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Philodoria lama Kobayashi, Johns & Kawahara, sp. n. Figs. 9C, 41L, 42K, 43K, 56L, 73E, F, 83. Philodoria sp. 9; Johns et al. 2018: fig. 2. Type locality. Kanepuu (Lanai). Etymology. The specific epithet, lama (pronounced 'La-mah') is a noun in apposition taken from the Hawaiian name of the host plant, Diospyros. Type material. Holotype ♂, Kanepuu, Lanai, 24.viii.2014 (stored), K. Bustamente leg., host: Diospyros sandwicensis, 5.viii. 2014, CJ382, SK744♂ in BPBM. Paratype 4 (3♂, 1♀), All types were preserved in BPBM. 3♂, 1♀, Lualailua, Maui, 24.v.2016 (stored), K. Bustamente leg., host: D. sandwicensis, Spring. 2016, CJ540, SK 813♀, SK814 ♂. The holotype was mounted by placing four wings without mountant under a coverslip (2/3 of two forewings and apical half of two hindwings); four paratypes: a female specimen mounted as a dry pinned specimen with the nearly entire right forewing without mountant under a coverslip; CJ540 / SK813); two male specimens mounted as dry pinned specimens with its near complete forewing without mountant under a coverslip; a male specimen mounted by placing two forewings without mountant under a coverslip (two near complete forewings slightly damaged at base; CJ540 / SK814). The head, antenna, thorax, and legs for all specimens were sacrificed for molecular analysis. Additional material. 1♂, Lualailua, Maui, 30.xii.2013 (stored), C.A. Johns leg., host: D. sandwicensis, 24.xii.2013, CJ161, PHIL0037♂ (abdomen only) in BPBM. Diagnosis. The forewing pattern, genital structure and larval habit are very similar to those of P. splendida Walsingham and P. basalis Walsingham, but are distinguished from them because of the basal oblique fascia of the forewing, slender saccus and longer coecum in male genitalia, and coecum which is about 1/4 length of phallus in P. lama, but about 1/6~1/5 length in P. splendida (Figs. 9C, 42I, 43I, 73E, F). Description: Adult (Figs. 9C, 73E, F). Forewing length 3.4, 3.5 mm in two paratypes. Head and frons brown; maxillary palpus reduced; labial palpus ocherous to brown with dark brown scales at apex. Antenna grayish fuscous, about 4.2 mm long in paratype. Thorax gray. Forewing dark shiny, pale leaden gray with brownish ocherous patches enclosing gray fascia: bp sometimes absent in Lanai specimens, from base along to dorsum 1/4, sometimes from near base of costal fold to near dorsum 1/ 4 in Maui specimens; tf from costal 1/3 to dorsal 1/2, tp after the middle to apical portion, distinctly narrowing in the dorsum, extending to dorsal 2/3, containing two short, shiny, steel gray costal spots; apical portion with black as and a large jet-black terminal patch from tornus to apex, containing two upright steel-gray spots near its opposite extremities; a single bright white spot in its middle; a very narrow ocherous line along the base of tornus; cilia shining, leaden gray with a blackish fringe line; two white costal streaks, long one extend apex; terminal cilia often blackish. Hindwing coppery brown; cilia tawny. Abdomen tawny brown, white below. Legs grayish fuscous, with white spots on the tarsal joints; spurs white with fuscous scales. Male genitalia (Figs. 41L, 42K, 43K) (n=3). Capsule 810 µm. Tegumen slightly shorter than length of valva; valva 460 µm long, similar to P. splendida except minute spines on dorsal process covered from apically to basally and on inner margin (Fig. 41L). Phallus 560 µm long (Fig. 43K). Female genitalia (Fig. 56L) (n=1). 1240 µm long. Lamella antevaginalis 160 µm in length. Corpus bursae 770 µm long. Similar to P. basalis and P. splendida. Distribution. Oahu, Lanai, and Maui. Host plants. Ebenaceae: Diospyros sandwicensis (A.DC.) Fosberg and/or D. hillebrandii (A.DC.) Fosberg. We could not separate the two Diospyros species, D. sandwicensis and the closely related D. hillebrandii. Biology. (Fig. 83). Biology unknown, but it is presumably similar to that of P. basalis and P. splendida. Larvae form an elongate blotch mine on the adaxial leaf surface (Fig. 83A–C). One mine per leaf (Fig. 83A). Most of the “Taco” cocoons were still attached to the leaf and leaves with cocoons were both on ground and on tree, based on observations at Lualailua, Maui. Remarks. We have observed Philodoria leaf mines on Diospyros on Oahu, but we were unable to find any mines with larvae in them (Fig. 83D). Swezey (1910a) described the Diospyros leaf-miner, Gracilaria [sic] (= Caloptilia) mabaella from Oahu and did not mention the unnamed Philodoria leaf miner on Diospyros. Zimmerman (1978a) noted that Caloptilia mabaella might be an introduced species from Japan or North America. We observed the P. lama and Caloptilia mabaella (CJ333) (Fig. 73D), occurring sympatrically at Kanepuu, Lanai. The mine of Caloptilia is distinguished from that of P. lama by the brown in coloration and oval cocoon which is situated at the end of the mine., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 43-46, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Johns, C. A., Toussaint, E. F. A., Breinholt, J. W. & Kawahara, A. Y. (2018) Origin and macroevolution of micro-moths on sunken Hawaiian Islands. Proceedings of the Royal Society B, 285, 20181047. https: // doi. org / 10.1098 / rspb. 2018.1047","Walsingham, Thomas, de Grey (Lord) (1907) Microlepidoptera. In: Sharp, D. (Ed.), Fauna Hawaiiensis. 1 (5). Cambridge University Press, Cambridge, pp. 469 - 759.","Swezey, O. H. (1940) New species of Hawaiian Lepidoptera. Proceeding Hawaiian Entomological Society, 10 (3), 461 - 465.","Swezey, O. H. (1946) New species of Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 12 (3), 625 - 628.","Meyrick, E. (1928) Some new species of Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 7 (1), 104.","Swezey, O. H. (1910 a). The ebony leaf-miner (Gracilaria [sic] mabaella). Proceedings of the Hawaiian Entomological Society, 2 (3), 88 - 90, pl. 3.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718."]}

Philodoria costalis Swezey, 1934 Figs. 10A, 57A, B, 61I–N. Philodoria costalis Swezey, 1934: 524–525; Zimmerman 1978a: 708, figs. 462, 469, 476. Type locality. Makaha ridge, 3000 ft, west side of Mount Kaala (Oahu). Type material. Lectotype ♂, Makaha ridge, 3000 ft, west side of Mount Kaala, Waianae Mountains, Oahu, 15.iv.1934, O.H. Swezey Collecter, [host:] Pipturus, | Holotype | Philodoria costalis Sewezey Det. by O. H. Swezey| Type no. 4269 in BPBM (here designated). Paralectotypes 2♂, 2♀, same locality and data as lectotype, | Paratype | 34153/Z-XII-20-60-2♂, SK 652♀ |♂ paratype to ECZ 7-17-61[= 17.vii.1961]| BPBM 34154, SK683♂| BPBM 34155, SK 653♀ | BPBM 34156. Described from presumably six specimens that Swezey (1934) examined, reared from numerous mines on Pipturus. One specimen was labeled ‘holotype’ and five others as ‘paratypes’. Given the manner in which the descriptions were written, Swezey likely considered these specimens to be the holotype and paratypes. But as a holotype was not specified in the description, the so-labeled holotype and paratypes are all to be considered syntypes under Article 73.2 of the Code (ICZN 1999), and any one is thus eligible for designation as lectotype under Article 74 of the Code (ICZN 1999). The ‘ Holotype, type no. 4269’, which Swezey indicated as holotype in the specimen label, is here designated as lectotype (Fig. 10A). The remaining syntypes are paralectotypes. Diagnosis. Swezey (1934) noted that this species is very similar to Philodoria micropetala Walsingham, but it is distinguishable by the white costal line and the lack of fuscous scales along middle of apical orange area (Fig. 10A). The forewing pattern and genitalia are similar to that of P. pipturiella Swezey, but distinguished by the rather oblong forewing with a small white spot at middle (Fig. 10A); the straight base of valva; and a rather short ductus bursae. Redescription: Adult (Fig. 10A). Wingspan 7–8 mm; forewing length 3.5 mm in lectotype, 2.4–3.0 mm in paralectotypes. Head tawny brown; frons white; maxillary palpus reduced; labial palpus white shaded with brownish externally on apex of median joint and most of the terminal joint. Antenna uniformly light brown, about 1.25 x length of forewing. Thorax and abdomen tawny brown, white beneath. Forewing tawny brown, suffused with orange patches at apical portion; a broadly oval to nearly circular white spot (ws) about middle or just beyond middle of fold, not reaching dorsum; a narrow white line (cl) on costa from base to about 2/3; white costal spot (cs) at 2/3, this line enclosing fuscous scales; two white bands on both extremities of shiny pale blue color band (bb 3 ): w 1 at costal 1/4, bent w 2 near opposite at dorsal 1/4; another blue apical spot (as) extending into apical cilia; costal cilia with two white spots (a, b), rest of apical and the terminal cilia brown. Hindwing and cilia brownish fuscous. Legs light brown, white beneath. Male genitalia (Fig. 61J–N). See Zimmerman (1978a: fig. 469). Tegumen 0.8–0.9 x length of valva; valva rather short, tapering along costal margin from 1/2 to apex with short spines at basal region; apical half digitiform, near straight; along the inner side of valva, a series of small spines (Fig. 61J, L). Saccus short and tapering toward the point (Fig. 61K). Phallus rather short and thick with two series of minute cornuti in vesica (Fig. 61M). Female genitalia (Fig. 57A, B) (n=2). 900 µm long. Ostium bursae large; antrum cup-shaped with a pair of slender lateral lobes; lamella antevaginalis 160 µm, weakly sclerotized, widening toward anterior margin of A7. Ductus bursae 300–350 µm; terminus of ductus bursae biforked; corpus bursae oblong, 520 µm; paired rows of wrinkles running longitudinally, some sclerotized. Distribution. Oahu (Swezey 1934). Host plants. Urticaceae: Pipturus sp. (Swezey 1934). Biology. Unknown, other than that an oval brown cocoon has been reported to be situated on the surface of a leaf (Swezey 1934)., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 46-48, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Swezey, O. H. (1934) New species of Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 8 (3), 523 - 525.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","ICZN (1999) International code of zoological nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 306 pp.","Swezey, O. H. (1915) New species of Hawaiian moths. P roceedings of the Hawaiian Entomological Society, 3 (2), 94 - 97. https: // doi. org / 10.5962 / bhl. part. 24594"]}

Philodoria splendida Walsingham, 1907 Figs. 9A, B, 26D, E, 41J, K, 42I, J, 43I, J, 56K, 64G–I, 66C, D, 73B, C, 82. Philodoria splendida Walsingham, 1907: 719–720, pl. 25, fig. 23; Swezey 1913b: 223; Zimmerman 1978a: 718, figs. 466, 471, 477, 481. Type locality. Lanai, 2000ft. Type material. Lectotype ♂, Lanai, 3000ft, ii.1894, Perkins 26452 |♂ genitalia on slide 101, Oct.17.1940| Philodoria splendida ♂ Wlsm PARATYPE 2/9| Fauna Hawaiiensis Collection| No. 34284 Hawaiian Coll. BISHOP Museum in BPBM (here designated). Paralectotype 2 (1♂, 1 sex unknown). 1 (sex unknown), Lanai, 2000ft, vii.1894, Perkins 26493 | Philodoria splendida Wlsm PARATYPE 3/9| in BPBM; 1♂, Kauai Mts., 3000–4000ft, vi.1894, Perkins 27188, | Philodoria splendida ♂ Wlsm PARATYPE 9/9| in BPBM. Described from ten specimens: ‘type ♂ (26430) and nine ‘paratypes’ from Kauai, Oahu, Molokai and Lanai. Given the manner in which the descriptions were written, Walsingham likely considered them as the holotype and paratypes, as indicated on their specimen labels. However, because a holotype was not specified in the description, the so-labeled types and paratypes are to be considered syntypes under Article 73.2 of the Code (ICZN 1999), and any one of them is thus eligible for designation as lectotype under Article 74 of the Code (ICZN 1999). We could not find ‘type ♂ (26430)’ from 2000 ft., Lanai, xii.1894 and six syntypes: 1 (sex unknown), Halemanu, 4000 ft, Kauai, v.1895; 1 (sex unknown), Waianae Mts., 2000 ft., iv.1892, beaten from “Lefua” (Metrosideros polymorpha ?); 1 (sex unknown), Mts., near Honolulu, 2000 ft., 25.x.1892; 1 (sex unknown), Kalae, 1500 ft., v.1893; 1 (sex unknown), Mts., about 4000 ft., 9.vi.1893; 1 (sex unknown), 3000 ft., Lanai, ii.1894. Zimmerman (1978a: 701, fig. 466) provided the last photograph for the ‘holotype’ (=type ♂ (26430)) in the NHMUK. The current curator of NHMUK, D.C. Lees informed us that the ‘holotype’ of P. splendida is not stored in the designated location of the NHMUK. We selected a lectotype from three syntypes (Fig. 9A), and the remaining syntypes are paralectotypes. Additional material. 28 (14 ♂, 11 ♀, 3 sex unknown). Kauai: 1♂, Kauai Na Pali, Kona Forest Reserve, Alakai Swamp trail, 3800’, 29ix.1973, K. & E. Sattler, B.M.1973-498, Philodoria sp. 12 (Kauai) Sattler Coll., color slide 140, D. C. Lees Sep. 2016, BMNH(E)1621068. Oahu: 1♂, Waianae, 5.i.1913, ex Metrosideros polymorpha, SK 697♂, BPBM 34316; 1♂, Waianae, 5.i.1913, ex Ohia (= M. polymorpha) in USNM;1♂, 1♀, Lauiluli, 25.x.1914, “Ohia Lehua” (= M. polymorpha), BPBM 34312; 3♂, 1♀, Malamalama, 8.x.1916, “Ohia Lehua”, 34293–34295, ♂ ♀ slide aB Aug26 1941 in USNM; 1♀, Konahuanui, 2300ft, 9.v.1943, Resting on Cladium, EC Zimmerman Collector, BPBM 34319; 1♀, Kalihi, Coll. O.H.S, “Ohia Lehua”, BPBM 34311; 1 (sex unknown), Olympus, Coll. O.H.S, no. “35” in USNM.1♂, Waialae, Nui, 16.ii.1919, Z-XII-19-62-20♂ in BPBM. 1 (sex unknown), Kaala Road, 23.iv.2013 (stored), C.A. Johns leg., Adult on Melicope clusiifolia, 21.iv.2013, CJ049, KT 982402 in BPBM; 1♂, 2♀, Kanahuanui, 30.i.& 4.ii.2014 (stored), C.A. Johns leg., host: M. polymorpha, 3.i.2014, CJ246ab, CJ247, SK780♂, 781♀, 773♀ in BPBM. In NHMUK: 1♂, Oahu 4000’, Waianae Range, Ma Kaala, 4.ix.1976, K. & E. Sattler, B.M.1976-605, Philodoria sp. 10D (Lanai, Mauai, Oahu) splendida ? Sattler Coll., D.C. Lees, Sep. 2016, BMNH(E)1621292; 1♀, Oahu 2200’, Waianae Range, above Makua Valley, 11.ix.1976, K. & E. Sattler, B.M.1976- 605, Philodoria sp. 10C cf lysimachiella (Oahu) Sattler Coll., D. C. Lees. Sep. 2016, BMNH(E)1621295. Lanai: 1♂, 2750’, Munro Trail, 2.x.1976, Philodoria sp. 10D (Lanai, Mauai, Oahu) splendida? Sattler Coll., D. C. Lees Sep. 2016, BMNH(E) 1621678 in NHMUK; Maui: 1♂, Haelaau, 17.xii.1928, O.H. Swezey, 34289 in BPBM; 1♂, 1♀, 1 (sex unknown), Flemming Arboretum Trail, [3.v.2013 (stored), C.A. Johns leg., host: M. polymorpha, 12.iv.]2013, SK 621♀, SK696♂, CJ105 (sex unknown) / KT 982410 in BPBM. 1♀, W. Mauai, 3000’, W. Maui Forest Res., Kaulalewelewe (Puu Kukui Trail), 26.viii.1976, K. & E. Sattler, B.M.1976-605, Philodoria sp. 10D (Lanai, Mauai, Oahu) splendida? Sattler Coll., D. C. Lees Sep. 2016, BMNH(E) 1621208 in NHMUK; Syzygium sp. , C.A. Johns leg., in BPBM: 2♀, Limahuli, Kauai, 21.vii.2015 (stored), 20.vi.2015, CJ488ab, SK 748♀, SK 752♀; 1♂, Waiaai Trail, Oahu, 30.vii.2014 (stored in RNAlater solution), 5.vii.2014, CJ312, SK747♂ (only abdomen). Diagnosis. This species and P. lama sp. n. have a brownish ocherous basal oblique fascia on forewing (Figs. 9A, B, 73B, C). Genital features very similar to those of P. basalis and P. lama, but distinguished by a rather slender dorsal process of valva in male. Kauai specimens (CJ488, male unknown) on Syzygium sp. have similar forewing pattern (Fig. 26D, E) and female genitalia, but the Oahu specimen (CJ312, thorax sacrificed for molecular analysis) have thickened dorsal process with slightly curved dorsal side (Fig. 41K) and slender saccus (Fig. 42J). Redescription: Adult (Figs. 9A, B, 26D, E, 73B, C). Wingspan 8–9 mm in type series; forewing length 4 mm in holotype, 3.9 mm in paratype. Head dark brown; frons white; maxillary palpus reduced; labial palpus white, externally fuscous at apical portion. Antenna grayish fuscous. Thorax leaden gray. Forewing shining, pale leaden gray with dark brownish ocherous patches enclosing gray fascia: bp from near base of costal fold to dorsal 1/4; tf from costal 1/3 to dorsal 1/2, tp after the middle to apical portion, distinctly narrowing in the dorsum, containing two short, shining, steel gray costal spot; apical portion with large jet-black terminal patch from tornus to apex, containing two upright steel-gray streaks near its opposite extremities; a single bright silvery dot in its middle; a very narrow ocherous line along the base of tornus; cilia shiny, leaden gray with a black fringe line; two white costal lines, long one extend apex; terminal cilia often black. Hindwing coppery brown; cilia tawny. Abdomen tawny brown, silver below. Legs grayish fuscous, with white spots on tarsal joints and at distal end of tibiae; spurs white. Male genitalia (Figs. 41J, K, 42I, J, 43I, J, 64G, I) (n=3). Capsule 830 µm. Tegumen same length as valva and rather wide; valva 490 µm long, similar to P. basalis except dorsal process rather slender (Fig. 41J). Saccus broad and straight (Fig. 42I). Phallus 660 µm long, sinuous in lateral view; cornuti large and well developed apically (Fig. 43I). Female genitalia (Fig. 56K) (n=4). 1400 µm long. Lamella antevaginalis 200 µm in length. Corpus bursae 800 µm long. Similar in the appearance to P. basalis. Distribution. Kauai, Oahu, Molokai, Lanai (Walsingham 1907), and Hawaii (Big Island) (Zimmerman 1978a). Host plants. Myrtaceae: Metrosideros polymorpha Gaudich. (Walsingham, 1907), Syzygium sp.: new record. Biology. (Figs. 66C, D, 82). Swezey (1913b: 223) reported the “taco” like cocoon and mature larval habit, which are similar to that of P. basalis. The biology is poorly studied, but it is presumably similar to that of P. basalis. In general, larvae form slender linear mine on the adaxial leaf surface (Fig. 82E), which gradually expands as lavae feed and grow (Fig. 82A). One or two mines per leaf. We often observed the fallen mined leaves under the host plant (Fig. 66C, D) with late instar larvae in the mine and/or pupae in a taco cocoon on or near the leaf. Parasitoids. (Fig. 101G, H). Bethylidae: Sierola philodoriae Fullaway, 1920; Eulophidae: Pauahiana metallica Yoshimoto, 1965 (Zimmerman 1978a). Remarks. We identified two adult moths (Coll ID CJ-049 / GenBank accession no. ID KT982402 and CJ-105 / KT982410) as P. splendida, based on the presence of a brown basal oblique fascia on forewing and host plant, from which whole bodies were sacrificed for molecular analysis (Johns et al. 2016)., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 39-42, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Walsingham, Thomas, de Grey (Lord) (1907) Microlepidoptera. In: Sharp, D. (Ed.), Fauna Hawaiiensis. 1 (5). Cambridge University Press, Cambridge, pp. 469 - 759.","Swezey, O. H. (1913 b) Leaf-miners of the Hawaiian Islands. Proceedings of the Hawaiian Entomological Society, 2 (5), 221 - 227.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","ICZN (1999) International code of zoological nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 306 pp.","Swezey, O. H. (1928) Some new species of lepidopterous leaf-miners in Hawaii. Proceedings of the Hawaiian Entomological Society, 7 (1), 187 - 191.","Swezey, O. H. (1946) New species of Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 12 (3), 625 - 628.","Yoshimoto, C. M. (1965) Synopsis of Hawaiian Eulophidae including Aphelininae (Hym.: Chalcidoidea). Pacific Insects, 7 (4), 665 - 699.","Johns, C. A., Moore, M. R. & Kawahara, A. Y. (2016) Molecular phylogeny, revised higher classification, and implications for conservation of endangered Hawaiian leaf-mining moths (Lepidoptera: Gracillariidae: Philodoria). Pacific Science, 70 (3), 361 - 372. https: // doi. org / 10.2984 / 70.3.7"]}

Philodoria keaensis Kobayashi, Johns & Kawahara, sp. n. Figs. 22A, B, 59K. Type locality. Mauna Kea (Big Island). Etymology. The specific epithet is derived from the type locality, Kea (pronounced ‘Keh-ah’). Type material. Holotype ♀, Mauna Kea, Hale Pohaku, 9,970ft (~ 3,038 m), Hawaii Is. [Big Island], 9.viii.2014 (stored), J. Eiben leg., adult on Tetramolopium humile, 27.vi.2012, CJ355 / SK 777♀ in BPBM. The holotype was mounted as a dry pinned specimen by placing two forewings without mountant under a coverslip. Diagnosis. Distinguished from other species by having a broad white streak at its costal fold and a transverse streak at 3/4 on the forewing (Fig. 22A, B). Philodoria wilkesiella Swezey and P. touchardiella (Swezey) have female genitalia that are similar in shape to P. keaensis sp. n., but the former two are separated from the latter by a rather broad or long antrum and distinct forewing pattern. Description: Adult (Fig. 22A, B). Forewing length 2.5 mm in holotype. Head and frons dark brown with white scales on lateral side; maxillary palpus white; labial palpus white above, brownish beneath. Thorax dark brown. Forewing brown to pale redish brown with white patches: a costal broad one along costal fold from base to costal 1/2; a dorsal broad outwardly oblique ds 2 at 1/2 enclosing fuscous lines, connecting white line extending dorsal base of wing along dorsal margin; a white transverse fascia at 3/4; a large black patch nearby transverse streak with shiny blue scales at center; apical portion orangish brown with a costal white spot (b) at apex of cilia; an apical black patch (as) with shiny blue scales; apical and terminal cilia fuscous, tornal cilia brown. Hindwing and cilia fuscous. Abdomen white below. Legs dark brown, tarsi spotted with white. Male genitalia Unknown. Female genitalia (Fig. 59K) (n=1). 960 µm long. Ostium bursae large; antrum cup-shaped with a pair of lateral lobes; lamella antevaginalis 160 µm, weakly sclerotized, semicircular, widening toward anterior margin of A7. Ductus bursae short, middle part weakly sclerotized, round and flat; terminus of ductus bursae tubular, biforked. Corpus bursae 570 µm, pyriform; anterior end of corpus bursae weakly sclerotized; paired rows of longitudinal, partly sclerotized wrinkles; very small paired signa with a pair of slender minute spines. Distribution. Hawaii (Big Island). Host plants. and Biology. Unknown. Remarks. Jesse Eiben collected the holotype from beating the plant, Tetramolopium humile (A. Gray) Hillebr. (Asteraceae) at Mauna Kea, 9,970ft (~ 3,038 m), Hawaii Island (Big Island). We returned to this location in October 2014 and searched Tetramolopium humile, Argyroxiphium sandwicense DC. subsp. sandwicense, Dubautia ciliolata (DC.) D. D. Keck, and D. linearis (Gaudich.) D. D. Keck but none of the plants we checked had visible leaf mines. We also set blacklight bucket trap lights for two nights within five feet of these plants, but were unable to collect any Philodoria adults. Some leaves of A. sandwicense subsp. sandwicense had its layers of leaf tissue separated, roughly resembling the pockets that form within mined leaves, but we were unable to find any frass. We therefore do not consider these separations as damage created by Philodoria larvae. Jesse Eiben has been conducting arthropod surveys in this area for nearly a decade. Given the rarity of this Philodoria species despite years of comprehensive arthropod sampling in the area, it is possible that this insect does not belong here, but instead was windblown from elsewhere as is often seen with arthropods on Mauna Kea. Future surveys focused specifically on this insect would benefit from sampling known Philodoria host plants and where they are more abundant, at a lower elevation.

