 47 Phyllocnistis spp. specimens collected in Eurasia and beyond on woody plants from six families (indicated in the figure). Host plant genus / species is given in the figure for those leaf miners which were obtained directly from their mines; for specimens collected in other ways the host plant species remained unknown.
Figure 2.
Figure 2.
A neighbour-joining tree based on the H3 and 28S sequences in Cornus-feeding Phyllocnistis spp. from Northeast Asia.
Figure 3.
Figure 3.
Phyllocnistis indistincta, mines on leaves of Cornus spp. in Japan (Honshu, Nara Prefecture). A–D Cornus controversa E–I C. florida J, K C. macrophylla A, B Konagao, Kumatawa, 670 m C, D, H–J type locality: Soni, Imai, Menashi, 630 m E, F Konagao, Tawamine, 685 m G, K Imai, Oku-Kochi Sanso, 455 m I mine on bract. Arrows show mines (A, C, G, J) and pupation site (B, D, E, F, H, I, K). Scale bar: 50 mm.
Figure 4.
Figure 4.
Biology of Phyllocnistis indistincta in Japan (Honshu) and its hostplant, Cornus controversa. A, B, D, K, I Nara Prefecture, Soni, 400–630 m C, E Yamanashi Prefecture, Kawaguchi-ko, 880–950 m F–J, M type locality: Nara Prefecture, Soni, Imai, Menashi, 630 m A habitat B, C serpentine mines, lower side of leaves D old mine, same side E young mines, upper side of leaves F old mines, same side G, H later instar larva I cocoon fold J pupa, dorsal view K same, ventral view L same, lateral sview M resting posture of the adult, dorsal-lateral view. Arrows show mines. Scale bars: 2 mm (G, H), 1 mm (I–M).
Figure 5.
Figure 5.
Biology of Phyllocnistis indistincta in Japan (Honshu) and its host plants. A–H type locality: Nara Prefecture, Soni, Imai, Menashi, 630 m, Cornus florida I, J Nara Prefecture,Obako-dake, 1000 m, C. kousa K Yamanashi Prefecture, Yamanaka-ko, C. kousa, 1000 m L Nara Prefecture, Soni, Konagao, 685 m, C. macrophylla A habitat B host plant C flowers D–F, G later instar larva H cocoon fold I host plant J mines and branches of host plant K, L serpentine mines on upper side of leaves. Arrows show mines. Scale bars: 2 mm (G, H).
Figure 6.
Figure 6.
Biology of Phyllocnistis indistincta on Cornus florida in Japan (Honshu, Nara Prefecture, Soni, Imai, Oku-Kochi Sanso, 455 m). A habitat B serpentine mines on upper side of leaves C flowers and bracts, an arrow shows mine D serpentine mines on upper side of bracts E same, mines and cocoon folds F cocoon fold. Arrows show mines (C, D) and pupation site (D, E). Scale bar: 2 mm (F).
Figure 7.
Figure 7.
Biology of Phyllocnistis verae on Cornus alba in Russia (type locality: Krasnoyarsk, village Borovoe, left bank of Yenisei River, 144 m). A habitat B branch with mined leaves on the lower side C mine with feeding larva D fragment of mine with young larva (transmitted light) E same, incident light F line of frass and feeding larva (transmitted light) G opened mine H sap-feeding larvae, dorsal view I, J pupation near leaf margin K, L pupa. Arrows show mines (B, C), larva (D, E, F, G), frass (G, F), pupation site (I–K). Scale bars: 3 mm (D, E), 5 mm (F, G), 1 mm (H, L), 2 mm (J, K).
Figure 8.
Figure 8.
Biology of Phyllocnistis saepta on Cornus macrophylla in China (type locality: Yunnan Province, Weixi, 2800 m). A host plant B, C serpentine mine on the upper side of the leaf D pupation site under leaf fold on the upper side of the leaf E serpentine mine on the low side of the leaf F pupation site between two veins at a leaf margin. Arrows show mine (B, C, E), pupation site (D, F). Scale bars: 18 mm (C), 4 mm (D), 8 mm (F).
Figure 9.
Figure 9.
Adults of Phyllocnistis indistincta (Japan: Honshu, Kyushu) and P. saepta (China). A–E P. indistincta A holotype male, ex Cornus controversa, Honshu, Nara Prefecture [Suppl. material 1, No. 3] B paratype female, ex C. florida, Nara Prefecture [Suppl. material 1, No. 92] C paratype male, ex C. controversa, Nagano Prefecture [Suppl. material 1, No. 38] D ex C. kousa, Kyushu, Fukuoka Prefecture [Suppl. material 1, No. 133] E paratype female, Nara Prefecture [Suppl. material 1, No. 127] F P. saepta, holotype male, ex C. macrophylla, Yunnan Province [Suppl. material 1, No. 160].
Figure 10.
Figure 10.
Adults of Phyllocnistis verae (Russia) and P. cornella (Japan: Hokkaido). A–B P. verae, ex Cornus alba, Krasnoyarsk A holotype male [Suppl. material 1, No. 163] B paratype female [Suppl. material 1, No. 164] C–D P. cornella, ex C. controversa, Sorachi, Kamifurano C female [Suppl. material 1, No. 1] D female [Suppl. material 1, No. 2]. Scale bar: 1 mm.
Figure 11.
Figure 11.