Philodoria kolea Kobayashi, Johns & Kawahara, 2018 Figs. 7F, 34B, C, 35, 41D, 42D, 43D, 56F, 79I���K. Philodoria kolea Kobayashi, Johns & Kawahara, 2018: 132���137, 3, 5K, L, 7A���D, I, 12, 13, 14D Type locality. Hawaii Volcanoes National Park (Big Island). Type material. Holotype ♂, Hawaii Volcanoes National Park, Hawaii (Big Island), 25.iv.2016, A. Kawakita leg., host: Myrsine lessertiana (understory shrub), Gen-Bank accession no. MF804825, IO-322, SK 851 in BPBM. The type series was mounted from emerged adult moths. Paratypes, in BPBM: 1♀, Kaumana Trail, Hilo, Hawaii (Big Island), 28.iv.2016, em., C.L. - Vaamonde & C. Doorenweerd leg., host: Myrsine sp., 20.iv.2016 (Cocoon), HILO016, SK 634♀. 1♀, Thurston lava tube (Nahuku), Hawaii Volcanoes National Park, Hawaii Is., 13.v.2016, em., S. Kobayashi leg., host: Myrsine lessertiana, 25.iv.2016 (larva), SKH-05-1, SK 632♀; 1♀, same locality and data as holotype, IO-323, SK852; 2♀, same locality as holotype, 2& 24.v.2016, em., C.L. - Vaamonde & C. Doorenweerd leg., host: Myrsine lessertiana, 22.iv.2016 (larva), HILO020/SKH-15, SK 630♀, 631♀. Diagnosis. Among Philodoria species having similar fuscous forewing coloration (i.e., P. epibathra (Walsingham), P. nigrella (Walsingham) P. pipturiana Swezey, and P. wilkesiella Swezey (See Zimmerman 1978a)), P. kolea is distinguished by the white and bronze color bands on the forewing (Fig. 7F). The forewing pattern and genitalia are similar to those of other Myrsine mining species, P. auromagnifica and P. succedanea, but P. kolea completely lacks the orange markings (Fig. 7F). Adult (Fig. 7F, 34B, C, 35). Wingspan 6.7 mm in holotype, 6.6, 8.5 mm in paratypes. Male genitalia (Figs. 41D, 42D, 43D) and Female genitalia (Fig. 56F). See also Kobayashi et al. (2018). Distribution. Hawaii (Big Island) (Kobayashi et al. 2018). Host plants. Primulaceae: Myrsine lessertiana A. DC. (Kobayashi et al. 2018). Biology. (Figs. 79I���K). Kobayashi et al. (2018: 136���137, figs. 12, 13, 14D) reported the leaf mine and pupal cocoon., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 27-28, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Kobayashi, S., Johns, C. A., Lopez-Vaamonde, C., Doorenweerd, C., Kawakita, A., Ohshima, I., Lees, D. C., Hanabergh, S. & Kawahara, A. Y. (2018) Hawaiian Philodoria (Lepidoptera, Gracillariidae, Ornixolinae) leaf mining moths on Myrsine (Primulaceae): two new species and biological data. ZooKeys, 773, 109 - 141. https: // doi. org / 10.3897 / zookeys. 773.21690","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718."]}

Philodoria lipochaetaella (Swezey, 1940) Figs. 23D–G, 24A. Parectopa lipochaetaella Swezey, 1940: 464. Philodoria lipochaetaella (Swezey, 1940); Zimmerman 1978a: 681, figs. 439, 442, 449, 455. Type locality. A gulch above Lahaina (Maui). Type material. Lectotype ♂, Gulch above Lahaina, Maui, 28.viii.1929, O.H. Swezey Collector, host: Lipochaeta larvarum, | Holotype Parectopa lipochaetaella Swezey |Type no. 4260 in BPBM. Paralectotype 7 (1♂, 2♀, 4 sex unknown): 1♂, 1♀, 1 (sex unknown), same mount of lectotype in BPBM, 1♀, 3 (sex unknown), same data and locality as lectotype, SK 700♀ | BPBM34187. Four paralectotypes missing. This species was described from 12 specimens from Maui. Zimmerman (1978a) designated Type no. 4260 (upper right of four syntypes) on a mount as lectotype (Fig. 23E), and the remaining syntypes are paralectotypes (Fig. 23D, F, G). Diagnosis. Very similar to P. sciallactis (Meyrick), but can be distinguished by its round valva (Zimmerman 1978a: fig. 449). Redescription: Adult (Figs. 23D–G, 24A). Wingspan 5–6 mm in type series; forewing length 3 mm in lectotype, 2.5–3.0 mm in paralectotype. Head and frons white; maxillary palpus white; labial palpus white with a black spot externally at apex of second, and third segments. Antenna white beneath, ringed with fuscous above. Thorax white. Forewing pale ocherous with white patches: a narrow white median line at basal half of fold; widely white basal patch (ds 1 + ds 2 ) extending inwardly and outwardly oblique to meet cs 1 at middle; a dorsal patch (ds 3 ) from 1/2 to 3/4, almost connecting with cs 2 and cs 3 ; three outwardly oblique costal ones (cs 1–3 ), cs 1 from base to 1/2, cs 2 at 3/4, cs 3 at 4/5; three white costal spots (a, b, c) near apex; apical portion white centered with a patch of pale blue scales (as); all white areas with a few marginal fuscous scales; cilia white with basal line of fuscous scales and a fuscous line near scale tips. Hindwing and cilia pale gray. Legs white, fore and middle legs and posterior tarsi with black spots. Abdomen white, anal tuft pale ocherous to white. Male genitalia. See Zimmerman (1978a, fig. 449). Tegumen 0.7–0.8 x length of valva; valva digitiform and slightly narrowing at middle, rounded at apex, slightly curved toward dorsal side, similar to P. hauicola. Saccus slender and needle-shaped in ventral view. Phallus not examined. Female genitalia. The female genitalia are very similar to that of P. sciallactis. See Zimmerman (1978a, fig. 455). Distribution. Maui (Swezey 1940 b). Host plants. Asteraceae: Lipochaeta lavarum (Gaudich.) DC. (Swezey 1940 b). Lipochaeta could not be found during our surveys of West Maui. Biology. Unknown. Remarks. We could not find the male specimen and male genitalia slide no. Z-I-27-61-3 of Zimmerman’s figs. 449 and 451. Swezey (1940, 1946) described this species twice from two series, collected from the same locality and same host plant in 1928 and 1929. Zimmerman (1978a) placed Parectopa lipochaetae Swezey, 1946 (= the specimens series in 1928) in synonymy with P. lipochaetaella, but P. lipochaetae is synonymous with P. sciallactis based on the male genitalia. Zimmerman (1978a) also noted in his taxonomic key that P. sciallactis has a subbasal oblique white costal streak (cs 1 ) which continues along the costa to or close to the wing base and the second oblique white costal streak (cs 2 ) is discontinued distinctly across wing. However, the forewing alone cannot be used to distinguish the two species. We treat the type series from 1929 and the male specimens that have a rounded valva as P. lipochaetaella., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 103-104, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Swezey, O. H. (1940) New species of Hawaiian Lepidoptera. Proceeding Hawaiian Entomological Society, 10 (3), 461 - 465.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","Swezey, O. H. (1946) New species of Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 12 (3), 625 - 628."]}

Philodoria platyphylliella Kobayashi, Johns & Kawahara sp. n. Figs. 22D, 50I, 51G, 52G, 75D–G, 87G–I. Philodoria sp. 14; Johns et al. 2018: figs. 1A, 2. Type locality. Nakula Natural Area Reserve (Maui). Etymology. The specific epithet, platyphylliella, refers to the species epithet of Dubautia platyphylla, its host plant. Type material. Holotype ♂, Nakula Nar, Maui Is., 11.viii.2014 (stored), C.A. Johns leg., host: Dubautia platyphylla, 26.vii.2014, CJ363, SK833♂ in BPBM. Paratype ♂, same locality and data as holotype, 10.viii.2014 (stored), CJ358, PHIL0026 (only abdomen) in BPBM. The holotype was mounted as a dry pinned specimen by placing two forewings without mountant under a coverslip. The head, antenna, thorax, and legs of paratype were sacrificed for molecular analysis. Additional material. 1♀, same locality and data as holotype, 12.viii.2014 (stored), CJ368, PHIL0025 genitalia missing in BPBM. Diagnosis. The forewing pattern similar to that of P. nigrelloides (Swezey) and P. touchardiella (Swezey), but is differs from them by having a darker brown forewing with a blue to fuscous patch at its apex (Fig. 22D). The male genitalia is similar to that of P. touchardiella, but is distinguished by its rather thick saccus. Description: Adult (Fig. 22D, 75D–G). Forewing length 3.4, 3.6 mm in holotype. Head bronze; frons white; maxillary palpus pale bronze; labial palpus white basal half, bronze at apex; Antenna bronze, about 1.1–1.2 x length of forewing. Thorax dark brown. Forewing dun to bronze with three oblique white streaks: ds 2 at about 2/5, extends to about middle of wing; a smaller ds 3 at 2/3. a small costal spot at near base; cs 3 from 3/4, extending about 2/3 across wing; apical portion dark orangish brown with a large as of blue scales; a blue to fuscous patch at 4/5; few shiny gray spot at the middle of apical portion; cilia fuscous mixed dark brown scales with three white costal spots (a, b, c) at apex; black bl 1 at terminal cilia. Hindwing and cilia fuscous. Abdomen bronze, white beneath. Legs fuscous. Male genitalia (Figs. 50I, 51G, 52G) (n=2). Capsule 800 µm. Tegumen 0.9 x length of valva; valva 590 µm long, tapering along costal margin from basal 1/2 to apex (Fig. 50I). Saccus digitiform in ventral view (Fig. 51G). Phallus 500 µm long and straight, slightly widening at phallobase with developed coecum; cornuti in vesica indistinct (Fig. 52G). Female genitalia Unknown. Distribution. Maui. Host plants. Asteraceae: Dubautia platyphylla (A.Gray) D.D.Keck. Dubautia platyphylla is rare and restricted to Maui. We surveyed a small population of this plant species within Nakula Natural Area Reserve, Maui, in July 2014 (Fig. 69C, D). This plant population was located on a steep drainage and was out of reach of ungulates, goat, pig, and deer. Biology. (Fig. 87G–I). Larvae form an oblong blotch mine (Fig. 87G–I). Pupation occurs in the white cocoon on the adaxial leaf surface (Fig. 87I)., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on page 101, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Johns, C. A., Toussaint, E. F. A., Breinholt, J. W. & Kawahara, A. Y. (2018) Origin and macroevolution of micro-moths on sunken Hawaiian Islands. Proceedings of the Royal Society B, 285, 20181047. https: // doi. org / 10.1098 / rspb. 2018.1047"]}

Philodoria obamaorum Kobayashi, Johns & Kawahara, sp. n. Figs. 19C–E, 27K–M, 50C, D, 51B, 52B, 59C, D, 63C–G. Gracilaria neraudicola Swezey, 1920: 385–386. Parectopa neraudicola Swezey, 1920; Swezey 1928: 191. Philodoria neraudicola (Swezey, 1920); Zimmerman 1978a: 689, figs. 449, 458. Philodoria sp. 16; Johns et al. 2018, fig. 2. Type locality. Pahoa (Big Island). Etymology. The specific epithet, obamaorum, is named after Barack Hussein Obama II, the 44th president of the United States, born in Honolulu, Hawaii, and his wife, First Lady Michelle LaVaughn Robinson Obama. They have have continued to support nature conservation efforts in Hawaii. Following recommendation 31.1.2. of the International Code of Zoological nomenclature (ICZN, 1999) the specific name is treated as a noun in the genitive plural case. Type material. Holotype ♂, Pahoa, Puna, Hawaii (Big Island), 20.ix.1918, O.H. Swezey, host: Pipturus albidus in USNM. Paratype 14 (4♂, 3♀, 7 sex unknown): 1♂, 1♀, same locality and data as holotype, SK 710♀, in USNM; 3♂, 2♀, 2 (sex unknown), S. of Hilo, Hawaii, 25.vii.1921, Swezey coll., Pipturus 34248, SK709♂, BPBM34242, no. 4257 |Z-1-26-61(3) ♂, Z-V-18- 61♀; 2 (sex unknown), Judd Trail, Hawaii, 13& 14.viii.1919, OHS, Pipturus, BPBM 34251, BPBM34252. Additional material. 4 (1♂, 3♀) 1♂, 2♀, Mauna Ulu Havo, Hawaii, 21.v.2015 (stored), C.A. Johns leg., host: Pipturus sp., 3.v.2015, CJ423 / SK 844♀, 403M / SK736♂, 403O(b-2) / SK 739♀ in BPBM; 1♀, Upper Hamakua Ditch trail, Hawaii, 15.v.2016 em., C.L.-Vaamonde & C. Doorenweerd leg., host: Pipturus sp., 24.iv.2016 (cocoon), CLV0017, SK 823♀ in BPBM; 2♀, Puu Makkaala Natural Reserve, Hawaii, 9.vi.2015 (stored), C.A. Johns leg., host: Pipturus sp. (B), 25.v.2015, CJ423 / SK 844♀, CJ 427 in BPBM. In NHMUK: 1♂, Hawaii 2800’, Kau District, Ocean View, 23.vi.1976, K. & E. Sattler, B.M.1976-605, Philodoria sp. 25 cf marginestrigata (Hawaii) Sattler Coll., D.C. Lees Sep. 2016, BMNH(E)1621262; 1♀, Hawaii 1300’, S. Hilo District, Upper Waiakea F. R., Stainback Highway, 12.vii.1976, K. & E. Sattler, B.M.1976-605, Philodoria sp. 21 (Hawaii) Sattler Coll., D.C. Lees Sep. 2016, BMNH(E)1621128. Diagnosis. Distinguished from P. neraudicola by having a triangular white patch at 3/4 and two white dorsal streaks that are nearly the same width as in the forewing (Fig. 19C, D); saccus and coecum half the length of those of P. neraudicola (Figs. 51B, 52B). Description: Adult (Fig. 19C–E). Wingspan 7.5 mm in holotype, 7.0– 7.5 mm in paratype; forewing length 3.3, 3,4 mm in holotype, 3.0– 3.5 mm in paratype. Head creamy white; frons white mixed with fuscous scales; maxillary palpus white mixed with fuscous scales; labial palpus white, median joint fuscous at apex, terminal joint with fuscous band at middle and at apex. Antenna brownish, basal segment white. Thorax brownish fuscous, pale in middle and posteriorly. Forewing brownish fuscous with white patches: two outwardly-curved white dorsal streaks: ds 1 at 1/4, slightly widened at base, ds 2 at 1/2, almost same width; a triangular white patch (ds 3 ) at 3/4, not reaching the middle of wing; a white slender outwardly-oblique cs 3 at 3/5; all streaks and patch margined with a few black scales; apical portion orangish ocherous; a round black as with a few pale blue scales, a few pale blue scales between this spot and ds 3 ; cilia brown with three white costal spots; apical and terminal cilia paler near base with black dl 1 , tornal cilia very paler at apex. Hindwing and cilia pale brown. Abdomen pale brownish, white below. Legs ocherous, anterior and middle tibiae fuscous, and middle and posterior tarsi fuscous. Male genitalia (Figs. 50C, D, 51B, 52B, 63E, F) (n=3). Capsule 700 µm. Tegumen 0.9 x length of valva; valva 480–500 µm long, tapering along costal margin from 2/5 to apex, similar to P. neraudicola (Fig. 50C, D). Saccus rather short and pointed at apex in ventral view (Fig. 51B). Phallus 520 µm long and nearly straight with developed coecum; two series of minute cornuti in vesica (Fig. 52B). Female genitalia (Figs. 59C, D, 63C) (n=5). Similar to P. neraudicola. 890–1180 µm long. Lamella antevaginalis 200 µm. Corpus bursae 570 µm. Distribution. Hawaii (Big Island). Host plants. Urticaceae: Pipturus albidus A.Gray ex H.Mann (Swezey 1920), and Pipturus sp. Biology. C.L.-Vaamonde and C. Doorenweerd collected larvae from leaves of Pipturus species in 2016. Larvae formed blotch mines and made six cocoons on the adaxial leaf surface; the orange oval cocoon is 6.0 mm in length and 2.5 mm in width. Two adults emerged from the cocoons, one was P. obamaorum and another P. floscula Walsingham. Remarks. This species and other Pipturus miners occur together on Hawaii (Big Island). It is unclear which species of Pipturus these moths were on at Hawaii Volcanoes National Park, the site that was surveyed in 2016. Some Philodoria species may prefer certain species of Pipturus, but the determination of host preference is impeded by the unsettled taxonomy of this host plant genus. Future studies of these moths should focus on rearing different species of Philodoria from Pipturus to understand host plant specificity., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 86-87, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Swezey, O. H. (1920) Some new Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 4 (2), 376 - 386. https: // doi. org / 10.5962 / bhl. part. 16149","Swezey, O. H. (1928) Some new species of lepidopterous leaf-miners in Hawaii. Proceedings of the Hawaiian Entomological Society, 7 (1), 187 - 191.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","Johns, C. A., Toussaint, E. F. A., Breinholt, J. W. & Kawahara, A. Y. (2018) Origin and macroevolution of micro-moths on sunken Hawaiian Islands. Proceedings of the Royal Society B, 285, 20181047. https: // doi. org / 10.1098 / rspb. 2018.1047","ICZN (1999) International code of zoological nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 306 pp."]}

Philodoria urerana (Swezey, 1915) Figs. 20A, B, 33G, 50E, 51C, 52C, 59E, 78E, F, 95A–C. Gracilaria urerana Swezey, 1915: 95–96. Parectopa urerana (Swezey, 1915); Swezey 1928: 191. Philodoria urerana (Swezey, 1915); Zimmerman 1978a: 691, figs. 446, 453, 460. Type locality. Mt. Tantalus (Oahu). Type material. Lectotype ♀, Mt.Tantalus, Oahu, 16.iii.1915, O.H. Swezey, ex Urera [sandwicensis], |TYPE OF 214 Gracilaria urerana Swezey | top one of two cotypes of the same type mount designated by Zimmerman (1978a, 691, fig. 446) in BPBM (Fig. 20A). Paralectotype 1 (sex unknown), bottom one of the same mount of lectotype (Fig. 20B). 17 including six ‘syntypes’ (2♂, 6♀, 9 sex unknown): 2♂, 4♀, 6 (sex unknown), same data and locality as lectotype, SK671♂, 672♂, 673♀ | BPBM 34363, BPBM 34634 (“ex blotch mine”), BPBM 34365– BPBM 34369; 2♀, 3 (sex unknown), Same label, “ex serpentine mine” (3 sex unknown) in USNM. This species is described from eight specimens reared from Urera sandvicensis and from the same lot of leaves as those collected for P. ‘ ureraella ’. We examined the lectotype and 18 specimens in BPBM and USNM that bear the same data as the lectotype. Additional material. Oahu: 1♂, Waianae, 28.iii.2016 (stored), K.Bustamente leg.,host: Urera glabra 9.xi.2015, KMB02, CJ533, SK670♂ in BPBM; 1♂, 1♀, Kaau Crater, 27.vii.2014 (stored), C.A. Johns leg., 10.vii.2014, CJ307, PHIL0032 ♀, PHIL0033♂, in BPBM (only genitalia slides remain). Diagnosis. Forewing brownish fuscous to brown with narrow oblique white ds 1–3 . Similar to P. hibiscella (Swezey), but distinguished by the darker forewing with pale lemon-yellow apex. Redescription: Adult (Figs. 20A, B, 33G). Wingspan 9–11 mm in type series; forewing length 4.5 mm in lectotype, 3.5–4.6 mm in paralectotypes. Head dull ocherous; and frons ocherous; maxillary palpus ocherous, fuscous at apical portion; labial palpus white, terminal segment fuscous externally. Antenna fuscous, paler at the apex, 1.5x length of forewing. Thorax pale brownish ocherous. Forewing brownish fuscous to brown with narrow oblique white streaks: ds 1 at 1/4, ds 2 at 1/2, ds 3 at 3/4, about equally spaced, wider at base, tapering to apex, bent outwards; cs 3 at 2/3, meeting ds 3 in the middle of wing; apical portion pale lemon-yellow with a pale blue spot, sometimes dorsal half with fuscous patch from the dorsum to termen. Cilia white, apical cilia with four fuscous costal lines, terminal cilia fuscous, yellowish at base. Hindwing and cilia light fuscous. Abdomen grayish fuscous. Legs ocherous, fore and mid tibiae fuscous above, tarsi barred with fuscous. Male genitalia (Figs. 50E, 51C, 52C) (n=3). Capsule 1030 µm. Tegumen 0.8–0.9 x length of valva; valva 690 µm long, tapering along costal margin from 2/5 to apex, slightly rounded at apex and curvied toward dorsally (Fig. 50E). Saccus slender and needle-shaped in ventral view (Fig. 51C). Phallus 670 µm long and straight with developed coecum; cornuti in vesica indistinct (Fig. 52C). Female genitalia (Fig. 59E) (n=1). 1450 µm long. Ostium bursae large; antrum deeply cup-shaped with a pair of thick lateral lobes; lamella antevaginalis 240 µm, weakly sclerotized, trapezoid in ventral view, widening toward anterior margin of A7. Ductus bursae slender and long, middle part weakly sclerotized, round and flat; terminus of ductus bursae tubular, biforked. Corpus bursae 740 µm, oblong, signa a pair of longitudinal, partly sclerotized wrinkles. Distribution. Oahu (Swezey 1915). Host plants. Urticaceae: Urera sandvicensis Wedd. (Swezey 1915) and U. glabra Wedd.: new record. Urera is now rare on the island of Hawaii (Big Island), but U. glabra can still be found in large numbers on Oahu. Biology. (Fig. 95A–C) Swezey (1915: 96) reported the biology: “The mine is very slender where it starts from an egg placed on the under side of the leaf, it gradually widens as the larva grows, becomes serpentine and towards the last enlarges to a blotch. The larva emerges to spin its whitish cocoon on the surface of the leaf. The moths emerged from the cocoons in about ten days.” We observed larvae mining leaves of U. glabra beginning with a slender, tortuous linear mine that became a blotch mine (Fig. 95A, B) that gradually expanded as larvae grew (Fig. 95A). Usually 2–3 mines per leaf. The final instar larva spun a reddish-brown cocoon at the leaf margin, the leaf margin was slightly curled upwards from the contraction of the cocoon silk (Fig. 95C). Mines were rarely observed during our field surveys, but nearly all mines that we did see were active or fresh. Parasitoids. Eulophidae: Sierola sp., Bethylidae; Euderus metallicus (Ashmead, 1901) (Zimmerman 1978a). Remarks. We discount the distribution records from the island of Hawaii (Big Island) determined by Swezey that were mentioned in Zimmerman (1978a). We could not identify the specimens from Kilauea, Hawaii (Big Island) stored in BPBM, USNM and NHMUK. We also investigated two populations of Urera plants on the island of Hawaii (Big Island), but these plants did not have any Philodoria mines. The type series was reared by Swezey from the same lot of leaves from which the type series of “ P. ureraella ” (P. ureraella and P. opuhe in the present study) was reared. In the present study, P. urerana was obtained from leaves of Urera glabra on Oahu., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 87-88, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Swezey, O. H. (1915) New species of Hawaiian moths. P roceedings of the Hawaiian Entomological Society, 3 (2), 94 - 97. https: // doi. org / 10.5962 / bhl. part. 24594","Swezey, O. H. (1928) Some new species of lepidopterous leaf-miners in Hawaii. Proceedings of the Hawaiian Entomological Society, 7 (1), 187 - 191.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718."]}

Philodoria basalis Walsingham, 1907 Figs. 8C–F, 26C, 34D, 37A, 40E–H, 41I, 42H, 43H, 61H, 81. Philodoria basalis Walsingham, 1907: 720, pl. 5, fig. 24; Zimmerman 1978a: 695, figs. 468, 475. Type locality. Olinda (Haleakala), 4000 ft (Maui). Type material. Lectotype ♀ (=“Type ♂ (26666) Mus. Wlsm” in original description), Olinda, 4000ft, Haleakala, Maui, iv. 1894, Perkins 26666, B.M. ♀ Genitalia slide no. 3952 | PHILODORIA BASALIS, Wlsm. Fn. Hawaii. I. TYPE ♂ descr. figd.| Walsingham Collection. 1910—427.| NHMUK010305331 in NHMUK (here designated). Paralectotype ♂, same locality as holotype, v. 1896, Perkins 28541 | Philodoria basalis ♂ Wlsm PARATYPE 1/1 descr. figd.| Walsingham Collection. 1910—427.| NHMUK 010305332 in NHMUK. Described from two specimens: ‘Type ♂ (26666)’ and ‘paratype’ from Maui. This may indicate that Lord Walsingham considered them as holotype and paratype, as indicated on the specimen label. But because a holotype was not specified in the description, the so-labeled holotype and paratype are to be considered syntypes under Article 73.2 of the Code (ICZN 1999), and any one is thus eligible for designation as lectotype under Article 74 of the Code (ICZN 1999). The syntype ‘Type ♂ (26666)’ (= ♀) is here designated as lectotype (Fig. 8C), and the other syntype is the paralectotype. Additional material. 41 (17♂, 20♀, 4 sex unknown). Maui, in BPBM: 1♀, 2 (sex unknown), Keanae, 22.viii.1918, BPBM 34151; 1 (sex unknown), Makawao forest Reserve, 7.v.2013 (stored), C.A. Johns leg., host: Metrosideros polymorpha, 13.iv.2013, CJ112, KT982411; 2♂, 2♀, Makawao For. Res., Flume Rd. 1296 m, N20.80720 W 156.25259, 10–11.v.2013 (UV Bucket trap 1), A. Prestes & W. Haines leg., SK635–638. Hawaii (Big Island): 1♂, S. Kona, 8.viii.1919, O.H. Swezey Collector, “Ohia”, BPBM 34148; 1♀, Upper Hamakua Ditch Trail, Kohala Mts., 4.ix.1919, O.H. Swezey Collector, “Ohia Lehua”, Z-XII-20-62-4 (“Slight vari”), BPBM 34152; host: M. polymorpha in BPBM: 1♂, Kohala Watershed Partnership, 4.vi.2015 (stored), C.A. Johns leg., 18.v.2015, CJ416, SK775♂; 2♂, 1♀, Puu Makaala Natural Area Reserve, Hawaii, 30.v.& 13.vi.2015 (stored), C.A. Johns leg., 7.v.2015, CJ410, CJ430 / SK776♂, CJ431 / SK778♂; 4♂, 12♀, 1 (sex unknown), same locality, 3.v.–6.vi.2016 em., A.Y. Kawahara leg., 1.v.2016, SKH-09-1 / SK 615♀, SK 616♀, 617♀; 1♂, Volcanoes National Park, Thurston Lava Tube, 24.v.2016 em., C.L.-Vaamonde & C. Doorenweerd leg., 22.iv.2016, HILO019; 4♂, 2♀, Same locality, 16–28.v.2016 em., A.Y. Kawahara leg., 27.iv.2016 (larva); 3♂, 1♀, same locality, 28–29.iv.2016 (light trap). In NHMUK: 1♂, Hawaii 1300’, S. Hilo Distr., Waiakea F. R., Stainback Highway, 21.vi.1976, K. & E. Sattler, B.M.1976-605, Philodoria sp. 11 (Hawaii) Sattler Coll., D. C. Lees Sep. 2016, BMNH(E)1621300; 1♂, Hawaii 1300’, S. Hilo District, Upper Waiakea F. R., Stainback Highway, 9.vii.1976, K. & E. Sattler, B.M.1976-605, Philodoria sp. 11 (Hawaii) Sattler Coll., D. C. Lees Sep. 2016, BMNH(E)1621134; 1♂, Hawaii 2800’, Kau District, Ocean View, 20.vi.1976, K. & E. Sattler, B.M.1976-605, Philodoria sp. 10B cf splendida (Hawaii [=Big Island]) Sattler Coll., D. C. Lees Sep. 2016, BMNH(E)1621065; 1♂, Hawaii 6200’, Kau District, Kahuku Ranch, Punaluu Kahawai, 4.viii.1976, K. & E. Sattler, B.M.1976-605, Philodoria sp. 15 (Hawaii [=Big Island]) Sattler Coll., D. C. Lees Sep. 2016, BMNH(E)1621320; 1♂ 1♀, Kilauea, Hawaii (Big Island), at light (= light trap), 20& 21.v.1915, A. Busck Collector, deposited in USNM. Diagnosis. A leaden gray patch covers the entire lower (anal vein) portion of forewing (Fig. 8C–F). The male genitalia has a broad valva with a short dorsal process, a rather broad and shorter capsule, and a broad phallus. The female has a sclerotized lamella antevaginalis and oblong pod-shaped signa similar to those of P. splendida Walsingham and P. lama sp. n. Redescription: Adult (Fig. 8C–F, 34D). Wingspan 8–10 mm; forewing length 4.5 mm in holotype, 3.8–4.3 mm in the additional specimens examined. Head leaden gray; frons white; maxillary palpus reduced; labial palpus white (Fig. 34D, 37A). Antenna grayish fuscous. Thorax: grayish fuscous. Forewing leaden gray suffused with pale brown patches enclosing dark gray fascia; tf from costal 1/3 to dorsal 1/2, tp after the middle to apical portion, distinctly narrowing near the dorsum, containing two short, shining, steel gray costal spots; apical portion with large jet-black terminal patch from tornus to apex, containing two upright steel-gray spots near its opposite extremities; a single bright silvery dot in its middle; a very narrow ocherous line along the base of tornus; cilia shiny, leaden gray with a blackish fringe line; two white costal streaks, longer streak extends to apex; terminal cilia often black. Hindwing tawny black; cilia tawny gray. Abdomen grayish fuscous above, banded with white beneath. Legs pale graysih fuscous, spurs white. Male genitalia (Figs. 41I, 42H, 43H): new record (n=4). Capsule 750 µm. Tegumen 1.1–1.2 x length of valva; valva 500 µm long, rather broad and having a knob-shaped dorsal process; dorsal process densely covered with minute spines ventrally and on inner margin (Fig. 41I). Saccus rather short, broad, slightly narrowing in the middle (Fig. 42H). Phallus 650 µm long, rather broad; two series of cornuti large and developed in vesica (Fig. 43H). Female genitalia (Fig. 40E–H, 61H; Zimmerman 1978a: fig. 474) (n=3). 1450 µm long. Ostium bursae broad; antrum very short and wide; lamella antevaginalis 200 µm, semicircular (Fig. 40F, H). End of ductus bursae broad, connecting in dorsal portion of anterior part of corpus bursae; corpus bursae about 1000 µm long; anterior end of corpus bursae weakly sclerotized; some lines consisting of wrinkles running longitudinally, some sclerotized; paired signa oblong and podlike with a series of minute spines (Fig. 40G, H). Distribution. Maui (Walsingham, 1907) and Hawaii (Big island) (Zimmerman 1978a). Host plants. Myrtaceae: Metrosideros polymorpha Gaudich. (present study), Metrosideros sp. (Zimmerman 1978a). Biology. (Fig. 81). Zimmerman (1978a: 695) reported the cocoon and mature larval habit based on Swezey's observations. We observed larvae on multiple occasions. Early instar larvae form a slender, linear mine on the adaxial leaf surface that gradually expands into a larger blotch mine as the larva feeds and grows (Fig. 81A, G). Mines ~ 5 cm in length, brownish pale green; older mines turn dark brown to pale grayish ocherous and redish brown (Fig. 81B). One or two mines per leaf (Fig. 81E). The larva nearly always causes the leaf to dehisce from the plant, leading to the mined leaf to fall to the ground. Fully grown larvae exit the mine, and cut an oval ring into the upper (adaxial) epidermis of the bloch mine (Fig. 81J). The oval epidermis is folded by the final instar larva by using silk to pulling two ends of the outer surface of the epidermis, creating a “taco”-like cocoon (5.0–6.0 mm in length and 1.5–2.0 mm in width), which hardens and becomes bronze-colored over time (Fig. 81B, K–M). The taco often falls out of the oval cut-out in the leaf and falls beneath the leaf litter (Fig. 81C), but can occasionally remain attached (Fig. 81C, D, F). We suspect it may help avoid parasitoids and other predators while being in a humid environment. The penultimate and final instar larvae are approximately 4–5 mm long and pale yellow (Fig. 81H, I). During the day, adults rest on adaxial side of leaves of Melicope species, Rutaceae (not a Philodoria host plant) at Mt. Kaala on Oahu. Parasitoids. Eulophidae: Euderus metallicus (Ashmead, 1901) (Zimmerman 1978a). Remarks. We identified one adult moth (Coll ID CJ-112 / GenBank accession no. ID KT982411) as P. basalis, based on a leaden gray patch covering the entire lower portion of forewing, from which whole bodies were sacrificed for molecular analysis (Johns et al. 2016). William Haines (pers. comm.) collected> 10 adults of this species in UV Bucket trap at one night in Makawao Forest Reserve, Maui (10–11.v.2013)., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 36-38, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Walsingham, Thomas, de Grey (Lord) (1907) Microlepidoptera. In: Sharp, D. (Ed.), Fauna Hawaiiensis. 1 (5). Cambridge University Press, Cambridge, pp. 469 - 759.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","ICZN (1999) International code of zoological nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 306 pp.","Swezey, O. H. (1920) Some new Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 4 (2), 376 - 386. https: // doi. org / 10.5962 / bhl. part. 16149","Johns, C. A., Moore, M. R. & Kawahara, A. Y. (2016) Molecular phylogeny, revised higher classification, and implications for conservation of endangered Hawaiian leaf-mining moths (Lepidoptera: Gracillariidae: Philodoria). Pacific Science, 70 (3), 361 - 372. https: // doi. org / 10.2984 / 70.3.7"]}

Philodoria marginestrigata (Walsingham, 1907) Figs. 23A–C, 27P–R, 37C, 40A–D, 50J–M, 51H, 52H, 60A–C, 77E, F, 93C. Gracilaria marginestrigata Walsingham, 1907: 721–722, pl. 25, fig. 26; Swezey 1913b: 223. Parectopa marginiestrigata (Walsingham, 1907); Meyrick 1912b: 48. (mistake in the character) Parectopa marginestrigata (Walsingham, 1907); Swezey 1928: 190. Philodoria marginestrigata (Walsingham, 1907); Zimmerman 1978a: 681, 685, figs. 432, 443, 450, 455. Type locality. Waianae Mts. (Oahu). Type material. Lectotype ♀, Waianae Mts., 1400ft., OAHU, Hawaiian Is., iv.1892, Perkins. 25990 | Gracilaria marginestrigata Wlsm. ♀ PARATYPE 3/15| Walsingham Collection, 1910-427.| BMNH(E) 1407669 in NHMUK. (here designated). Paralectotype 5 (1♂, 4♀; BPBM ones are all from above Walsingham accession and ‘PARATYPE’ below is short for ‘ Gracilaria marginestrigata Wlsm. PARATYPE’ as printed on large black-margined labels, with the 5-digit Walsingham specimen numbers whose first digit is ‘2’ written on locality label): 1♂, Waianae Mts., 1400ft., OAHU, Hawaiian Is., iv.1892, Perkins. 25989| Gracilaria marginestrigata Wlsm. ♂ PARATYPE 2/15| Fauna Hawaiiensis Collection| BPBM 34197|; 1♀, Waianae Mts., 1400ft., OAHU, Hawaiian Is., iv.1892, Perkins. 25991| Gracilaria marginestrigata Wlsm. ♀ PARATYPE 4/15| Fauna Hawaiiensis Collection| BPBM 34195|; 1♀, Waianae Mts., 1400ft., OAHU, Hawaiian Is., iv.1892, Perkins. 25996| Gracilaria marginestrigata Wlsm. ♀ PARATYPE 8/15| Fauna Hawaiiensis Collection| BPBM 34198|; 1♀, Malaweli (sea level), KAUAI, Hawaiian Is., ex V. 1895 Perkins. 27969| Gracilaria marginestrigata Wlsm. ♀ PARATYPE 14/15| Fauna Hawaiiensis Collection| BPBM 34194|; 1♀, Mts. 3–4000 ft, KAUAI, Hawaiian Is., VI. 1895 Perkins. 27096| Gracilaria marginestrigata Wlsm. ♀ PARATYPE 12/15| Fauna Hawaiiensis Collection| BPBM 34196|. Described from 17 specimens: ‘type ♀ (25992); ♂ (25991)’ and 15 ‘paratypes’ from Kauai, Oahu, Molokai. Given the way the description was written and the labels on the specimens, Walsingham likely considered them as holotype, allotype and paratypes. However, because a holotype was not formally specified in the original description, the so-labeled types should all be considered syntypes under Article 73.2 of the Code (ICZN 1999), and any of them can be designated as lectotype under Article 74 of the Code (ICZN 1999). Zimmerman (1978a, 701, fig. 443) provided the last known photograph for the ‘type ♀ (25992)’, which he treated as holotype in the NHMUK. The present curator of the NHMUK, D.C. Lees informed the authors that syntypes, ‘type ♀ (25992); ♂ (25991)’ and 11 ‘paratypes’ are not stored where they should and are missing. The syntype (25990) is here designated as lectotype (Fig. 23A) and the remaining syntypes are paralectotypes. Type material in original description. ‘ Type ♀ (25992); ♂ (25991) Oahu, Mus. Wlsm. HAB. Makaweli, sealevel, Kauai, larva mining various plants (2) excl. v.1895. Makaweli, Mts, 3000–4000ft., Kauai, larva mining various plants (2) excl. vi.1894. Waianae Mts., 1400ft., Oahu, on the leaves of a berry-bearing plant called “Kikania” [Xanthium strumarium ?](11) iv.1892. Sea-level, Molokai, (1) v.1893; above 3000ft, (1) 5.vi.1893.’ (Walsingham 1907: 722). Additional material. 75 (23♂, 16♀, 36 sex unknown). Nihoa, in BPBM: 12 (sex unknown), Nihoa I., 13.vi.1923, E.H. Bryan Jr. Collector, [host:] Sida, 34199–34210; 2 (sex unknown), Nihoa Island, 10.vi.1962, J.W. Beardsley, ex Sida, BPBM 34122| Parectopa marginestrigata Walsingham det by J.W. Beardsley| 34123; 3 (sex unknown), Hawaiian Is, Nihoa I: Miller Val. above campsite, 25.vi.1990 at UV light, J. Suazanac coll., 36525–36527; 1 (sex unknown), Nihoa I, 15.vi.1980 at light, Aoc. No. 1980.301, S. Conant Coll., BPBM 36528. Oahu: 1♂, 2♀, 2 (sex unknown), Koko Hd. [Koko head?, Oahu], 22.ii.1927, O.H. Swezey Collector, ex Abutilon incanum, BPBM 34211– 34215; 2 (sex unknown), Kaimuki, Oahu, Coll. O.H.S., BPBM 34216, 34217; 2♂, 2♀, Kaimuki, Oahu, Coll. O.H.S., SK662♂, SK663♂, SK 664♀, SK 665♀, BPBM 34220; 5 (sex unknown), Kaimuki, Oahu, Coll. O.H.S., BPBM 34221; 3♂, 1 (sex unknown), Kaimuki, Oahu, Coll. O.H.S. ex Sida in USNM; 4♂, 2 (sex unknown), Kaimuki, Oahu, Coll. O.H.S., “♂ genitalia twice? on slide aB? Aug.22.1941” in USNM; 1 (sex unknown), Hawaiian Ids, Oahu, Honolulu, 9.iii.1917, J.C. Bridwell, BPBM 34219; 4♂, 5♀, 5 (sex unknown), Punahou, Oahu, xi.1963, M. Chong Collector, ex. Abutilon mlle in USNM.1♂, Mokio, Molokai, 30.xii.2013 (stored), C.A. Johns leg., host Lipochaeta rockii 16. xii.2013, CJ165, SK760♂ in BPBM. Host: Abutilon menziesii: 1♂, 1♀, Mokio, Molokai, 2& 4.i.2014 (stored), C.A. Johns leg., 16.xii.2013, CJ175b / SK 772♀, CJ187 / SK771♂ in BPBM. Host: Sida fallax: C.A. Johns leg., all specimens were spreserved in BPBM: 1♂, Kokee, Kauai, 21.ix.2015 (stored), CJ521, SK784; 1♀ 1 (sex unknown), Kaena Point, Oahu, 17& 21.v.2013 (stored), 19.iv.2013, CJ135(sex unknown) / KT982412, CJ 138♀; 1♂, Same locality, 12.vii.2014 (stored), 1.vii.2014, CJ278b / SK785♂; 1♂ 1♀, Mokio, Molokai, 30.xii.2013 (stored), 16.xii.2013, CJ168 / SK782♂, CJ173 / SK 783♀; 2♂, 2♀, Kanepuu, Lanai, 16.viii.2016 (stored), 5.viii.2014, CJ379a / SK786♂, CJ379b / SK 787♀, CJ379c♂, CJ379d ♀. Diagnosis. The forewing pattern is similar to that of P. lipochaetaella (Swezey) and P. sciallactis (Meyrick), but distinguished by the narrowing ds 1–3 and cs 1–3 . Genitalia are similar to that of P. sciallactis, but P. marginestrigata has a rather short and slender saccus. Redescription: Adult (Fig. 23A–C, 37C). Wingspan 6.5 mm in ‘Type ♀ (25992)’, 6–7 mm for paralectotype; forewing length 2.4 mm in lectotype, 2.8 mm in paralectotype, 2.5–3.0 mm for the additional specimens examined. Head ocherous; frons white; maxillary palpus white with fuscous scales at the paex; labial palpus white, a dark spot at the end of the median joint and another before the apex of the terminal joint beneath (Fig. 37C). Antenna ocherous, faintly annulate with white. Thorax ocherous. Forewing ocherous, with six oblique white streaks, narrowing black-margined; three costal streaks (cs 1–3 ): cs 1 and cs 2 small, nearly meeting the extremities of the larger ds 1 and ds 2 below them, which arise in each instance nearer to the base; cs 3 at costal 3/4, slender, extending beyond the apex of the triangular, somewhat curved, ds 3 below it; apical portion with shiny, steel-gray spot (as) with a patch of black scales; an elongate patch shining blue-black scales from ds 3 to the apex; cilia shiny, blue-gray, with three short costal spots (a, b, c) and a slender bl 1 of black scales, bronze gray at the tornus. Hindwing and cilia bronze gray. Abdomen leaden gray. Legs gray with white spots at the extremities of the tibiae and tarsi. Male genitalia (Figs. 40A–C, 50J–M, 51H, 52H) (n=7). Capsule 600 µm. Tegumen 0.8–0.9 x length of valva; valva 440–460 µm long, basal portion broad along dorsal margin, tapering along costal margin from basal 2/5–1/2 to apex; apical half slender and slightly curved toward inner margin with small spines at apex; a set of short spines arranged along inner side of basal region (Figs. 40C, 50J–M). Saccus short and triangular in ventral view (Figs. 40B, 51H). Phallus 460 µm long and nearly straight, slightly widening at phallobase with small coecum; cornuti in vesica indistinct (Figs. 40A, 52H). Female genitalia (Fig. 60A–C) (n=4). 930 µm long. Ostium bursae large; antrum low cup-shaped with a pair of lateral lobes; lamella antevaginalis 110–160 µm, weakly sclerotized, sometimes slightly inflexed on the posterior margin, widening toward anterior margin of A7 (Fig. 60B, C). Ductus bursae short and broad, middle region weakly sclerotized, round and flat; terminus of ductus bursae tubular, biforked; middle region of ductus seminalis broad. Corpus bursae 570 µm, oblong with a very small spiny signa; anterior end of corpus bursae weakly sclerotized; paired rows of longitudinal, partly sclerotized wrinkles. Distribution. Nihoa, Kauai, Oahu, Molokai, Hawaii (Big Island) (Zimmerman 1978a) and Lanai: new record. Host plants. Asteraceae: Dubautia sp. and Xanthium strumarium (Swezey 1928: 190) (= X. echinatum (Zimmerman 1978a)), Xanthium sp. (Swezey 1913a: 223), and Lipochaeta rockii Sherff is new record; Malvaceae: Abutilon grandifolium (Willd.) Sweet [= Abutilon molle (Zimmerman 1978a: 681)], A. incanum (Link) Sweet (Swezey 1928: 190), Abutilon sp. (Swezey 1913a: 223), Abutilon menziesii Seem. (Endangered, IUCN) is new record, Sida cordifolia L., S. fallax Walp., S. meyeniana Walp., S. rhombifolia L. (Swezey 1928:190), Sida sp. (Swezey 1913a: 223). Biology. (Fig. 93C). Swezey (1913b: 223) and Zimmerman (1978a: 685) reported its biology: “this species is a common leaf-miner in the lowlands and there may be as many as 12 mines in a single leaf, and the cocoon is made within the mine; the larvae feed introduced plants, as unusual for an endemic insect (Zimmerman 1978a: 685)”. We collected larvae from three genera in two plant familes, Asteraceae (Lipochaeta) and Malvaceae (Abutilon and Sida). Larvae that mine leaves of Sida fallax formed a tortuous linear mine; 1–4 mines per leaf (Fig. 93C). Parasitoids. (Figs. 108, 109). Braconidae: Mirax sp.; Eulophidae: Cirrospilus sp., Diglyphus begini (Ashmead, 1904), Euderus metallicus (Ashmead, 1901), Neochrysocharis formosus (Westwood, 1833), Pnigalio externa (Timberlake, 1927), Sympiesis vagans (Timberlake, 1926), Zagrammosoma flavolineatum (Crawford, 1913); Pteromalidae: Lyrcus tortricidis (Crawford, 1921) (Zimmerman 1978a). Remarks. Walsingham (1907) presented an inaccurate and misleading figure of this moth (pl. 25, fig. 26), which does not show the distinct first white costal streak, which looks like a dorsal patch instead (after Zimmerman 1978a). Zimmerman (1978a) also noted that observations by Perkins and Walsingham are mixed and there may be more than one species involved in the literature records. We treat the first Nihoa record reported by Bryan (1926) from the 1923 Tanager Expedition as P. marginestrigata, but that specimen has three costal and three dorsal white streaks. There is some phenotypic variation in wing pattern and male genitalia (particularly in the shape of the valva (Figs. 27P–R, 50J–M)) between populations on different islands and those feeding on different host plants. DNA barcoding may help with the delimitation of this species., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 101-103, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Walsingham, Thomas, de Grey (Lord) (1907) Microlepidoptera. In: Sharp, D. (Ed.), Fauna Hawaiiensis. 1 (5). Cambridge University Press, Cambridge, pp. 469 - 759.","Swezey, O. H. (1913 b) Leaf-miners of the Hawaiian Islands. Proceedings of the Hawaiian Entomological Society, 2 (5), 221 - 227.","Meyrick, E. (1912 b) Adelidae, Micropterygidae, Gracilariadae. In: Wagner, H. (Ed.), Lepidopterorum Catalogus. Vol. 6. W. Junk, Berlin, pp. 48. https: // doi. org / 10.5962 / bhl. title. 122538","Swezey, O. H. (1928) Some new species of lepidopterous leaf-miners in Hawaii. Proceedings of the Hawaiian Entomological Society, 7 (1), 187 - 191.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","ICZN (1999) International code of zoological nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 306 pp.","Swezey, O. H. (1913 a) A day's collecting at Punaluu, Oahu. Proceedings of the Hawaiian Entomological Society, 2 (5), 197 - 199.","Bryan, E. H. (1926) Insects of Hawaii, Johnston Island and Wake Island. Introduction. Bernice P. Bishop Museum Bulletin, 31, 1 - 16."]}