Forewing colouration and patterns of Phyllocnistis indistincta from Cornus florida, C. controversa and C. kousa (Japan: Honshu). A, D, E ex Cornus florida B, C, F, G ex C. controversa H ex C. kousa I adult emerged from mined bract of C. florida A, D, E type locality: Nara Prefecture, Soni [Suppl. material 1, Nos 93 (A, D) 111 (E)] B, G Osaka Prefecture, Mt. Mikusa [Suppl. material 1, No. 46] C, F Nagano Prefecture, Ohshirakawa [Suppl. material 1, No. 38] H Nara Prefecture, Kawakami [Suppl. material 1, No. 126] I Tottori Prefecture, Mt. Daisen [Suppl. material 1, No. 129]. Arrows show variable characters. Scale bars: 1 mm.
Figure 12.
Figure 12.
Forewing colouration and patterns of Phyllocnistis indistincta from Cornus controversa and C. macrophylla (Japan: Honshu, Shikoku, Kyushu). A–I ex C. controversa J–L ex C. macrophylla A–C type locality, Honshu, Nara Prefecture, Soni, Imai, Menashi [Suppl. material 1, Nos 67 (A), 69 (B), 71 (C)] D Yamanashi Prefecture, Fuji-Kawaguchiko [Suppl. material 1, No. 44] E Nagano Prefecture, Ohshirakawa [Suppl. material 1, No. 37] F Tochigi Prefecture, Nasu Imperial Villa [Suppl. material 1, No. 14] G Aichi Prefecture, Kirara [Suppl. material 1, No. 45] H Shikoku, Tokushima Prefecture, Naka [Suppl. material 1, No. 141] I Kyushu, Fukuoka Prefecture, Hikosan [Suppl. material 1, No. 142] J–L Honshu, Nagano Prefecture, Okada [Suppl. material 1, Nos 135 (J), 133 (K), 134 (L)]; Arrows show variable characters. Scale bars: 1 mm.
Figure 13.
Figure 13.
Forewing venation of Phyllocnistis indistincta, ex Cornus florida, Japan (type locality: Honshu, Nara Prefecture, Soni, Imai, Menashi), wing vein slide SK580 [Suppl. material 1, No. 108].
Figure 14.
Figure 14.
Male genitalia of Phyllocnistis indistincta (Japan: Honshu). A–C paratype, ex Cornus controversa, Nara Prefecture, gen. slide SK163 [Suppl. material 1, No. 52] D–F paratype, ex C. controversa, Nagano Prefecture, gen. slide TRB4220 [Suppl. material 1, No. 33] A, E ventral view B, F lateral view C, D phallus B, F left valva. Scale bars: 0.2 mm (under B also refers to A, C, under D also refers to E, F).
Figure 15.
Figure 15.
Male genitalia of Phyllocnistis verae (Russia), P. septae (China), and P. cornella (Russia). A–C P. verae, holotype, ex Cornus alba, Krasnoyarsk, gen. slide TRB4200 [Suppl. material 1, No. 163] D, E P. saepta, holotype, ex C. macrophylla, Yunnan, gen. slide IsO-793 [Suppl. material 1, No. 160] F P. cornella, holotype, ex. C. controvera, Kunashir, drawing from Ermolaev (1987) B, E, F ventral view C lateral view A, D phallus C left valva. Scale bars: 0.2 mm (under A also refers to B, C under D also refers to E, F).
Figure 16.
Figure 16.
Female genitalia of Phyllocnistis indistincta (Japan: Honshu, Nara Prefecture). A, C paratype, ex Cornus controversa, gen. slide SK146 [Suppl. material 1, No. 51] B C. macrophylla, gen. slide TRB4148 [Suppl. material 1, No. 137] D ex C. kousa, gen. slide TRB4151 [Suppl. material 1, No. 128] E ex C. kousa, gen. slide TRB4153 [Suppl. material 1, No. 127]. Abbreviations: ap apophysis posterioris aa apophysis anterioris db ductus bursae ds ductus seminalis ob ostium bursae si signum. Scale bars: 0.2 mm.
Figure 17.
Figure 17.
Female genitalia of Phyllocnistis verae (Russia) and P. cornella (Japan: Hokkaido). A–C P. verae A paratype, ex Cornus alba, Krasnoyarsk, gen. slide TRB4116 [Suppl. material 1, No. 164] B paratype, ex C. alba, Krasnoyarsk, gen. slide TRB4152 [Suppl. material 1, No. 166] C paratype, ex C. alba, Krasnoyarsk, gen. slide NK-39-15-4 [Suppl. material 1, No. 168] D, E P. cornella, ex C. controversa, Sorachi, Kamifurano, gen. slides CDGP0275 and CDGP0136 [Suppl. material 1, Nos 1–2]. Scale bars: 0.2 mm.
Figure 18.
Figure 18.
Pupa of Phyllocnistis indistincta from Cornus controvera, Japan (Honshu, Nara Prefecture) [Suppl. material 1, No. 78]. A head, ventral view B cocoon cutter, ventral view C head, lateral view D cocoon cutter, lateral view E head, dorsal view F cocoon cutter, dorsal view G spines and a pair of setae of abdominal tergum H same, lateral view I A7–A10, lateral view J A8–A10, ventral view K same, lateral view L same, dorsal view. Scale bars: 400 µm (A, E), 300 µm (I), 200 µm (C, G, J, L), 100 µm (B, D, F, H, K).
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