Philodoria wilkesiella Swezey, 1940 Figs. 12D, 26N, 47A, 48A, 49A, 58A, 88A. Philodoria wilkesiella Swezey, 1940: 464–465; Zimmerman 1978a: 718, figs. 467, 473, 480. Type locality. Puu Kukui (Maui). Type material. Lectotype ♂, Puu Kukui, Maui, 20.xii.1928, O.H. Swezey Collector, Wilkesia grayana (= Argyroxiphium grayanum), | Holotype Philodoria wilkesiella Sweszey |, Type no. 4274 in BPBM. Paralectotype missing. Described from nine specimens reared from mines in leaves of “ Wilkesia grayana ”, Puu Kukui, Maui, December 20, 1928. Given the manner in which the description was written, Swezey considered these specimens as holotype and paratypes, as indicated on their specimen label. However, because a holotype was not formally specified in the description, the so-labeled holotype is to be considered syntype under Article 73.2 of the Code(ICZN 1999), and either is eligible for designation as the lectotype under Article 74 of the Code (ICZN 1999). We formally designate the specimen bearing the label ‘Holotype’ as the lectotype (Fig. 12D) and the remaining syntypes are paralectotypes. Zimmerman (1978a) illustrated the genital character with photographs: paratype male, slide no. Z-XII-19-62-18 8, fig. 473; paratype female, Z-XII-19-62-19, ostial plate (corpus bursae missing), fig. 480. We were unable to find the paratype specimens and genital slides in BPBM. Additional material. 5 (3♂, 2♀) All specimens preserved in BPBM. 3♂, 2♀, Haelaau, Maui, 10.iv.2013 (stored), C.A. Johns leg., adult on Argyroxiphium grayanum, CJ 007 / SK647♂, CJ009 / SK648♂, CJ010 / SK649, CJ012 / SK650, CJ013 / PHIL0007♂. Diagnosis. Brownish fuscous ground color with a number of white spots on the forewing (Fig. 12D). Philodoria pipturiana Swezey is similar in color pattern of its forewing, but distinguished from P. wilkesiella by its dark fuscous ground color and a white longitudinal basal streak (Fig. 12A–C). Genital characters of P. wilkesiella resemble Asteraceae-feeding Philodoria species, e.g., P. epibathra Walsingham and P. naenaeiella (Swezey). However, it differs in having a valva with a broad apical half and a shorter male saccus. The female has a semicircular shaped lamella antevaginalis and shorter ductus bursae. Redescription: Adult (Fig. 12D, 26N). Wingspan 7 mm in type series; forewing length 3.25 mm in lectotype. Head white; frons white with some brownish scales; maxillary palpus reduced; labial palpus white with a few brownish scales externally. Antenna grayish fuscous, 1.2–1.3x length of forewing. Thorax white with brownish scales at the sides, white beneath. Forewing brownish fuscous with a number of white spots: three dorsal ones, the smaller one near base, two outwardly oblique larger spots at 1/3 and 3/5; a narrow elongate spot at about middle of the costa, two or three white costal spots towards apex, a small transverse white spot at apex and sometimes two or three small spots preceding this in apical area of wing. Cilia gray, dark at base, with a few white scales in vicinity of white spots. Hindwing uniform gray. Abdomen gray. Legs gray, with tarsi banded with white. Male genitalia (Figs. 47A, 48A, 49A) (n=3). Capsule 620 µm. Tegumen 0.7 x length of valva; valva tapering along costal margin from base 2/5 to apex; along the inner side of valva, a series of small spines arranged; apical half of valva straight in lateral view (Fig. 47A). Saccus short, stick-shaped (Fig. 48A). Phallus 570 µm long, slender with developed coecum (Fig. 49A). Female genitalia (Fig. 58A) (n=3). 890–900 µm long. Ostium bursae large; antrum cup-shaped with wavy pair of lateral lobes; lamella antevaginalis 160–170 µm, weakly sclerotized, semicircular in ventral view, widening toward anterior margin of A7. Ductus bursae thick, 270–300 µm. Corpus bursae small 560 µm; paired rows of wrinkles, some sclerotized. Distribution. Maui (Swezey 1940). Host plants. Asteraceae: Argyroxiphium grayanum (Hillebr.) O.Deg. (Swezey 1940). The host plant is located on the windy and rainy bogs of the summit of West Maui. Biology. (Fig. 88A). Similar to Dubautia miners, in that its first and second instar larvae mine along the length of the leaf, following leaf veins. Pupation occurs in the cocoon inside of the mine terminus, usually with one or both extremities of the white cocoon exposed (Fig. 88A). Three mines per leaf. Adults were observed on the host plant surface during sunny and less windy periods, and then they retreated to the substrate beneath the plant once these conditions subsided. Remarks. Zimmerman (1978a: 718) mentioned that “one of the few moths known to attack the extraordinary endemic composite Argroxiphium ”. We surveyed other members of Argyroxiphium with glabrous leaves, investigated type locality of this moth, and a population of A. grayanum on east Maui, but leaf mines were not found. We also checked some herbarium plant specimens of this east Maui population, and specimens of the extinct species, A. virescens. No leaf mines were found on these herbarium specimens., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 59-60, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Swezey, O. H. (1940) New species of Hawaiian Lepidoptera. Proceeding Hawaiian Entomological Society, 10 (3), 461 - 465.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","ICZN (1999) International code of zoological nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 306 pp."]}

Philodoria neraudicola (Swezey, 1920) Figs. 19A, B, 27I, J, 50A, B, 51A, 52A, 59A, B, 85E, 86C. Gracilaria neraudicola Swezey, 1920 b: 385–386. Parectopa neraudicola (Swezey, 1920); Swezy 1928: 191. Philodoria neraudicola (Swezey, 1920); Zimmerman 1978a: 689, fig. 444. Type locality. Punaluu (Oahu). Type material. Lectotype ♂, Punaluu, Oahu, 11.vi.1916, Coll. O.H.S. (= O.H. Swezey), ex Neraudia [melastomifalia], | Gracilaria neraudicola Swezey | Holotype | Type no. 4261 in BPBM (here designated). Paralectotype 5 (1♂, 1♀, 3 sex unknown): Host N. melastomifalia, | Paratype | O.H. Swezey: 1 (sex unknown), same data and locality as lectotype, missing abdomen, BPBM 34243; 1♂, 1♀, 2 (sex unknown), Waiahole, Oahu, 13.viii.1916, SK 708♀ | BPBM 34244, BPBM 34245, BPBM 34246, and 1♂, SK707♂ in USNM. Described from six specimens from Oahu. The holotype was not specified in the original description, therefore the so-labeled ‘holotype’ and ‘paratypes’ can be considered syntypes under Article 73.2 of the Code (ICZN 1999), and any of them is eligible for designation as lectotype under Article 74 of the Code (ICZN 1999). The syntype ‘holotype’ is here designated as lectotype (Fig. 19A) and the remaining syntypes are paralectotypes. Additional material. 3 (sex unknown), Kamiloloa, Mol., 3200, 20.xii.1925, O.H. Swezey, Pipturus, BPBM 34247, BPBM 34249, BPBM 34250; 4♀, Iao Valley, Maui, 9& 10.viii.2014 (stored), C.A. Johns leg., host: Pipturus albidus, 28.vii.2014, CJ336, SK 838♀ in BPBM; 1♀ 1 (sex unknown), Same locality, 10& 12.viii.2014 (stored), C.A. Johns leg., host: Pipturus sp., CJ360, CJ 369 in BPBM. Diagnosis. The forewing pattern is similar to P. hesperomanniella sp. n. and P. urerana (Swezey), but P. neraudicola has an orangish ocherous apical region; the saccus is slender and pointed. Swezey (1920) noted that a series of five specimens reared from leaves of Pipturus albidus at Pahoa, Puna on the island of Hawaii (Big Island), September 20, 1918 seems to be the same species. However, the five syntypes from Swezey (1920) have distinguishable forewing pattern from that of P. neraudicola, and therefore we treat them here as new species, P. obamaorum sp. n. (See diagnosis of P. obamaorum). Redescription: Adult (Fig. 19A, B). Wingspan 6.5–8.1 mm in type series; forewing length 3.75 mm in lectotype, 2.9–3.8 mm in paralectotype. Head creamy white; frons white; maxillary palpus white mixed with fuscous scales; labial palpus white, median joint fuscous at apex, terminal joint with fuscous band at middle and at apex. Antenna brownish, basal segment pale. Thorax brownish fuscous, pale in middle and posteriorly. Forewing brownish fuscous, with three outwardly-oblique white dorsal streaks (ds 1–3 ) widened at base and margined with a few black scales, somewhat curved apically, the ds 3 nearly connected with a white slender outwardly-oblique blackmargined line (cs 1 ) at three-fourths of costa, beyond this white line, three or four white costal spots (a, b, c); a round black spot at apex with a few pale blue scales, a few pale blue scales between this spot and the ds 3 , sometimes this area ocherous; apical and terminal cilia brown, paler near base and black bl 1 at extreme base, tornal cilia- very pale brown. Hindwing and cilia pale brown. Abdomen pale brownish. Legs ocherous, anterior and middle tibiae and tarsi and posterior tarsi fuscous marked. Male genitalia (Figs. 50A, B, 51A, 52A) (n=1). Capsule 780 µm. Tegumen 0.9 x length of valva; valva 540 µm long, tapering along costal margin from 2/5 to apex, slightly narrowing at middle with small spines at apex; a set of short spines arranged along inner side of basal region (Fig. 50A, B). Saccus slender and pointed at apex in ventral view (Fig. 51A). Phallus 570 µm long and thickened at phallobase with well-developed coecum; two series of minute cornuti in vesica (Fig. 52A). Female genitalia (Fig. 59A, B) (n=2). 1100–1200 µm long. Ostium bursae large; antrum cup-shaped with a pair of lateral lobes; lamella antevaginalis 220 µm, weakly sclerotized, trapezoid in ventral view, widening toward anterior margin of A7. Ductus bursae slender, middle region weakly sclerotized, round and flat; terminus of ductus bursae tubular, biforked. Corpus bursae 670 µm, oblong; signa a pair of longitudinal, partly sclerotized wrinkles. Distribution. Molokai (Zimmerman 1978a), Oahu (Swezey 1920), Maui: new record and possibly Kauai. Host plants. Urticaceae: Neraudia melastomifolia Gaudich. (Swezey 1920), Pipturus albidus A.Gray ex H.Mann and Pipturus sp. Biology. (Figs. 85E, 86C) Little is known about its biology. The larva on Pipturus form a blotch mine on the adaxial leaf surface that gathers the frass in the center (Fig. 85E); numerous mines and cocoons per leaf; pupation occurs in a brownish-white cocoon outside of the mine, which is usually on the adaxial leaf surface (Fig. 86C). Parasitoids. Eulophidae: Euderus metallicus (Ashmead, 1901), Pnigalio externa (Timberlake, 1927); (Zimmerman 1978a). Remarks. We could not locate the Kauai specimens and therefore the record from Kauai cannot be confirmed. Leaf mines on Pipturus in Iao Valley along Iao Stream were abundant during our survey in 2014. We also collected similar blotch mines (Fig. 86B) and tortuous linear to blotch mines (Fig. 85C, D) on West Maui and Hawaii (Big Island). Unfortunately, adults did not emerge, but these mines may be from larvae of P. floscula or P. haelaauensis sp. n. The host plant genus Neraudia has become exceedingly rare on the Hawaiian Islands with several species being listed as endangered. We were able to locate fewer than ten plants of N. melastomifolia throughout our surveys. None of these plants showed signs of leaf mine activity. However, surveys of Neraudia herbarium specimens (BPBM) revealed Philodoria leaf mines on several plants., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 85-86, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Swezey, O. H. (1920) Some new Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 4 (2), 376 - 386. https: // doi. org / 10.5962 / bhl. part. 16149","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","ICZN (1999) International code of zoological nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 306 pp."]}

Genus PHILODORIA Walsingham, 1907 Hawaiian Name: Hunelele ‘elilau Figs. 3–95, 97. Philodoria Walsingham, 1907: 717. Zimmerman 1978a: 644–718, figs. 427–481; Johns et al. 2016: 66–67. Synonymy Subgenus Eophilodoria Zimmerman, 1978: 659, fig. 432. Type species: Philodoria succedanea Walsingham, 1907. Subgenus Philodoria Zimmerman, 1978: 659, figs. 433–434. Type species: Philodoria marginiestrigata (Walsingham, 1907). Original description: “Antennae a little longer than the forewings, simple, without pecten. Labial Palpi long, curved, drooping, divergent, smooth; terminal joint almost as long as the median. Maxillary Palpi obsolete. Haustellum well-developed. Ocelli present. Head and thorax smooth. Forewings narrow, elongate, lanceolate: neuration 12 veins all separate; 2, 5 and 6 weak; 7 to costa. Hindwings narrow, lanceolate, acute; cilia 3 1/2: neuration 7 veins (3 and 4 coincident); 2 and (3 +4) stalked; cell open between 4 and 5; 5 and 6 stalked out of 7; 8 short. Abdomen slender. Legs, hind tibiae smooth. This genus is closely allied to Gracilaria, Hw., but differs in the absence of the maxillary palpi.” (Walsingham 1907: 717). Type species. Philodoria succedanea Walsingham, 1907 by original description. Systematic history. The systematic history of the genus was reviewed by Johns et al. (2016). In summary, the genus was placed originally in the Tineidae by Walsingham (1907), and incorrectly assigned to the Glyphipterygidae by Meyrick (1912a). Walsingham characterized the genus from others as lacking a maxillary palpus. Genera he thought were closely related, Gracillaria and Elachista, have a maxillary palpus and/or blackish forewing. Swezey (1910b, 1913bc, 1915, 1920, 1928, 1940 & 1946) and Meyrick (1928) similarly assigned species within Philodoria with a developed maxillary palpus to other genera, i.e. Parectopa and Gracillaria. Meyrick (1928) placed some Philodoria species described by Walsingham as Gracillaria in the Gracillariidae. Zimmerman (1978a) taxonomically reviewed Hawaiian Gracillariidae and placed some species assigned to other genera in Philodoria. Zimmerman also divided the genus into two subgenera, Philodoria (Eophilodoria) and Philodoria (Philodoria) based on the length of the maxillary palpus. Recently, Johns et al. (2016) synonymized the subgenus Eophilodoria Zimmerman 1978 with the genus Philodoria Walsingham, 1907. Philodoria has been treated as a genus of the Parectopa group, subfamily Gracillariinae (Kumata et al. 1988; Davis & Robinson 1998; De Prins & De Prins 2005). Recently, Kawahara et al. (2017) placed Philodoria in the subfamily Ornixolinae Kuznetzov & Baryshnikova, 2001 based on a phylogeny that utilized 22 genes and 94 gracillariid species. Johns et al. (2018) published the first molecular phylogeny of Philodoria utilizing 507 loci and 33 Philodoria species. They estimated the origin of the genus to the now partially sunken Hawaiian islands of Laysan or Lisianski, approximately 21 Ma, and hypothesized that the ancestral moth was likely associated with host plants in the families Ebenaceae, Malvaceae or Primulaceae. While the study of Johns et al. (2018) was a major breakthrough in understanding the evolutionary patterns of Philodoria, many of the species included in that study did not have formally assigned names and species boundaries were never formally examined. Two species of Philodoria were described by Kobayashi et al. (2018) but many more remained undecribed until the present revision. Diagnosis. Adults. The association of this genus to Ornixolinae is substantiated by the presence of three autapomorphies for the subfamily: 1) the presence of moderately long, four-segmented maxillary palpi; 2) forewing with three branches of the medial vein and two branches of the cubital vein (Fig. 38); and 3) an ostium bursae opening at the seventh sternum in female (Fig. 40E, F). However, 17 Philodoria species possess very reduced four-segmented maxillary palpi, of which the apical two segments are very small and indistinct (Figs. 34A,D, 37A, B) or nearly absent (Figs. 34B, C, 35). Diagnostic features of the genus include: forewing with lustrous and metallic apical portion; hindwing with small frenular costal bristles; in the male, a dorsal flap extending tergum VIII; in the female, terminus of ductus bursae tubular and sclerotized; corpus bursae with a pair of lateral signa that are either a pair of spines or series of small spines; inception of ductus seminalis at the posterior part of corpus bursae. Larvae and pupae were not examined in the present study and therefore do not include a diagnosis for them.. Redescription: Adult (Figs. 3–6, 34–40). Head and frons smooth. Haustellum well-developed. Ocelli and chaetocemata absent (despite Walsingham’s [1907] claim that ocelli are present, Philodoria lack ocelli). Maxillary palpus developed, 4-segmented (Figs. 34E, F, 36) or reduced 2-segmented and obscure small 3rd and 4th segments (Fig. 34A, D), sometimes greatly reduced, vestigial, obsolete (Figs. 34B, C, 35). Labial palpus developed, upcurved, 3-segmented. Antenna about 1.0–1.5 length of forewing. Thorax smooth. Legs slender, smooth. Basal color of forewing dark lustrous and metallic, fuscous or brown. Forewing pattern (Fig. 3) consisting of orange ocherous patches with white bands or spots (Fig. 3A, B) or oblique white to brown streaks (cs 1–3 and ds 1–3 ) (Fig. 3C, D). Hindwing a little shorter in length than forewing. Wing venation (Figs. 38, 62H). Forewing lanceolate, with thirteen veins (Fig. 38A). Sc along costal fold, ending at one-third of costa; R1 from base to ending at the middle of costa; branched R2 at two-third of the wing; R3, R4 and R5 from end of cell to costa; M1, M2, M3 and CuA1 from end of cell to tornus; CuA2 rather week, at about two-thirds of the wing; CuP weak, from base and reaching dorsal area; A1+2 from base to beyond midde of dorsum. Hindwing lanceolate, with eight veins (Fig. 38B); Sc along anterior margin, ending at one-fourth of costa with about four small frenulums; R1 weak, along anterior margin, ending at near the middle of the wing; RS from base to reaching apex; branched M1 and M2; CuA from base to ending at the middle of dorsum; branched M3; CuP from base to ending at one-fourth of dorsum. Abdomen. Tergum smooth without spiniform setae. Male 8th abdominal segment (Fig. 39) with one or two pairs of coremata; a dorsal flap extending tergum VIII, elongate, triangular, covered with slender, round and flat scales (Fig. 39, 40D). Male genitalia (Fig. 40A–C) Uncus absent. Tegumen as long as valva in length (Fig. 40B). Tuba analis membranous with weakly sclerotized subscaphium; gnathos V-shaped transverse band, terminal margin weakly joining subscaphium and anterior process connecting ventral base of tegumen (Fig. 40B). Valva elongate, at the middle, tapering along costal margin (Fig. 40C), some species have broad valva with a process; small spines at the apex, surrounded by a set of long setae; a set of short median sized spines arranged on the basal region. Vinculum U-shaped; saccus developed. Phallus tubular, slender; vesica with cornuti; coecum developed (Fig. 40A). Female genitalia (Fig. 40E–H) Apophyses anteriores and apophyses posteriores slender. Ostium bursae broad, opening at the middle of 7th abdominal segment (Fig. 40E); antrum cup-shaped with a pair of lateral lobes; lamella antevaginalis semicircular and sometimes sclerotized; ductus bursae slender, tubular, posterior section very slender and membranous, curved inside of body and median section weakly sclerotized and flattened (Fig. 40E, F); anterior section tubular sclerotized, terminus of ductus bursae biforked in ventral view and flanked pairs of sclerotized wrinkles reaching anterior end of corpus bursae; inception of ductus seminalis on the posterior part of ductus bursae; corpus bursae pyriform; signa both lateral, a pair of spines or many series of small spines (Fig. 40G, H). Distribution. United States: Hawaiian Islands (Nihoa, Kauai, Oahu, Molokai, Lanai, Maui and Hawaii [Big Island]). Host plants. Asteraceae, Ebanaceae, Malvaceae, Myrtaceae, Primulaceae and Urticaceae. Biology. (Figs. 79–95). Philodoria larvae mine leaves of Hawaiian endemic plants in at least 20 genera and 52 species. Of these, 16 plant species are herbaceous. The larva forms a slender linear or serpentine mine on the adaxial side of the host leaf, and gradually expands its mine as it feeds and the mine later becomes the form of a blotch. Larvae of nearly all Philodoria species are presumed specialists on one plant genus (for example, 16 Philodoria species were recorded from one plant species), except for P. neraudicola P. sciallactis and P. splendida, that utilize two genera in one plant family. Philodoria marginestrigata is unusual in that it utilizes five genera in two plant families. Twenty-seven species are known to create a cocoon outside of the mine, usually pupating on the surface of the leaf. Three species, P. dubautiella , P. marginestrigata, and P. wilkesiella pupate within larval mines (Fig. 88A; Swezey 1913: 279; Zimmerman 1978a: 685). Natural enemies. (Table 3, Figs. 98–110). Natural enemies, other than parasitoids, are not well known. Eighteen parasitoid species from four families have been recorded from 14 Philodoria species (Yoshimoto 1965; Zimmerman 1978a), these are: Bethylidae: Sierola planiceps, S. philodoriae, S. pulchra, Sierola sp.; Braconidae: Mirax sp., Pholetesor bedelliae; Eulophidae: Cirrospilus sp., Closterocerus sp., Diglyphus begini, Diglyphus sp., Euderus metallicus, Neochrysocharis formosus, Pauahiana maculatipennis, P. metallica, Pnigalio externa, Sympiesis vagans, Zagrammosoma flavolineatum; Pteromalidae: Lyrcus tortricidis. We reared and examined 98 parasitoid wasps that emerged from 15 Philodoria species. Remarks. A total of 51 species are described or redescribed in the present study (See Table 4 and discussion). In addition, some potentially unnamed species have been reported, for example, a “ Philodoria (Philodoria) species” on Maui associated with Clermontia (Campanulaceae) (Zimmerman 1978a), and two unidentified “ Philodoria (Eophilodoria) species” on the island of Hawaii (Big Island) associated with Myoporum (Scrophulariaceae) (Mines by Swezey 1954:136) and Pisonia (Nyctaginaceae) (See Swezey 1954:167) respectively. We observed small minelike markings on Pisonia on Maui and Oahu, and window-like leaf mines in Clermontia on East and West Maui, but these mines contained no larvae and it is unclear whether these are made by Philodoria. We also could not observe Philodoria leaf mines on Myoporum, despite there being records of Philodoria on this plant. Myoporum leaves from Maui and Hawaii (Big Island) had leaf mines, but those were made by agromyzid flies. Etymology: According to Walsingham (1907), the scientific name “ Philodoria ” has the meaning “bounteousness.” It remains unknown what aspect of the moths this word refers to. We present a Hawaiian common name because there are many Hawaiian names for endemic plants and animals, yet this group of moths lacks one. Early Western entomologists in Hawaii did not record Hawaiian names of the many living plants and animals nor did they often take advantage of native guides in their entomological explorations. Roughly translated, hunelele means "tiny flier" (hune + lele) which reflects the microlepidopteran nature of these moths. The second part ʻelilau means "leaf excavator" and alludes to the leaf mining habits of the larval form. We encourage anyone who works with Philodoria species to use this name when referring to the genus in the vernacular., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 10-16, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Walsingham, Thomas, de Grey (Lord) (1907) Microlepidoptera. In: Sharp, D. (Ed.), Fauna Hawaiiensis. 1 (5). Cambridge University Press, Cambridge, pp. 469 - 759.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","Johns, C. A., Moore, M. R. & Kawahara, A. Y. (2016) Molecular phylogeny, revised higher classification, and implications for conservation of endangered Hawaiian leaf-mining moths (Lepidoptera: Gracillariidae: Philodoria). Pacific Science, 70 (3), 361 - 372. https: // doi. org / 10.2984 / 70.3.7","Meyrick, E. (1912 a) Lepidoptera, Heterocera, Fam. Gracilariadae. In: Wystman, P. (Ed.), Genera insectorum. Vol. 128. L. Desmet-Verteneuil, Bruxelles, pp. 36.","Swezey, O. H. (1910 b) Some new species of Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 2 (3), 106.","Meyrick, E. (1928) Some new species of Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 7 (1), 104.","Kumata, T., Kuroko, H. & Ermolaev, V. P. (1988) Japanese species of the Acrocercops - group (Lepidoptera: Gracillariidae). Part I. Insecta Matsumurana, New Series 38, 1 - 111.","Davis, D. R. & Robinson, G. S. (1998) The Tineoidea and Gracillarioidea. In: Kristensen, N. P. (Ed.), Lepidoptera, Moths and Butterflies. 1. Evolution, Systematics, and Biogeography. Handbook of Zoology. Vol. 4. Walter de Gruyter, Berlin, pp. 91 - 117. https: // doi. org / 10.1515 / 9783110804744.91","De Prins, W. & De Prins, J. (2005) Gracillariidae (Lepidoptera). In: Landry, B. (Ed.), World Catalogue of Insects. Vol. 6. Apollo Books, Stenstrup, pp. 1 - 502.","Kawahara, A. Y., Plotkin, D., Ohshima, I., Lopez-Vaamonde, C., Houlihan, P., Breinholt, J. W., Kawakita, A., Xiao, L., Regier, J. C., Davis, D. R., Kumata, T., Sohn, J. - C., De Prins, J. & Mitter, C. (2017) A molecular phylogeny and revised higher-level classification for the leaf-mining moth family Gracillariidae and its implications for larval host-use evolution. Systematic Entomology, 42, 60 - 81. https: // doi. org / 10.1111 / syen. 12210","Johns, C. A., Toussaint, E. F. A., Breinholt, J. W. & Kawahara, A. Y. (2018) Origin and macroevolution of micro-moths on sunken Hawaiian Islands. Proceedings of the Royal Society B, 285, 20181047. https: // doi. org / 10.1098 / rspb. 2018.1047","Kobayashi, S., Johns, C. A., Lopez-Vaamonde, C., Doorenweerd, C., Kawakita, A., Ohshima, I., Lees, D. C., Hanabergh, S. & Kawahara, A. Y. (2018) Hawaiian Philodoria (Lepidoptera, Gracillariidae, Ornixolinae) leaf mining moths on Myrsine (Primulaceae): two new species and biological data. ZooKeys, 773, 109 - 141. https: // doi. org / 10.3897 / zookeys. 773.21690","Yoshimoto, C. M. (1965) Synopsis of Hawaiian Eulophidae including Aphelininae (Hym.: Chalcidoidea). Pacific Insects, 7 (4), 665 - 699.","Swezey, O. H. (1954) Forest entomology in Hawaii. An annotated check-list of the insect faunas of the various components of the Hawaiian forests. Bishop Museum Special Publication 44. University of Hawaii Press and Bishop Museum Press, Honolulu, 266 pp."]}

Philodoria lysimachiella Swezey, 1928 Figs. 8A, 26H, 37B, 38, 41E, 42E, 43E, 56G, 62A–E. Philodoria lysimachiella Swezey, 1928: 188; Zimmerman 1978a: 708, figs. 463, 469. Type locality. Puu Hapapa, Waianae Mountains (Oahu). Type material. Lectotype ♂, Windward slope of Puu Hapapa, 2000 ft, half a mile south from Kolekole Pass, Waianae Mts., Oahu, 16.i.1927, O.H. Swezey Collector, Lysimachia [rotundifolia], | Holotype | Philodoria lysimachiella Swezey Det. by O. H. Swezey| Type no. 4270, in BPBM (here designated). Paralectotype 5♂, 2♀, same data and locality as holotype, | Paratype | 34188|♂ genitalia slide Z-XII-20-62-1| BPBM 34189 ♀, BPBM 34190 ♀, BPBM 34191♂, BPBM 34192♂ and 2♂ in USNM. Described from nine specimens: the holotype was originally in the collection of the Hawaiian Entomological Society; paratypes in author’s collection, Bishop Museum and the U.S. National Museum. We found eight “types” in the BPBM and USNM. The holotype was not specified in the description, and the so-labeled holotype and paratype are to be considered syntypes under Article 73.2 of the Code (ICZN 1999), and any one is therefore eligible for designation as lectotype under Article 74 of the Code (ICZN 1999). The syntype labeled, ‘Holotype’ is formally designated here as lectotype (Fig. 8A). The remaining syntype is the paralectotype. Additional material. 2♂, 3♀, 3 (sex unknown), Waianae (21.501325, -158.16538), Oahu, 28.iii.2016 (stored), K. Bustamente leg., host: Lysimachia hillebrandii, 9.xi.2015 /KMB01,CJ530 / SK 654♀, SK 655♀, SK656♂, SK657♂, SK751(wing vein); deposited in BPBM. Diagnosis. Very similar to P. molokaiensis Swezey feeding on same host plant, Lysimachia, but is distinguished by the median white streak reaching from base to near middle of wing and the lacking an oblique transverse fascia (cl + bb 2 ) from the base via costal fold, curved at costal 1/4 to dorsal 1/ 2 in the forewing (Fig. 8A) and the broad valva with short slender dorsal process and the rather slender pointed saccus in the male genitalia (Figs. 41E, 42E); oval-shaped lamella antevaginalis rather small and signa with rather small spines in the female genitalia (Fig. 56G). Redescription: Adult (Figs. 8A, 26H, 37B, 38). Wingspan 6.7–7.7 mm in paratypes; forewing length 2.5 mm in holotype, 3.0– 3.7 mm in paratypes. Head light gray; frons white; maxillary palpus reduced; labial palpus white, slightly grayish externally on the apical half of middle joint, terminal joint blackish in front (Fig. 37B). Antenna light gray, white below on basal third, very long, about 1.5 x length of forewing. Thorax light gray, patagium ocherous-orange. Forewing shiny, metallic bronze with ocherous-orange patches: a large oblique patch from base to 1/2, containing two longitudinal streaks, a median white one from base to near middle of wing lying slightly in front of the fold, bent dorsally in the outer 1/4; another bronze one same length, on the dorsal margin; a large tp after the middle to costal 3/4, distinctly narrowing in the dorsum, extending to dorsal 2/3, containing white cs; one white band on the middle of the first bronze color band (bb 2 ), another on costa to the middle of second band (bb 3 ), dorsally suffused with shining black scales; apical portion with a fuscous patch extending toward the termen and apex with a bronze as with orange-ocherous color encroaching on the apex and a small white spot at tornus; cilia light gray, with a white spot at beginning of costal cilia, remainder of cilia with black scales at base. Hindwing light gray, slightly infuscated towards apex, cilia light gray. Abdomen light gray, white beneath. Legs mostly light gray, white beneath. Male genitalia (Figs. 41E, 42E, 43E, 62A, B) (n=1). Capsule 900 µm. Tegumen 0.5–0.6 x length of valva; valva 480–500 µm long, very broad, apically rounded and having a slender dorsal process (Fig. 41E). Saccus short and tapering toward the point (Fig. 42E). Phallus 780–800 µm long and sinuous at phallobase in lateral view with two series of spiny minute cornuti in vesica (Fig. 43E). Female genitalia (Fig. 56G) (new record) (n=2). 1290 µm long. Ostium bursae small; antrum slender, cylindrical; lamella antevaginalis 200 µm, well-developed, oval-shape. Corpus bursae 600 µm long; paired signa with a pair of claw-shaped spines. Distribution. Oahu (Swezey 1928). Host plants. Primulaceae: Lysimachia hillebrandii Hook. f. (Swezey 1928). Biology. Unknown, but it is presumably similar to that of P. molokaiensis. Remarks. Leaf mines were once collected from Lysimachia along a ridge of Kalihi Valley on Oahu, but no adult moths were reared from them (Swezey 1928). Tim Kroessig (pers. comm.) found Philodoria leaf mines on L. remyi ssp. subherbacea, which were collected in Wai’anae mountains, Oahum,, but larvae from these mines were not reared., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 32-33, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Swezey, O. H. (1928) Some new species of lepidopterous leaf-miners in Hawaii. Proceedings of the Hawaiian Entomological Society, 7 (1), 187 - 191.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","ICZN (1999) International code of zoological nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 306 pp."]}

Philodoria pipturicola Swezey, 1915 Figs. 11A–D, 44C, D, 45B, 46B, 57D, E. Philodoria pipturicola Swezey, 1915: 96–97; Zimmerman 1978a: 709, figs. 464. Type locality. Punaluu (Oahu). Type material. Lectotype ♂, Punaluu, Oahu, 13.ix.1914, O.H. Swezey, ex Pipturus, |215 TYPE OF Philodoria pipturicola Swezey | right top species on a mount with three others is the lectotype, Type no. 215 in BPBM designated by Zimmerman (1978a). Paralectotype 9 (4♂, 4♀, 1 sex unknown): 1♂, 2♀, same mount of lectotype in BPBM; 1♂, 1♀, same data as locality as lectotype, | Philodoria pipturicola Paratypes Swezey| SK715♂ (right specimen), SK 716♀ (left specimen), BPBM 34257; 2♂, 1♀, 1 (sex unknown), same data and locality as lectotype, | Philodoria pipturicola Swezey |, SK 717♀ (top right one), SK829♂ (top left specimen), SK830♂ (bottom left specimen) in USNM. This species was described from ten specimens: four specimens on type mount no. 215 and six ‘paratypes’ reared from mines in the leaves of Pipturus. Given the manner in which the descriptions were written, Swezey likely considered these specimens as the holotype and paratypes, as indicated on their specimen labels. Zimmerman (1978a, fig. 464) designated a lectotype selected from the series of four syntypes on the type mount in the BPBM. Diagnosis. The forewing pattern is similar to P. pipturiella Swezey, but distinguished by an oblique white patch at near middle and two small white bands in the forewing (Fig. 11A–D). Redescription: Adult (Fig. 11A–D). Wingspan 6–7 mm in type series; forewing length 3 mm in lectotype. Head grayish fuscous; frons white; maxillary palpus reduced; labial palpus white, median joint slightly fuscous externally. Antenna dark fuscous. Thorax slate grey fuscous. Forewing fuscous suffused with orange patches at apical portion, with white spots: a cl from near base to about 2/3, apical potion bends inward; two white bands (w 1, 2 ) on both extremities of pale blue color band (bb 3 ) at 3/4, widest in the the middle; an oblique ws near middle of the wing, narrowly toward dorsum and not reaching dorsal margin; cilia fuscous with two white spots in costal cilia, a at the end of w 1 , b smaller and a little nearer the apex; a large pale-bluish patch (as) in terminal cilia; Hindwing and cilia dark fuscous. Abdomen dark fuscous, white beneath. Legs fuscous, white beneath. Male genitalia (Figs. 44C, D, 45B, 46B) (n=3). Capsule 700 µm. Tegumen 0.8–0.9 x length of valva; valva 510–530 µm long, slender and tapering toward the apex; dorsal margin of valva slightly angled with short spines; a series of small spines arranged along the inner margin of valva (Fig. 44C, D). Phallus 500 µm long, straight in lateral view with two series of minute cornuti in vesica (Fig. 46B). Female genitalia (Fig. 57D, E) (n=2). 1050 µm long. Ostium bursae large; antrum cup-shaped with a slender pair of lateral lobes; lamella antevaginalis 160 µm, weakly sclerotized, widening toward anterior margin of A7. Ductus bursae 270–300 µm; terminus of ductus bursae biforked; corpus bursae oblong, 640 µm; paired rows of wrinkles running longitudinally, some sclerotized, indistinct at middle. Distribution. Oahu (Swezey 1915). Host plants. Urticaceae: Pipturus sp. (Swezey 1915). Biology. Swezey (1915: 97) reported the mine form as “at first serpentine, later a blotch. The larva emerges to spin a light brownish cocoon on some convenient surface.” Immature stages reported by Swezey (1915: 96): “ Mature larva. 7 mm. Pale yellowish in colouration; head with a dark brown or blackish spot in each lobe; deeply bilobed and mostly retracted into segment 2, which is wider than the rest, other segments gradually tapering backwards; distinct constrictions between segments; cervical shield somewhat roughened, two pale brownish longitudinal streaks; thoracic legs feeble; abdominal prolegs on segments 7-9. Pupa. 3 mm. very pale brownish, abdomen yellowish in colouration; wing-sheaths pointed, extending to 5th abdominal segment; posterior leg-sheaths extend to apex of abdomen; antenna-sheaths extend beyond apex of abdomen and curve over dorsally and forward about two segments.” Parasitoids. Eulophidae: Euderus metallicus (Ashmead, 1901), Pnigalio externa (Timberlake, 1927) (Zimmerman1978a). Remarks. Maui records were misidentified, see description of P. haelaauensis. Johns et al. (2016) recorded this species from Palikea, Oahu (Coll ID CJ-101 / GenBank accession no. ID KT982409). CJ101 was sacrificed entirely for molecular analysis and CJ093 which is in the same series as CJ101, only has its genitalia and abdomen remaining. Female genitalia of CJ093 / PHIL0020 lacks signa on its corpus bursae and is similar to that of P. costalis, P. pipturiella, P. pipturicola. Because we do not have sufficient data, we here do not assign a formal new name to this species., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 49-54, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Swezey, O. H. (1915) New species of Hawaiian moths. P roceedings of the Hawaiian Entomological Society, 3 (2), 94 - 97. https: // doi. org / 10.5962 / bhl. part. 24594","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","Walsingham, Thomas, de Grey (Lord) (1907) Microlepidoptera. In: Sharp, D. (Ed.), Fauna Hawaiiensis. 1 (5). Cambridge University Press, Cambridge, pp. 469 - 759.","Swezey, O. H. (1928) Some new species of lepidopterous leaf-miners in Hawaii. Proceedings of the Hawaiian Entomological Society, 7 (1), 187 - 191.","Swezey, O. H. (1934) New species of Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 8 (3), 523 - 525.","Swezey, O. H. (1923) The leaf-miners of Pipturus. Proceedings of the Hawaiian Entomological Society, 5 (2), 293 - 296.","Swezey, O. H. (1940) New species of Hawaiian Lepidoptera. Proceeding Hawaiian Entomological Society, 10 (3), 461 - 465.","Swezey, O. H. (1920) Some new Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 4 (2), 376 - 386. https: // doi. org / 10.5962 / bhl. part. 16149","Swezey, O. H. (1946) New species of Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 12 (3), 625 - 628.","Meyrick, E. (1928) Some new species of Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 7 (1), 104.","Johns, C. A., Moore, M. R. & Kawahara, A. Y. (2016) Molecular phylogeny, revised higher classification, and implications for conservation of endangered Hawaiian leaf-mining moths (Lepidoptera: Gracillariidae: Philodoria). Pacific Science, 70 (3), 361 - 372. https: // doi. org / 10.2984 / 70.3.7"]}

Philodoria auromagnifica Walsingham, 1907 Figs. 7E, 26B, 41B, C, 42B, C, 43B, C, 56C, 61A–G, 79H. Philodoria auromagnifica Walsingham, 1907: 718, pl. 25, fig. 20; Swezey 1913b: 223; Zimmerman 1978a: 695, figs. 461, 468, 474; Kobayashi et al. (2018): 130 –132, figs. 2E, F, 5E–H, 6E–H, 7G, 8B, 11, 14C. Type locality. mountains, 2000 ft near Honolulu (Oahu). Type material. Holotype ♂, Mts. 2000 ft near Honolulu, Oahu, 25.x.1892, Perkins. 25857|BM slide no. 472|Walsingham Collection. 1910–427.| NHMUK010305330 | in NHMUK. Diagnosis. Very similar to P. succedanea, but distinguished from it by the dark brownish orange patches and brownish orange basal patch in the forewing; a fuscous patch with dark orangish scales in the apical portion (Fig. 7E); in the male genitalia by the rather long valva narrowing in the middle, vinculum large, inflexed on the ventral side, broad and straight saccus (Figs. 41B, C, 42B, C); in the female genitalia by signa with rather blunt spines (Fig. 55C) (Kobayashi et al. 2018). See also diagnosis of P. succedanea. Adult (Fig. 7E). Wingspan 8 mm in holotype. Male genitalia (Figs. 41B, C, 42B, C, 43B, C, 61A–C) and Female genitalia figured (Figs. 56C, 61G). See also Kobayashi et al. (2018). Distribution. Kauai (Kobayashi et al. 2018), Oahu (Walsingham 1907), Molokai (Swezey & Bryan 1929), and Hawaii (Big Island) (Zimmerman 1978a). Host plants. Primulaceae: Myrsine lessertiana A. DC. and M. sandwicensis A. DC. (Johns et al. 2016), M. wawraea (Mez) Hosaka (Kobayashi et al. 2018), Myrsine sp. (Swezey 1913a). Biology. (Fig. 79H). Kobayashi et al. (2018: 131, figs. 8B, 11, 14C) reported the leaf mine and pupal cocoon. Parasitoids. Eulophidae: Euderus metallicus (Ashmead, 1901) (Zimmerman 1978a)., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 25-26, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Walsingham, Thomas, de Grey (Lord) (1907) Microlepidoptera. In: Sharp, D. (Ed.), Fauna Hawaiiensis. 1 (5). Cambridge University Press, Cambridge, pp. 469 - 759.","Swezey, O. H. (1913 b) Leaf-miners of the Hawaiian Islands. Proceedings of the Hawaiian Entomological Society, 2 (5), 221 - 227.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","Kobayashi, S., Johns, C. A., Lopez-Vaamonde, C., Doorenweerd, C., Kawakita, A., Ohshima, I., Lees, D. C., Hanabergh, S. & Kawahara, A. Y. (2018) Hawaiian Philodoria (Lepidoptera, Gracillariidae, Ornixolinae) leaf mining moths on Myrsine (Primulaceae): two new species and biological data. ZooKeys, 773, 109 - 141. https: // doi. org / 10.3897 / zookeys. 773.21690","Swezey, O. H. & Bryan, E. H. (1929) Further notes on the forest insects of Molokai. Proceedings of the Hawaiian Entomological Society, 7 (3), 293 - 314.","Johns, C. A., Moore, M. R. & Kawahara, A. Y. (2016) Molecular phylogeny, revised higher classification, and implications for conservation of endangered Hawaiian leaf-mining moths (Lepidoptera: Gracillariidae: Philodoria). Pacific Science, 70 (3), 361 - 372. https: // doi. org / 10.2984 / 70.3.7","Swezey, O. H. (1913 a) A day's collecting at Punaluu, Oahu. Proceedings of the Hawaiian Entomological Society, 2 (5), 197 - 199.","Swezey, O. H. (1934) New species of Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 8 (3), 523 - 525.","Swezey, O. H. (1923) The leaf-miners of Pipturus. Proceedings of the Hawaiian Entomological Society, 5 (2), 293 - 296."]}

Philodoria micropetala Walsingham, 1907 Fig. 10F Philodoria micropetala Walsingham, 1907: 719, pl. 25, fig. 22; Zimmerman 1978a: 709, figs. 463, 476. Type locality. Halemanu (Kauai). Type material. Holotype ♀, Halemanu, 4000ft, Kauai, v.1895., Perkins 27560, | PHILODORIA MICROPETALA, Wlsm Fn Hawaii I descry. figd | Walsingham Collection. 1910–427.| NHMUK010305336 |TYPE|BM ♀ Genitalia slide no. 3951| in NHMUK. Described based on a single specimen from Kauai. The ‘type’ specimen, designated by Walsingham is here thus the holotype following article 73.1.2 (ICZN 1999). Diagnosis. The forewing pattern is similar to that of P. floscula, but distinguished by an erect dorsal ws in the forewing (Fig. 10F). The female genitalia were incompletely described by Zimmerman (1978a: fig. 476) who mentioned only the ostium and apophyses. Description: Adult (Fig. 10F). The following is from Walsingham (1907: 719): “Antennae brownish. Palpi white, a dark spot at the end of the median joint, the terminal joint also shaded with fuscous. Head and Thorax bronze. Forewings bronze, with four white spots; the first on the outer half of the fold, not touching the dorsum; the second on the costa beyond the middle, pointing obliquely outward and dark-margined externally; the other two situated as in floscula, but dark-margined on their inner sides, the metallic spot between them being bright steelblue; beyond this an orange spot, with three white streaks in the costal cilia above it, alternate with brownish, a few fuscous scales scattered across the middle of this patch terminate in a blackish spot at its outer edge, followed by shining, steel-blue, which extends through the ternlinal and apical cilia. Exp. al. 8 mm. Hindwings tawny brown; cilia tawny. Abdomen fuscous above, whitish beneath. Legs brownish, spurs and a few tarsal spots whitish.” Male genitalia Unknown. Female genitalia See Zimmerman (1978a: fig. 476). Ostium bursae large; antrum wide with a slender pair of lateral lobes; lamella antevaginalis weakly sclerotized, widening toward anterior margin of A7. Ductus bursae and corpus bursae not figured. Distribution. Kauai (Walsingham 1907). Host plants. Urticaceae: Pipturus sp. (Zimmerman 1978a). Biology. Unknown. Parasitods. Eulophidae: Euderus metallicus (Ashmead, 1901), Neochrysocharis formosus (Westwood, 1833) (Zimmerman 1978a). Remarks. Zimmerman (1978a) noted that the records from outside Kauai, i.e. from the islands of Hawaii (Big Island), Maui, Molokai, and Oahu mentioned by Swezey (1913b: 222) were likely in error and attributable to other Pipturus miners; P. micropetala appears to be restricted to Kauai. We accept Zimmerman’s suggestion in recognizing that only the specimens from Kauai are attributable to this species., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 57-58, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Walsingham, Thomas, de Grey (Lord) (1907) Microlepidoptera. In: Sharp, D. (Ed.), Fauna Hawaiiensis. 1 (5). Cambridge University Press, Cambridge, pp. 469 - 759.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","ICZN (1999) International code of zoological nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 306 pp.","Swezey, O. H. (1913 b) Leaf-miners of the Hawaiian Islands. Proceedings of the Hawaiian Entomological Society, 2 (5), 221 - 227."]}

Philodoria Walsingham, 1907 is a threatened, Hawaiian endemic genus of leaf-mining gracillariid moths that feeds as larvae on many threatened and endangered Hawaiian endemic plants. These moths are poorly studied and species lack detailed descriptions of morphology, distribution data, and natural history information of adults and immatures. Based on extensive fieldwork from 2013 to 2016, and examination of museum specimens, we describe or redescribe 51 species, 13 which are new species and provide biological and distribution data for 41 species. The 13 new species and their host plants are: P. alakaiensis Kobayashi, Johns & Kawahara, sp. n. (Asteraceae: Dubautia sp.), P. funkae Kobayashi, Johns & Kawahara, sp. n. (Asteraceae: Wilkesia gymnoxiphium), P. haelaauensis Kobayashi, Johns & Kawahara, sp. n. (Urticaceae: Pipturus albidus, P. rockii, Pipturus sp.), P. hesperomanniella Kobayashi, Johns & Kawahara, sp. n. (Asteraceae: Hesperomannia arborescens and H. swezeyi), P. keaensis Kobayashi, Johns & Kawahara, sp. n. (host unknown), P. keahii Kobayashi, Johns & Kawahara, sp. n. (Asteraceae: Remya mauiensis), P. knudseniiella Kobayashi, Johns & Kawahara, sp. n. (Asteraceae: Dubautia knudsenii subsp. nagate and D. latifolia), P. lama Kobayashi, Johns & Kawahara, sp. n. (Ebanaceae: Diospyros sandwicensis and/or D. hillebrandii), P. limahuliensis Kobayashi, Johns & Kawahara, sp. n. (Malvaceae: Hibiscus waimeae subsp. hannerae), P. napaliensis Kobayashi, Johns & Kawahara, sp. n. (Ebanaceae: Diospyros sandwicensis and/or D. hillebrandii), P. obamaorum Kobayashi, Johns & Kawahara, sp. n. (Urticaceae: Pipturus albidus, Pipturus sp.), P. opuhe Kobayashi, Johns & Kawahara, sp. n. (Urticaceae: Urera kaalae and U. sandvicensis) and P. platyphylliella Kobayashi, Johns & Kawahara, sp. n. (Asteraceae: Dubautia platyphylla). Types of 30 species were examined, lectotypes of 16 species are designated, and a key to all Philodoria species and all Hawaiian leaf-mining moths are provided. We also present a new Hawaiian name for Philodoria, which we call Hunelele ʻelilau, meaning “tiny flier, leaf excavator”, referring to their life history and behavior. Philodoria feeds on five herbraceous and woody host plant families, and 41 species persist in localized populations in Hawaii as of 2016. Twelve species (P. alakaiensis, P. funkae, P. haelaauensis, P. hespermanniella, P. kauaulaensis, P. keaensis, P. keahii, P. knudsniiella, P. limahuliensis, P. platyphylliella, P. sciallactis, P. wilkesiella) appear to be severely threatened, as these species are rare or feed exclusively on plants that are endangered or rare. We were unable to document 10 species (P. costalis, P. lipochaetaella, P. micropetala, P. nigrella, P. nigrelloides, P. opuhe, P. pipturiana, P. pipturicola, P. pittosporella, P. spilota) after many efforts to sample in or near their historical habitats. We believe these species may be extinct. Given their restricted distribution and the fact that many species feed on rare and endangered plants, there is a risk that many more Philodoria species may go extinct unless immediate conservation measures are taken.

Philodoria touchardiella (Swezey, 1928) Figs. 21E, 27N, O, 50H, 51F, 52F, 59J, 94. Parectopa touchardiella Swezey, 1928: 189. Philodoria touchardiella (Swezey, 1928); Zimmerman 1978a: 690, figs. 445, 452, 460. Type locality. Black Gorge, north side of the Iao Valley (Maui). Type material. Lectotype ♀, North side of Iao Valley, Black Gorge, 1800 ft., Maui, 15.i.1926 O.H. Swezey Collector, host: Tonchardia latifolia, Type no. 4265 in BPBM (here designated). Described from two specimens from Maui. Swezey noted that ‘ holotype and paratype in the collection of the Hawaiian Entomological Society’. However, because a holotype was not specified in the original description, the so-labeled holotype and paratype can be treated as syntypes under Article 73.2 of the Code (ICZN 1999), and any specimen can designated as lectotype under Article 74 of the Code (ICZN 1999). The syntype labeled ‘holotype’, which Zimmerman (1978a) treated as the holotype, is here formally designated as lectotype (Fig. 21E). The other syntype is the paralectotype, but we could not find ‘Paratype’ and its genital slide no. Z-VII-14-61 (Zimmerman 1978a, fig. 452) in BPBM, we only found a slip of paper in the specimen’s box that read, ‘ paratype ♂ sent E.C.Z 6-5-61’. Additional material. (3♂, 2♀, 1 sex unknown). Adult: 1♂, 1♀, Kaluaaha, Molokai, 14.iii.2015 (stored), K. Bustamente leg., host: Tonchardia latifolia, 5.xii.2014, KMB04|CJ401a / SK 764♀, CJ401b / SK765♂ in BPBM; Maui Is: 2♂, Iao Valley, 11.viii.2014 (stored), host: Tonchardia latifolia, CJ 365 / SK642♂, 643♂ in BPBM; 1♀, East Wailuaiki Stream, 26.vii.2015 (stored), K. Bustamente leg., host: Tonchardia latifolia, 26.vi.2015, KMB01|CJ507b / SK 766♀ in BPBM. Pupa: 1 (sex unknown), same locality and data as holotype, BPBM34326. Diagnosis. The forewing pattern and male genitalia are very similar to that of P. nigrella (Walsingham), but is distinguishable by its rather brown forewing with ds 1 that does not reach the middle of the wing and slightly broad white ds 2 (Fig. 21E). The female genitalia differ from other congeners in having a larger corpus bursae and a pair of small linear signa in the middle with a series of minute spines (Fig. 59J). Redescription: Adult (Fig. 21E). Wingspan 6 mm in type series; forewing length 3.25 mm in holotype. Head light buff; frons white; maxillary palpus white; labial palpus white, median joint slightly infuscated externally at apex, terminal joint infuscated in front. Antenna light buff, infuscated on apical half, a little longer than forewing. Thorax, abdomen and legs olive brown, pale beneath. Forewing olive brown with white patches: three large triangular white dorsal streak (ds 1–3 ) extending obliquely outward: ds 1 at near base, ds 2 at about middle, longer and narrower reaching to middle of wing, and ds 3 near tornus; an outwardly oblique white cs 3 at 3/4, with a few blue scales at its terminus; apical portion beyond the cs 3 with a few yellow scales, and a few blue scales at extreme apex; cilia olive brown, with three white spots (a, b, c) in costal cilia alternating with black spots, terminal cilia with bl 1 at base. Hindwing and cilia nearly uniform olive brown. Male genitalia (Figs. 50H, 51F, 52F) (n=3). Capsule 820 µm. Tegumen 0.8–0.9 x length of valva; valva 570 µm long, tapering along costal margin and slightly sinuous from basal 2/5 to apex (Fig. 50H). Saccus rather long and stick-shaped in ventral view (Fig. 51F). Phallus 520 µm long and straight, widening at phallobase with small coecum; cornuti in vesica indistinct (Fig. 52F). Female genitalia (Fig. 59J) (n=2). 1280 µm long. Ostium bursae large; antrum cup-shaped with a pair of lateral lobes; lamella antevaginalis 210 µm, weakly sclerotized, semicircular in ventral view. Ductus bursae rather wide, middle part weakly sclerotized, round and flat; terminus of ductus bursae tubular, biforked. Corpus bursae 720 µm, pyriform; anterior end of corpus bursae weakly sclerotized with some minute wrinkles; two short linear signa with a series of minute spines. Distribution. Molokai: new record, and Maui (Swezey 1928). Host plants. Urticaceae: Touchardia latifolia Gaudich. (Swezey 1928). Biology. (Fig. 94). Larvae form a broad linear mine (Fig. 94). Usually two or more mines per leaf. Pupation occurs in the cocoon outside of the mine on the leaf surface. The leaf mine quickly becomes discolored, turning black or dark gray, and can be mistaken for other forms of leaf damage. When Touchardia is present, leaf mines tend to be abundant. Remarks. It is possible that this moth occurs on other Hawaiian Islands, but we were unable to survey Touchardia on the islands of Hawaii (Big Island), Lanai, and Oahu., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 98-99, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Swezey, O. H. (1928) Some new species of lepidopterous leaf-miners in Hawaii. Proceedings of the Hawaiian Entomological Society, 7 (1), 187 - 191.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","ICZN (1999) International code of zoological nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 306 pp."]}

Philodoria sciallactis (Meyrick, 1928) Figs. 24A–G, 53A, B, 54A, 55A, 60D, 78A–C, 91A–G. Parectopa sciallactis Meyrick, 1928:104. Parectopa lipochaetae Swezey, 1946: 627. A junior subjective synonym of Parectopa lipochaetaella Swezey, 1940, synonymized by Zimmerman (1978a: 681). Philodoria sciallactis (Meyrick, 1928); Zimmerman 1978a: 690, figs. 445, 451, 458; Johns et al. 2016: 369, figs. 1, 2; Johns et al. 2018: fig. 2. Philodoria sp. 6; Johns et al. 2018: fig. 2. Type locality. Kaena Point (Oahu). Type material. Lectotype ♂, Kaena Point, Oahu, 11.xi.1927, O.H. Swezey Collector, Lipochaeta [= Melanthera integrifolia], | Holotype | Parectopa sciallactis Mey. teste Meyr.| in BPBM. (here designated) Paralectotype 6 (3♂, 3♀). [Same data and locality as lectotype with following label: | Paratype | Parectopa sciallactis Mey. teste Meyr.|]: 2♂, 1♀, ♂| BPBM 34280, SK 713♀ | BPBM 34281, SK711♂| BPBM 34282; 1♂, |B.M. Genitalia slide no. 2754|Meyrick Coll. B.M. 1938-290|BMNH(E) 1407859| NHMUK 010305339| Paratype | in NHMUK; 1♀, | PARECTOPA Clem. | sciallactis Meyr. |BMNH(E)1407866| NHMUK 010305340| Paratype | in NHMUK; 1♀, SK 712♀ in USNM. Described from seven specimens reared from ‘fleshy leaves of Lipochaeta integrifolia [= Melanthera integrifolia], growing in rocky places near the seashore’ (Meyrick 1928). Specimen labels suggest that Meyrick considered one of these specimens to be the holotype and the remaining specimens as paratypes. However, because a holotype specimen was not formally designated in the original description, the so-labeled holotype and paratypes can be considered syntypes under Article 73.2 of the Code (ICZN 1999), and any of them are eligible for designation as the lectotype under Article 74 of the Code (ICZN 1999). The syntype labeled ‘Holotype’ is here formally designated as lectotype (Fig. 24B) and the remaining syntypes are paralectotypes. Additional material. (5♂, 1♀, 2 sex unknown) 1♂, Mokio, Molokai, 29.xii.2013 (stored), C.A. Johns leg., host: Melanthera rockii [= Lipochaeta rockii], 16.xii.2013, CJ159, SK759♂ in BPBM; Maui: 1♂, 1♀, 2 (sex unknown), Lahaina, 23.xii.1928, O.H. Swezey Collector, ex Lipochaeta larvarum, BPBM 34184, SK699♂| BPBM 34185, BPBM 34186,| Holotype Parectopa lipochaetae Swezey |Type no. 4259|; 3♂, Lualailua, 30.xii.2013 & 4.i.2014 (stored), C.A. Johns leg., host: Melanthera kamolensis, 24.xii.2013, CJ163 / SK688♂, CJ164 / PHIL0024♂, CJ189 / SK689♂ in BPBM. 1♀, 2 (sex unknown), Kaena Point, Oahu, 6–11.vii.2014 (stored in RNAlater solution), C.A. Johns leg., host: Melanthera integrifolia, 1.vii.2014, AHE_56, CJ271 / PHIL0034 ♀, CJ273 (no abdomen), CJ274 (no abdomen), ‘ Philodoria sp. 6 of Johns et al. (2018) ’, in FLMNH. Diagnosis. The forewing pattern is similar to that of P. marginestrigata (Walsingham), but this specimen can be distinguished by its large white patches on the forewing (Fig. 24A–G). See also diagnosis of P. lipochaetaella. Redescription: Adult (Fig. 24A–F). Wingspan 4.5–6 mm in paralectotype; forewing length 1.8–2.5 mm in paralectotype. Head and frons whitie; maxillary palpus white; labial palpus white, sometimes with fuscous rings at apex of second segment and in middle of terminal segment. Antenna white ringed fuscous. Thorax white to graywhite. Forewing white, sometimes partially suffused light gray with light ocherous to yellow patches, margined with fuscous scales: a median streak (mp) from base to 1/3, meeting apex of an oblique narrow line from base to 1/8 of near dorsum; a small spot at dorsal 1/7 (dp 1 ); three oblique costal patches (cp 1–3 ): cp 1 about 1/2, cp 2 at 3/4, reaching half across wing; cp 3 short, before apex; an oblique dp 2 at 1/2, meeting cp 1 and cp 2 ; apical portion white with light ocherous scales at middle, containing a black as; cilia white or white gray, with bl 1 and bl 2 from termen to tornus. Hindwing gray; cilia white. Abdomen ocherous. Legs white, tarsi with fuscous scales. Male genitalia (Figs. 53A, B, 54A, 55A) (n=6). Capsule 590 µm. Tegumen 0.9 x length of valva; valva 420 µm long, tapering along costal margin from 1/2 to apex; apical half digitiform, slender and slightly curved toward inner margin with small spines at apex; a set of short spines arranged along inner side of basal region (Fig. 53A, B). Saccus short and triangular in ventral view (Fig. 54A). Phallus 400 µm long and slightly widening at phallobase with developed coecum; cornuti in vesica indistinct (Fig. 55A). Female genitalia (Fig. 60D) (n=2). 1060 µm long. Ostium bursae large; antrum large, cup-shaped with a pair of lateral lobes; lamella antevaginalis 140 µm, weakly sclerotized, inflexed on the posterior margin, widening toward anterior margin of A7. Ductus bursae rather short, middle region weakly sclerotized, round and flat; terminus of the ductus bursae tubular, biforked. Corpus bursae 640 µm, pyriform; anterior end of corpus bursae weakly sclerotized; signa absent. Distribution. Oahu (Meyrick, 1928), Molokai and Maui: new record. Host plants. Asteraceae: Melanthera integrifolia (Nutt.) W.L.Wagner & H.Rob. (Meyrick 1928; Johns et al. 2016), M. kamolensis (O.Deg. & Sherff) W.L.Wagner & H.Rob. (Endangered, IUCN) and Lipochaeta rockii Sherff are new record. Biology. (Fig. 91A–G) The larva mines the fresh leaves of Melanthera integrifolia forming a blotch mine which expands to cover the entire leaf as the larva grows (Fig. 91A, B). Pupation occurs in the cocoon outside of the mine; cocoon is white ellipsoid (Fig. 91C). Mines on M. integrifolia are very difficult to find. Larvae that feed on Lipochaeta rockii form a blotch mine and have a white, ellipsioid pupal cocoon (Fig. 91D, E). On a leaf of M. kamolensis, we observed a similar cocoon to that of L. rockii. Cocoons were often found on the adaxial leaf surface, centered on the midvein. Remarks. We surveyed some Melanthera populations in the lowlands of Maui but did not find any Philodoria mines. The East Maui specimens feeding on Melanthera kamolensis have generally darker brown patches (Fig. 24E–G), similar to specimens from the type series of P. lipochaetaella, but these specimens are P. sciallactis based on male genitalia. The East Maui specimens are from the same series as P. sciallactis of Johns et al. (2018: fig. 2, AHE_4/CJ-145). Pholodoria sp. 6 of Johns et al. (2018) is here treated as P. sciallactis based on female genitalia: three specimens of P. sp. 6 (CJ271, 273 & 274) were collected from the type locality of P. sciallactis and female genitalia of CJ271 is poor condition, but similar to that of paralectotype of P. sciallactis. Two specimens (CJ273 & 274) were entirely sacrificed for molecular analysis (Johns et al. 2018), and no voucher remains. Among specimens from the populations on Oahu, Molokai and Maui, there is phenotypic wing pattern variation and a detailed DNA barcoding study in the future may prove beneficial to futher determine the species boundaries of P. sciallactis and P. lipochaetaella., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 104-109, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Meyrick, E. (1928) Some new species of Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 7 (1), 104.","Swezey, O. H. (1946) New species of Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 12 (3), 625 - 628.","Swezey, O. H. (1940) New species of Hawaiian Lepidoptera. Proceeding Hawaiian Entomological Society, 10 (3), 461 - 465.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","Johns, C. A., Moore, M. R. & Kawahara, A. Y. (2016) Molecular phylogeny, revised higher classification, and implications for conservation of endangered Hawaiian leaf-mining moths (Lepidoptera: Gracillariidae: Philodoria). Pacific Science, 70 (3), 361 - 372. https: // doi. org / 10.2984 / 70.3.7","Johns, C. A., Toussaint, E. F. A., Breinholt, J. W. & Kawahara, A. Y. (2018) Origin and macroevolution of micro-moths on sunken Hawaiian Islands. Proceedings of the Royal Society B, 285, 20181047. https: // doi. org / 10.1098 / rspb. 2018.1047","ICZN (1999) International code of zoological nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 306 pp."]}

Philodoria epibathra (Walsingham, 1907) Figs. 13A–C, 26O, P, 47B, 48B, 49B, 58B. Gracilaria epibathra Walsingham, 1907: 722, fig. 27. Parectopa epibathra (Walsingham, 1907); Swezey 1928: 190; Zimmerman 1978a: 679–680, fig. 459. Parectopa epibathra (Walsingham, 1907) (= Philodoria naenaeiella); Swezey 1928: 190 (Misidentification). Type locality. Molokai. Type material. Holotype ♀ (= ‘ Type ♂ (26056)’), Molokai, ‘000ft.’[elevation blank], Hawaiian Is., 12.vi.1893, Perkins 26056, | B.M. Genitalia slide no. 2884| GRACILARIA EPIBATHRA, Wlsm. Fm. Hawaii I TYPE ♂ descr. figd.| Walsingham Collection, 1910—427.| BMNH(E) 1407695| NHMUK010305333 | with capsulated left forewing in in NHMUK. Described based on a single specimen from Oahu. The ‘type’ specimen, labeled on the specimen by Walsingham is here formally designated as the holotype following article 73.1.2 (ICZN 1999). Additional material. Molokai: 2♂, 2♀, Olokui, 26.vii.2015 (stored), KB Collection, host: Dubautia laxa, 28.i.2015, KMB19, CJ506a–CJ506d, SK678–681. 1♀, Waiehu, 21.vii.2015 (stored), C.A. Johns leg., host: Dubautia sp., 7.vii.2015, CJ492, SK 837♀. All specimens preserved in BPBM. Diagnosis. Similar to P. nigrella (Walsingham) and P. touchardiella (Swezey) in that forewing has cs 3 and ds 1–3 , but differs from them by the wide ds 2 and rather long ductus bursae and lack of spines on signa in the female (Figs. 13B, C, 58B). Male genitalia of this species differs from other congeneric species in having a very long saccus. We collected four adult moths reared from Dubautia; these adults have similar forewing pattern and female genitalia as the holotype except for the indistinct or lack of white ds 1 at basal 1/3 (Figs. 13B, C, 26O, P). Description: The following is from Walsingham (1907: 722): “Antennae brownish. Palpi white, a small spot on the outer side of the median joint at its apex. Head shining, dirty whtish, a few brownish scales posteriorly. Thorax olive-brow. Forewings olive-brown with three outwardly oblique broad pearly white dorsal streaks; the first near the base, somewhat rounded at its apex; the second at the middle, attenuate; the third near the tornus, quadrate; above the latter is an outwardly oblique slender white costal streak, followed by two smaller wllite streaks in the costal cilia before the apex; at the end of the oblique costal streak is a small patch of shining, blue scales, narrowly surrounded by pale ocherous; with a black spot at the apex, followed by shining blue at the base of the cilia; through these runs a curved metallic blue line, the space between which and an outer curved brownish line being whitish, tornal cilia smoky brownish. Exp. al. 7 mm. Hindwings brownish, with slightly paler cilia. Abdomen dark brown. Legs brownish, with whitish tarsal spots.” Male genitalia (Figs. 47B, 48B, 49B) (n=2). (new record). Capsule 910 µm. Tegumen 0.8 x length of valva; valva tapering along costal margin from base 2/5 to apex; basal portion humped along dorsal margin in lateral view (Fig. 47B). Saccus 230 µm, very long and wavy, slightly curved toward dorsal side (Fig. 48B). Phallus 690 µm long, straight in lateral view Fig. 49B). Female genitalia. (Fig. 58B) (n=4). (Zimmerman 1978a: fig. 459). 1240 µm long. Ostium bursae large; antrum low-cup-shaped with a slender pair of lateral lobes; lamella antevaginalis 100–110 µm, weakly sclerotized, semicircular in ventral view, widening toward anterior margin of A7. Ductus bursae thick, posterior extremity widening and membranous; terminus of ductus bursae biforked. Corpus bursae 540–550 µm, pyriform; paired rows consisting of wrinkles running longitudinally, some sclerotized. Distribution. Molokai (Walsingham 1907). Host plants. Asteraceae: Dubautia laxa Hook. & Arn.: new record and Dubautia sp. Biology. Larvae mine leaves of Dubautia species, according to collecting data labels of adult specimens. Remarks. “ P. epibathra ” had been frequently assigned to several Dubautia -feeding Philodoria species by Swezey. The Kauai record (Swezey 1928:190) is erroroneous (Swezey 1940; Zimmerman 1978a) and applies to Philodoria knudseniiella sp. n. The host plant records of Dubautia sp. (Swezey, 1910a: 89, footnote), Dubautia laxa, Hesperomannia arborescens (Swezey 1928: 190) are also in error and apply to host plants of Philodoria naenaeiella (Swezey, 1940: 463) and P. hesperomanniella sp. n. respectively. Some leaf mines were collected on Dubautia at Kamakou Boardwalk, Molokai. Some mines begin as spiral-shaped (Fig. 88L), but no adults were successfully reared from this plant., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 60-61, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Walsingham, Thomas, de Grey (Lord) (1907) Microlepidoptera. In: Sharp, D. (Ed.), Fauna Hawaiiensis. 1 (5). Cambridge University Press, Cambridge, pp. 469 - 759.","Swezey, O. H. (1928) Some new species of lepidopterous leaf-miners in Hawaii. Proceedings of the Hawaiian Entomological Society, 7 (1), 187 - 191.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","ICZN (1999) International code of zoological nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 306 pp.","Swezey, O. H. (1940) New species of Hawaiian Lepidoptera. Proceeding Hawaiian Entomological Society, 10 (3), 461 - 465.","Swezey, O. H. (1910 a). The ebony leaf-miner (Gracilaria [sic] mabaella). Proceedings of the Hawaiian Entomological Society, 2 (3), 88 - 90, pl. 3."]}

Philodoria pipturiana Swezey, 1923 Fig. 12A–C. Philodoria pipturiana Swezey, 1923: 295; Zimmerman 1978a: 709, figs. 465, 466. Type locality. Upper Hamakua Ditch Trail, Kohala Mts. (Big Island). Type material. Lectotype ♂, Upper Hamakua Ditch Trail, [Kohala Mountains, Hawaii (Big Island)], 31.vii.1921, Swezey Coll., Pipturus, | Philodoria pipturiana Swezey | Holotype |♂ genitalia on slide Z-I-10-67-A |Type no. 4272 in BPBM. Described from two specimens: the holotype was originally in the collection of the Hawaiian Entomological Society and the paratype in the collection of the Hawaiian Sugar Planter’s Association. The ‘holotype’ is now preserved in BPBM. Given the manner in which the description was written, Swezey considered these specimens as holotype and paratype, as indicated on their specimen labels. But because the holotype was not formally specified in the description, the so-labeled holotype and paratype are to be considered syntypes under Article 73.2 of the Code (ICZN 1999), and either is eligible for designation as lectotype under Article 74 of the Code (ICZN 1999). The ‘ Holotype, type no. 4272’, which Swezey indicated as holotype in the specimen label, is here designated as lectotype (Fig. 12A–C). The other syntype is the paralectotype, but we could not locate that specimen. Diagnosis. The forewing pattern is similar to P. wilkesiella Swezey, but is distinguished by its longitudinal basal streak and oval white spot in the middle in the forewing (Fig. 12A–C). The male genitalia are similar to those of the Asteraceae-feeding Philodoria species (e.g., P. wilkesiella and P. hesperomaniniella sp. n.) in having a broad dorsal margin of valva, but P. pipturiana differs from later two species by its slender saccus. Redescription: Adult (Fig. 12A–C). Wingspan of specimens in the type series 8–9 mm; forewing length 4 mm in lectotype. Head dirty white; frons white; maxillary palpus reduced; labial palpus white, streaked with fuscous externally. Antenna pale fuscous, apical portion white. Thorax dark fuscous, white below. Forewing dark fuscous to nearly black, with several white patches: one outwardly oblique white costal spot at about 1/2, another at 2/3, recurves to the costa, terminating in a few white scales in the costal cilia; a curved transverse white streak at apical portion of wing, nearly interrupted in the middle, with a few white scales in the costal cilia (not present in the paratype); a longitudinal white streak from base to 1/3, followed by a large oval white spot about the middle of wing; a large oval white spot on dorsal 3/4; cilia fuscous except for the white scales previously noted in costal cilia and a few white scales at base of apical cilia. Hindwing and cilia dark fuscous. Abdomen fuscous. Legs pale fuscous, tarsi white spotted. Male genitalia (Zimmerman 1978a: fig. 465). Tegumen oblong, 0.8 x length of valva; valva tapering along costal margin from base 2/5 to apex; dorsal margin of valva broad. Saccus slender. Phallus straight in lateral view with spiny cornuti in vesica. Female genitalia Unknown. Distribution. Hawaii (Big Island) (Swezey 1923). Host plants and Biology. Unknown. Remarks. Swezey (1923) described this species from two specimens collected on a Pipturus plant which had leaves containing leaf-miners, but according to Zimmerman (1978a) these specimens were not reared. Therefore, we have removed this plant genus as a possible host. Much of the type locality habitat has been destroyed due to non-native plant species, and few host plants remained in sections of the Upper Hamakua Ditch Trail that we surveyed., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 58-59, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Swezey, O. H. (1923) The leaf-miners of Pipturus. Proceedings of the Hawaiian Entomological Society, 5 (2), 293 - 296.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","ICZN (1999) International code of zoological nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 306 pp."]}

Philodoria alakaiensis Kobayashi, Johns & Kawahara sp. n. Figs. 14F, 15A–E, 36, 39D, 47E, 48E, 49E, 58F, G. Philodoria sp. 15; Johns et al. 2018: fig. 2. Type locality. Alakai (Kauai). Etymology. The specific epithet is derived from the type locality, Alaka’i (pronounced ‘Ala-Kayee’) Wilderness Preserve, also known as Alaka’i Swamp, an important wet forest area for Hawaiian endemic plants. Type material. Holotype ♂, Alakai, Kauai Is., 9.vii.2015 (stored), C.A. Johns leg., host: Dubautia laxa or D. paleata, 18.vi.2015, CJ473, SK 836 in BPBM. Paratype 7 (3♂, 3♀, 1 sex unknown): 1♂, same data and locality as holotype, CJ 468 in BPBM; 2♂, 2♀, 1 (sex unknown), same data and locality as holotype, 11.vii.2015 (stored), CJ479/SK658♂, SK 659♀, SK660♂, SK 661♀ in BPBM; 1♀, same locality as holotype, 11.vii.2015 (stored), 18.vi.2015, host: Dubautia laxa or D. paleata, CJ 481/SK 864♀, SK870 (SEM) in BPBM; 1♂, Kauai, 4520’, Alakai Swamp, Kelekua Hut, 13.ix.1973, K. & E. Sattler, B.M.1973-498, Philodoria sp. 30 cf nigrelloides, Colour slides 133-4, 139 Pl. 3 Fig. 5 Zimmerman 1978a Sattler Coll., D.C. Lees Sep. 2016, BMNH(E) 1621107 in NHMUK; 1♀, Kauai, 4520’, Alakai Swamp, Kelekua Hut, 5.vii.1973, K. & E. Sattler, B.M.1982-342, Philodoria sp. 28, Sattler Coll., D.C. Lees, Sep. 2016, BMNH(E) 1621109 in NHMUK. The holotype was mounted as a dry pinned specimen with its four wings placed without mountant under a coverslip: forewings slightly damaged at base, basal 2/3 of hindwings broken. Diagnosis. Dark brownish forewings with four outwardly oblique white streaks: two dorsal ones and two slender costal ones (Figs. 14F, 15A–E). Differs from other Philodoria species in having two costal streaks, but lacks a ds 1 and a short cs 1 in the forewing; valva slender and slightly rounded apically. An Alakai species, P. nigrelloides (Swezey) (male unknown, feeding on Dubautia) differs from the new species in lacking the cs 1 and having a tubular antrum (Zimmerman 1978a: fig. 459). Description: Adult (Figs. 14F, 15A–E, 36). Forewing length 3.8–3.9 mm in holotype, 3.3–3.9 mm in paratype. Head dark brown; frons white; maxillary palpus white; labial palpus white but with dark brown scales below at apex (Fig. 36); antenna dark brown, white beneath, about 1.1x length of forewing. Thorax brown. Forewing brown to dark brown with outwardly oblique white streaks that are found at: cs 2 short one at about 1/2, cs 3 slender at 3/4; ds 1 very small at base along the dorsum; ds 2 large at 1/3; ds 3 at 3/4; some fuscous scales scattering from 1/2 to 3/4; apical portion orangish brown; three white costal spots (a, b, c) near apex; a spot of bluish scales at apex (as) and a few bluish scales in a more or less fuscous patch between apex and end of ds 3 ; cilia fuscous with two white spot at termen; bl 1 black from termen to tornus, fuscous fringe at tornus. Hindwing and cilia fuscous. Legs pale fuscous with dark brown scales. Male genitalia (Fig. 39D, 47E, 48E, 49E) (n=2). Capsule 800 µm. Tegumen 0.7–0.8 x length of valva; valva 590 µm long, tapering along costal margin from base 2/5–1/2 to apex, slightly rounded at apex and narrowing once at middle; a series of small spines arranged, along the inner side of valva (Fig. 47E). Saccus triangular in ventral view (Fig. 48E). Phallus 550 µm long, rounded at phallobase with developed coecum; two series of spiny minute cornuti in vesica (Fig. 49E). Female genitalia (Fig. 58F, G) (n=3). 1280 µm long. Ostium bursae large; antrum low, cup-shaped with a slender pair of lateral lobes; lamella antevaginalis 210 µm, weakly sclerotized, trapezoid in ventral view, slightly inflexed on the posterior margin, widening toward anterior margin of A7. Ductus bursae 400–450 µm; terminus of ductus bursae biforked. Corpus bursae 690–700 µm, oblong with paired rows of wrinkles running longitudinally, some sclerotized. Distribution. Kauai. Host plants. Asteraceae: Dubautia sp. (D. laxa or D. paleata). Several species of Dubautia co-occur in the Alakai swamp area and it is unclear whether the specimens found by N. Tangalin were on Dubautia laxa or D. paleata. These mines were very abundant during our surveys in the summer of 2015. Biology. The larva mines fresh fuzzy leaves. Parasitoids. Braconidae sp.? (Fig. 105A–L)., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 68-71, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Johns, C. A., Toussaint, E. F. A., Breinholt, J. W. & Kawahara, A. Y. (2018) Origin and macroevolution of micro-moths on sunken Hawaiian Islands. Proceedings of the Royal Society B, 285, 20181047. https: // doi. org / 10.1098 / rspb. 2018.1047","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718."]}

Philodoria nigrella Walsingham, 1907 Figs. 13D, E. Philodoria nigrella Walsingham, 1907: 721, pl. 25, fig. 25); Zimmerman 1978a: 690, figs. 444, 451. Type locality. Mt. Kilauea, Hilo (Big Island). Type material. Lectotype ♂, Mt. Kilauea, 2000ft, Hilo, Hawaii (Big Island), xii.1895, Perkins 28604, | B.M. ♂ Genitalia slide no. 4173| PHILODORIA NIGRELLA, ♂ Wlsm. Fn. Hawaii I. TYPE descr. figd.| Walsingham Collection, 1910—427| NHMUK010305337| in NHMUK (here designated). Paralectotype 1 ♂, Same locality, vii.1895, Perkins 27401| Philodoria nigrella, ♂ Wlsm. PARATYPE 1/1| Walsingham Collection, 1910—427| NHMUK010305338| in NHMUK. Described from two specimens, and Walsingham likely considered them as holotype and paratype, as indicated on their specimen labels. However, because a holotype was not specified in the description, the so-labeled types are syntypes under Article 73.2 of the Code (ICZN 1999), and any one of them is can be designated as lectotype under Article 74 of the Code (ICZN 1999). The syntype ‘type (28604)’, which Walsingham listed and figured, is here designated as lectotype (Fig. 13D) and the other syntype is the paralectotype. Diagnosis. Blackish forewing with three white dorsal streaks and an outwardly white costal streak at 3/4. The forewing pattern is similar to P. touchardiella, but the latter has brownish forewing ground color and rather broad valva base. Description: Adult (Fig. 13D, E). The following is from Walsingham (1907: 721): “Antennae fuscous, white at the apex. Palpi white, the median joint streaked with fuscous externally, the terrninal joint fuscous beneath. Head fuscous; face yellowish white. Thorax blackish. Forewings black, with a slight brownish gloss, a white spot at the extreme base below the middle and three short, outwardly oblique, white dorsal streaks, one near the base reaching to the fold, the second before the middle, crossing the fold, the third, shorter, at about the end of the fold; a little beyond the third dorsal is an oblique, narrow, spatulate leaden gray costal streak, which is succeeded by three white streaks in the costal cilia before the apex; at the apex is a black spot, separated beyond it by leaden gray and below it by chestnut-brown, from a black curved line around the base of the leaden gray cilia which blend with tawny fuscous about the tornus. Exp. al. 9 mm. Hindwing blackish; cilia tawny fuscous. Abdomen blackish, white beneath. Legs blackish, wllitish beneath; hind tarsi spotted with whitish.” Male genitalia (Zimmerman 1978a: fig. 451). Tegumen oblong, 0.8 x length of valva; valva oblong, dorsal margin indistinct and genitalia mounted in a poor condition. Saccus slender. Phallus straight in lateral view; cornuti in vesica indistinct. Female genitalia Unknown. Distribution. Hawaii (Big Island) (Walsingham 1907). Host plants and Biology. Unknown. Remarks. This species has not been collected since the original two male specimens in 1895 and we believe it may be extinct., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on page 61, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Walsingham, Thomas, de Grey (Lord) (1907) Microlepidoptera. In: Sharp, D. (Ed.), Fauna Hawaiiensis. 1 (5). Cambridge University Press, Cambridge, pp. 469 - 759.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","ICZN (1999) International code of zoological nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 306 pp."]}

Philodoria hibiscella (Swezey, 1913) Figs. 16A, B, 27C–E, 47G, 48G, 49G, 58H, 77B–D, 93A, B, D, E. Graclaria hibiscella Swezey, 1913: 279–280 (first mentioned as “ Gracilaria hibiscella Sw. ” in Swezey (1913b: 224) without any description). Parectopa hibiscella Swezey, 1913; Swezey 1928: 191. Philodoria hibiscella (Swezey, 1913); Zimmerman 1978a: 680–681, figs. 437, 440, 442, 448, 455. Type locality. Mt. Tantalus (Oahu). Type material. Lectotype ♀, Mt. Tantalus, Oahu, x.1911, Coll. O.H.S., “from mines in leaes (=leaves) of native Hibiscus ”, Type no. 211, bottom one of two cotypes on the type mount in BPBM. Paralectotype ♂, top one of two cotypes on same mount of lectotype in BPBM. Described from three specimens collected from type locality from mines in leaves of Hibiscus. We found the lectotype and a paralectotype on same type mount designated by Zimmerman (1978a) but could not find the second paralectotype (sex unknown) (Fig. 16A, B). Additional material. 13 (5♂, 3♀ 5 sex unknown): Oahu: 1♂, 2 (sex unknown), Honolulu, 1.iii.1914 em., Coll. O.H.S., ex Hibiscus, 24.ii.1914 (Cocoon), SK684♂ BPBM 34179, BPBM 34180, BPBM 34183; 1♀, Manoa, 26.v.1932, O.H. Swezey Collector, ex leaf of Hibiscus, BPBM 34181; 1♂, Waiahole, 13.viii.1933, O.H. Swezey Collector, Hibiscus, genitalia slide no. Z-1-26-61(2), BPBM 34176; 3♂, 2♀, 3 (sex unknown), Tantalus, Coll. O.H.S., ex native Hibiscus, BPBM 34177, SK 685♀ | BPBM 34178, ♀ | BPBM 34182, J.F.G.C. #3753♂, 3754 ♀ and 1♂, 2 (sex unknown) in USNM. Molokai: 1♀, Waiehu Cliffs, 14.iii.2015 (stored), K. Bustamente leg., host: H. arnottianus subsp. immaculatus, 14.i.2012, 20141201KMB01, CJ399, SK 723♀ in BPBM. Maui: 1♂, 1♀, Iao Valley, 29.iv.2013 (stored), C.A. Johns leg., host: Hibiscus tiliaceus, 14.iv.2013, CJ088 / SK 725♀, CJ089 / SK724♂ in BPBM. Diagnosis. Ocherous forewing with four outwardly oblique white streaks: slender ds 1–3 and one slender cs 3 (Fig. 16A, B). Among Philodoria species having similar ds 1–3 and cs 3 , i.e. P. naenaeiella (Swezey), P. hesperomanniella sp. n., P. neraudicola (Swezey), P. obamaorum sp. n. and P. urerana (Swezey), P. hibiscella is most similar to P. urerana (feeding on Urera), but is distinguishable by the ocherous forewing; males have a rather broad saccus and slender and sinuous phallobase; females signa as two short rows of minute spines in the central area of the corpus bursae. Redescription: Adult (Fig. 16A, B). Wingspan 9–10 mm in type series; forewing length 4 mm in paralectotype, 3.8–4.5 mm in the additional material examined. Head pale brownish-ocherous; frons white; maxillary palpus brown with white scales; labial palpus pale ocherous-white, terminal segment somewhat fuscous on outer side. Antenna pale ocherous barred with dark fuscous, 1.3–1.4x length of forewing. Thorax pale brownish-ocherous. Forewing ocherous, outwardly-oblique ds 1–3 , widened and black-margined at dorsum; A slender outwardly-oblique narrowly black-margined cs 3 at costal 3/4; 3 or 4 black costal line on apical cilia; a black wide subterminal streak with a few bluish scales; cilia at apex blacky on termen gray with a black line at base. Hindwing and cilia grayishfuscous. Abdomen gray. Legs pale ocherous, anterior legs fuscous on outer side. Male genitalia (Figs. 47G, 48G, 49G) (n=3). Capsule 920 µm. Tegumen 0.9–1.0 x length of valva; valva 630 µm long, similar to P. limahuliensis except basal portion of valva humped along dorsal margin (Fig. 47G). Saccus slender and digitiform in ventral view (Fig. 48G). Phallus 580 µm long, slender, straight and sinuous at phallobase with developed coecum; cornuti in vesica indistinct (Fig. 49G). Female genitalia (Fig. 58H) (n=4). 1410 µm long. Ostium bursae large; antrum wide tubular with a slender pair of lateral lobes; lamella antevaginalis 220 µm, weakly sclerotized, trapezoid in ventral view, slightly inflexed on the posterior margin, widening toward anterior margin of A7. Ductus bursae slender, 400–450 µm; terminus of ductus bursae biforked. Corpus bursae 840 µm, pyriformwith paired rows of wrinkles running longitudinally, some sclerotized; two short linear signa with minute spines. Distribution. Oahu (Swezey 1913c), Maui, Molokai: new record, Hawaii (Big Island) (Zimmerman 1978a: 680). Host plants. Malvaceae: Hibiscus arnottianus A. Gray, H. rosasinensis L. (Swezey 1928), Hibiscus sp. (Swezey 1913b: 224, 1913c: 280), H. arnottianus subsp. immaculatus and H. tiliaceus L.: new record, endangered subspecies. Biology. (Fig. 93A, B, D, E). Swezey (1913c: 280) reported its biology: “The mine usually begins towards base of leaf, proceeding upward irregularly and following the margin for a part of its course, it eventually reaches the apex, then follows down the opposite margin of the leaf rapidly widening until the larya, has finished its growth. It then breaks through the epidermic to form its white oval cocoon on the surface of the leaf. Pupal stage about a week.” Zimmerman (1978a) noted that larvae occasionaly cause severe local damage to ornamental Hibiscus, and extensively mined leaves may drop from the plants and the white, oval cocoon is made on the surface of a leaf. We also observed larvae forming first a slender mine along the leaf margin on the adaxial surface that elongated into a blotch mine (Fig. 93B) and expanded as the larva grew (Fig. 93A, B). The frass is brownish green and is deposited in a spiral line. One mine per leaf. Leaves with old mines remained on the host plant and we did not find any indication that larvae cause mined leaves to fall to the ground as in other Philodoria species. The late instar larva is pale yellow to green (Fig. 93D) and pupation occurs in the white cocoon outside of the mine on the leaf adaxial surface (Fig. 93E). Parasitoids. Eulophidae: Diglyphus sp., Euderus metallicus (Ashmead, 1901), Neochrysocharis formosus (Westwood, 1833), Pauahiana maculatipennis (Ashmead, 1901), Pnigalio externa (Timberlake, 1927) Sympiesis vagans (Timberlake, 1926) (Zimmerman 1978a). Remarks. The mature larva and pupa described by Swezey (1913c: 280): “ Mature larva. The full-grown larva is about 9 mm.; pale bluish-green; head with blackish mouth-parts, eyes, and 2 lines bordering the paraclypeus, much retracted into segment 2 which is widened, and has a large black spot ventrally and 2 black spots near the anterior margin dorsally; thoracic legs minute; abdominal prolegs on segments 7–9. Pupa. 5 mm.; pale testaceous-greenish, with a few fuscous markings ventrally; wing-sheaths extend to apex of fifth abdominal segment, free beyond fourth segment, dark fuscous at tip; posterior leg-sheaths extend to apex of abdomen; antenna sheaths extend beyond apex of abdomen, recurved over the back forward to base of fourth abdominal segment.”, Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 72-73, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Swezey, O. H. (1913 b) Leaf-miners of the Hawaiian Islands. Proceedings of the Hawaiian Entomological Society, 2 (5), 221 - 227.","Swezey, O. H. (1928) Some new species of lepidopterous leaf-miners in Hawaii. Proceedings of the Hawaiian Entomological Society, 7 (1), 187 - 191.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","Swezey, O. H. (1913 c) One new genus and eighteen new species of Hawaiian moths. Proceedings of the Hawaiian Entomological Society, 2 (5), 269 - 280."]}

Philodoria napaliensis Kobayashi, Johns & Kawahara, sp. n. Figs. 9D, 41M, N, 42L, 43L. Philodoria sp. 4; Johns et al. 2018: fig. 2. Type locality. Nâ Pali Trail (Kauai). Etymology. The specific epithet, napaliensis is derived from the type locality, Nâ Pali (pronounced 'Nah-Pahlee') Coast. Type examined. Holotype (sex unknown), Na Pali Trail, Kauai, 13& 21.vii.2015 (stored), C.A. Johns leg., host: Diospyros sandwicensis, 24.vi.2015, CJ491a (abdomen missing) in BPBM. Paratype 2♂, same locality and data as holotype, CJ482 / SK767♂, CJ491b / SK745♂ (apex of tegumen broken); deposited in BPBM. Three specimens are incomplete but they have features that are distinctive enough to allow this species description. What remains of three specimens were: holotype mounted by placing three wings without mountant under a coverslip: two full forewings slightly damaged at base and small apical region of one hindwing; two paratypes: a specimen mounted by placing four wings without mountant under a coverslip (two forewings [1/3 of left wing and 1/2 right wing] and 1/3 of two hindwing; CJ482); a specimen mounted by placing two forewings and one leg without mountant under a coverslip (almost full left wing and apical portion and basal half of right wing and one hind leg; CJ491b). The head, antenna, thorax, and legs for all specimens were sacrificed for molecular analysis. Additional material. 2 larvae (CJ-445/AHE_44) entirely sacrificed for molecular analysis (Johns et al. 2018: fig. 2): 2 larvae, same locality and data as holotype, 26.vi.2015 (stored), CJ445, in FLMNH. Diagnosis. Very similar to Philodoria that feed on Myrtaceae and P. lama sp. n., but differs from them in having a valva with a dorsal process with a thick inner margin (Fig. 41M, N). Description: Adult (Fig. 9D). Forewing length 3.4–3.5 mm in holotype. Head, antenna and thorax unknown. Forewing dark shiny, pale leaden gray with brownish ocherous patches enclosing gray fascia: bp from base to dorsum 1/3, contact to basal tf; tf from costal 1/3 to dorsal 1/2, tp after the middle to apical portion, distinctly narrowing in the dorsum, extending to dorsal 2/3, containing two short, shiny, steel gray costal spots; apical portion with black as and a large jet-black terminal patch from tornus to apex, containing two upright steel-gray spots near its opposite extremities; a single bright white spot in its middle; a very narrow ocherous line along the base of tornus; cilia shining, leaden gray with a blackish fringe line; two white costal streaks, long one extend apex; terminal cilia often blackish. Hindwing coppery brown; cilia tawny. Abdomen tawny brown, white below. Legs unknown. Male genitalia (Figs. 41M, N, 42L, 43L) (n=2). Capsule 600 µm. Tegumen 0.7–0.8 x length of valva; valva 450–460 µm long, similar to P. basalis and P. lama except inner margin of dorsal process thicken (Fig. 41M, N). Phallus 550 µm long (Fig. 43L). Female genitalia Unknown. Distribution. Kauai. Host plants. Ebanaceae: D. hillebrandii (A.DC.) Fosberg and/or Diospyros sandwicensis (A.DC.) Fosberg. We could not distinguish the two Diospyros species, D. hillebrandii and the closely related D. sandwicensis. Diospyros sandwicensis is found on all of the main Hawaiian islands. It often occurs as a dominant plant in dry to moist forests and occasionally in wet forests. It grows at elevations ranging from close to sea level to 1200 m (4000ft) (Wagner 1990). Biology. Unknown, but it is presumably similar to those of P. basalis, P. lama and P. splendida, whose larvae pupate in a “taco” like cocoon., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 42-43, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Johns, C. A., Toussaint, E. F. A., Breinholt, J. W. & Kawahara, A. Y. (2018) Origin and macroevolution of micro-moths on sunken Hawaiian Islands. Proceedings of the Royal Society B, 285, 20181047. https: // doi. org / 10.1098 / rspb. 2018.1047","Wagner, W. L., Herbst, D. R., & Sohmer, S. H. (1990) Manual of the flowering plants of Hawai'i. 2 Vols. Bishop Museum Special Publication 83. University of Hawaii Press and Bishop Museum Press, Honolulu, pp. 585 - 587."]}

Philodoria haelaauensis Kobayashi, Johns & Kawahara, sp. n. Figs. 11E, F, 26K, L, 39C, 44E–H, 45C, D, 46C, D, 57F, 63H, 64A–D, 74, 84. Philodoria pipturicola Swezey, 1915; Zimmerman 1978a: 709, figs. 471, 476, 479 (Misidentification). Philodoria sp. 10; Johns et al. 2018: fig. 2. Type locality. Haelaau (Maui). Etymology. The specific epithet is derived from the type locality, Haela`au (pronounced ‘Haye-La-ahoo’). Type material. (6♂, 1♀, 7 sex unknown). Holotype ♂, Haelaau, Maui Is., 18.xii.1928, O.H. Swezey Collector, Pipturus, SK 718, BPBM34265. Paratype 19 (6♂, 1♀, 12 sex unknown): 1♂, 7 sex unknown, same data and locality as holotype, BPBM34260, 34261, | Philodoria pipturicola Det. By O.H. Swezey | BPBM34262, BPBM34263, BPBM34264, | 1 ♂ Same date ex to Tribert, 1993| BPBM34265, BPBM34266, BPBM34267. 1♂, 1♀, 2 sex unknown, Wailuku, 2000ft, Maui, 9.xii.1922, O.H. Swezey, Pipturus, Z-XII-19-62-23♂, Z-XII-19-62- 24♀, BPBM34253; 4♂, 5 sex unknown (missing abdomen), Same data and locality, ‘♂ + ♀ genitalia on slide 99, aB Oct 15 1940 ’, SK822♂(middle), SK827♂(top middle), SK828♂(top right), BPBM34254. Additional material. 14 (5 ♂, 6 ♀, 3 sex unknown). Molokai: 1♀, Kamiloloa, 19.xii.1925, 3200ft, O.H. Swezey Collector, Pipturus, | Philodoria floscula Det by O.H. Swezey|, SK 705♀, BPBM 34170; 1♂, Kaibalu, 2000– 3000ft, 28.vii.1928, O.H. Swezey Collector, on Pipturus, | Philodoria floscula Det by O.H. Swezey|, SK704♂, BPBM 34171; 1♂, 1♀, 3 sex unknown, Kamiloloa, Mol., 19.xii.1925, 3200ft, O.H. Swezey Collector, Pipturus, BPBM 34172–34176. Maui: 1♂ 2♀, Makawao Forest Reserve, 11& 15.i.2014 (stored), W. Haines leg., host: Pipturus rockii, 28.xii.2013, CJ209 / PHIL0016 ♀, CJ216 / SK 821♀, CJ227 / PHIL0019♂ (Fig. 74) in BPBM. 2♂, 2♀, Honokohau, 9.viii.2014 (stored), K. Bustamente leg., host: Pipturus albidus, Summer. 2014., CJ348b / SK789♂, CJ348c / SK790♂, CJ348d / SK 791♀, CJ348e / SK 792♀ in BPBM. Diagnosis. The forewing pattern is similar to P. pipturicola Swezey but distinguished by an oblique white patch in the middle of the forewing, that reaches the dorsal margin (Fig. 11E, F); male valva same width at apical half; female with spiny pyriform signa and lacking sclerotized structure at the joint of the ductus bursae and corpus bursae. The new species and P. alakaiensis both have two pairs of coremata in the male 8th segment (Fig. 39C, D). Swezey (1923) misidentified specimens of P. haelaauensis from Wailuku, Maui as P. pipturicola and Zimmerman (1978a: figs. 471, 476, 479) described in error that the genitalia of P. haelaauensis as those of P. pipturicola. Description: Adult (Figs. 11E, F, 26K, L). Wingspan 7.1 mm in holotype, 6.3–7.2 mm in paratypes; forewing length 3.3, 3.4 mm in holotype, 2.8–3.4 mm in paratypes. Head slate grey fuscous; frons white; maxillary palpus reduced; labial palpus white, median joint slightly fuscous externally. Antenna dark fuscous. Thorax slate grey fuscous. Forewing fuscous suffused with orange patches at apical portion: an obscure cl from near base to about 2/3, apical potion bends inward; two white bands (w 1, 2 ) on both extremities of pale blue color band (bb 3 ) at 3/4, widest in the the middle; a large oblique ws near middle of wing, reaching dorsal margin; a large pale-bluish patch (as) at apex; cilia fuscous with two white spots (a, b) in costal cilia; Hindwing and cilia dark fuscous. Abdomen dark fuscous, white beneath. Legs fuscous, white beneath. Male genitalia (Figs. 44E–H, 45C, D, 46C, D, 64A, B) (n=5). Capsule about 750–800 µm. Tegumen 0.8–0.9 x length of valva; valva 530, 550 µm long, similar to pipturicola except apical half of valva not tapering at apex, almost same width in lateral view (Fig. 44E–H). Saccus triangular (Fig. 45C). Phallus 310 µm, straight in lateral view with two series of minute cornuti in vesica (Fig. 46C, D). Female genitalia (Figs. 57F, 63H) (n=3). 1250–1300 µm long. Ostium bursae large; antrum wide with a slender pair of lateral lobes; lamella antevaginalis 220–230 µm, weakly sclerotized, widening toward anterior margin of A7. Ductus bursae 600–650 µm, posterior extremity widening and membranous, and median section narrowing weakly sclerotized and flattened; terminus of ductus bursae membranous. Corpus bursae pyriform, 540–550 µm; signum a conical plate with minutely spined surface. Distribution. Maui (Swezey 1923 a) and Molokai: new record. Host plants. Urticaceae: Pipturus sp. (Swezey 1923 a), P. albidus A.Gray ex H.Mann and P. rockii Skottsb.: new record. Biology. (Fig. 84). Larvae form a tortuous linear to blotch mine (Fig. 84A–D) that expands as they feed and grow (Fig. 84E, G); the frass line is black. 1–5 mines per leaf. Pupation occurs in the cocoon outside of the mine, on the abaxial leaf surface; cocoon orangish brown (Fig. 84F)., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 54-55, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Swezey, O. H. (1915) New species of Hawaiian moths. P roceedings of the Hawaiian Entomological Society, 3 (2), 94 - 97. https: // doi. org / 10.5962 / bhl. part. 24594","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","Johns, C. A., Toussaint, E. F. A., Breinholt, J. W. & Kawahara, A. Y. (2018) Origin and macroevolution of micro-moths on sunken Hawaiian Islands. Proceedings of the Royal Society B, 285, 20181047. https: // doi. org / 10.1098 / rspb. 2018.1047","Swezey, O. H. (1923) The leaf-miners of Pipturus. Proceedings of the Hawaiian Entomological Society, 5 (2), 293 - 296."]}

Philodoria pittosporella (Swezey, 1928) Fig. 21D. Parectopa pittosporella Swezey, 1928: 189–190. Philodoria pittosporella (Swezey, 1928); Zimmerman 1978a: 690, figs. 445, 452. Type locality. Mt. Kaala, above 3000ft (Oahu). Type material. Holotype ♂, Mt. Kaala, Waianae range, [above 3000 ft.], Oahu, 1.iii.1925, Swezey Coll., ex cocoon on leaf of Pittosporum, | Holotype | Parectopa pittosporella Swezey Det. by O.H. Swezey | ♂ holotype abdomen sent E.C.Z 11-8-60,| [Type] no. 4264 in BPBM. Described based on a single specimen from Oahu. Diagnosis. The forewing pattern is similar to that of P. funkae sp. n., but it is distinguished by its light ocherous brown forewing and orange to light brown apex (Fig. 21D). See also the diagnosis of P. keahii sp. n. Redescription: Adult (Fig. 21D). Wingspan 8.5 mm in holotype; forewing length 4 mm in holotype. Head and frons white; maxillary palpus white; labial palpus white, a little brownish externally. Antenna light ocherous brown, whitie at base, about 1.2–1.3x length of forewing. Thorax white. Forewing light ocherous brown with white patches: a large white ds 2 at 1/2, connecting ds 1 at 1/5, distad portion widened to fold, then with an oblique pointed extension continuing 2/3 across wing, another from 3/5 nearly to apex, an extension obliquely towards the end of white costal streak, not reaching it, with a few brown scales at 3/4; an outwardly oblique white cs 3 at 4/5, margined with fuscous scales; a few blue scales at apex; apical portion damaged, cilia gray with three white costal spots (a, b, c). Hindwing and cilia grayish fuscous. Legs grayish fuscous, somewhat banded with white. Abdomen grayish fuscous. Male genitalia See Zimmerman (1978a: fig. 452). Tegumen oblong, 0.7–0.8 x length of valva; valva tapering along costal margin from basal 1/2 to apex; apical half slender and slightly curved toward inner margin. Saccus short and pointed at apex in ventral view. Phallus straight with small coecum; vesica with well-developed minute cornuti. Female genitalia Unknown. Distribution. Oahu (Swezey 1928). Host plants. and Biology. Unknown. Remarks. Zimmerman (1978a) incorrectly reported Pittosporum as the host plant and noted that the larva emerges from its mine to spin a cocoon on the leaf surface. Swezey (1928) noted that the holotype was reared from a cocoon found on a leaf of Pittosporum without mines on the leaves and pointed that Pittosporum was not the actual host plant. We observed leaf mines on Pittosporum on Kauai and Oahu, but it is unclear what insect makes these leaf mines as we have not been successful in finding a mine with a larva. It is likely that this mine is not made by Philodoria, but by another insect., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 97-98, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Swezey, O. H. (1928) Some new species of lepidopterous leaf-miners in Hawaii. Proceedings of the Hawaiian Entomological Society, 7 (1), 187 - 191.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718."]}

Philodoria dubauticola (Swezey, 1940) Figs. 14A, 27A, 34E, 47C, 48C, 49C, 58C, D, 88B–F. Parectopa dubauticola Swezey, 1940: 463–464. Philodoria dubauticola Swezey, 1940; Zimmerman 1978a: 667, figs. 441, 447; Johns et al. (2016): 63, 64, fig. 1. Type locality. Ridge above Haelaau (Maui). Type material. Lectotype ♀, Ridge above Haelaau, Maui, 20.xii.1928, O.H. Swezey Collector, Dubautia, | Holotype Parectopa dubauticola Swezey | Type no. 4258 in BPBM (here designated). Described from three specimens reared from Dubautia leaf mines collected on the ridge above Haelaau, Maui. Swezey considered them as holotype and paratypes as indicated on their specimen labels. However, because a holotype was not specified in the original description, the so-labeled types can all be considered syntypes under Article 73.2 of the Code (ICZN 1999), and any of them is eligible for designation as lectotype under Article 74 of the Code (ICZN 1999). The syntype ‘Holotype’ is here designated as lectotype (Fig. 14A) and the remaining syntypes are paralectotypes, but we could not find them in BPBM. Additional material. 7 (3♂, 3♀, 1 sex unknown): 2♂, 3♀, 1 (sex unknown), Haelaau, Maui, 29.iv.& 7.v.2013 (stored), C.A. Johns leg., host: Dubautia laxa, 10.iv.2013, CJ077 / KT982408 (see Remarks), CJ083 / PHIL0023♂, CJ084, CJ094 / SK 645♀, CJ111 / PHIL0008 ♀, CJ256 / SK646♂ in BPBM; 1♂, Helu, Maui, 14.iii.2015 (stored), K. Bustamente leg., host: D. laxa, 18.xii.2014, KMB01, CJ402, SK812♂ in BPBM. Philodoria sp. nr P. dubauticola: 3 (sex unknown), Nakula Natural Reserve, Maui, 1.viii.2014 (stored in RNAlater solution), C.A. Johns leg., host: Dubautia platyphylla, 26.vii.2014, CJ 315 in FLMNH (Fig. 75A). Diagnosis. Brownish fuscous forewing with a white longitudinal streak on the basal half of the wing. The species is similar in appearance to P. dubautiella (Swezey), but distinguishable from the latter by having two dorsal white streaks instead of three on the forewing, a rather broad saccus, and a large and well-sclerotized terminus of the ductus bursae. Redescription: Adult (Fig. 14A, 34E). Wingspan 8 mm; forewing length 4 mm in holotype. Head dirty white; frons white; maxillary palpus white with few dark scales externally; labial palpus whitish, second segment broadly infuscated externally at apex, third segment with a few dark scales externally at apex and towards base (Fig. 34E). Antenna light fuscous, basal fourth white beneath, about 1.3–1.4x length of forewing. Thorax grayish brown, white beneath. Forewing brownish fuscous with white streaks margined with a few fuscous scales: a white longitudinal streak from base to about 2/5; two dorsal outwardly oblique ones (ds 2, 3 ) extending the middle of wing, the ds 2 at 2/5, the ds 3 at 3/5; white narrow costal line on basal 1/3; two outwardly oblique white costal streaks (cs 2, 3 ), cs 2 at 1/2 and wider at costal side (not always distinct), the cs 3 at 3/4, slender and of even width terminating in a patch of pale blue scales, a similar patch of blue scales at apex of wing (as); remainder of apical portion of wing suffused with ocherous; costal cilia with three white costal spots (a, b, c) before apex, separated by light fuscous scales. Cilia light fuscous with a distinct dark fuscous fascia at base. Hindwing uniform light fuscous, cilia paler. Legs light fuscous, tarsi banded with white. Abdomen grayish fuscous, white beneath. Male genitalia (Figs. 47C, 48C, 49C) (n=3). Capsule 770 µm. Tegumen 0.8–0.9 x length of valva; valva tapering along costal margin from base 2/5 to apex; along the inner side of valva, a series of small spines arranged; apical half slender (Fig. 47C). Saccus short and thick (Fig. 48C). Phallus 500 µm long, slender and sinuous in lateral view with developed coecum; cornuti in vesica indistinct (Fig. 49C). Female genitalia (Fig. 58C, D) (new record) (n=3). 1120 µm long. Ostium bursae large; antrum cup-shaped with a slender pair of lateral lobes; lamella antevaginalis 280 µm, large and weakly sclerotized, trapezoid in ventral view, slightly inflexed on the posterior margin, widening toward anterior margin of A7. Ductus bursae 400 µm; terminus of ductus bursae biforked. Corpus bursae 600 µm, pyriform; paired rows of wrinkles running longitudinally, some sclerotized. Distribution. Maui (Swezey 1934). Host plants. Asteraceae: Dubautia plantaginea Gaudich.? (Zimmerman 1978a) and Dubautia laxa Hook. & Arn. (Johns et al. 2016). Biology. (Fig. 88B–F). Larvae form a slender linear mine on the adaxial leaf surface (Fig. 88B–D), that gradually expands as larvae feed and grow (Fig. 88E). One mine per leaf. Pupation occurs in the cocoon outside of the mine, on the adaxial surface of leaf (Fig. 88F). Remarks. We identified one adult moth (Coll ID CJ-077 / GenBank accession no. ID KT982408) as P. dubauticola, based on the presence of a white longitudinal streak on forewing, and this specimen was sacrificed for molecular analysis (Johns et al. 2016)., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 63-64, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Swezey, O. H. (1940) New species of Hawaiian Lepidoptera. Proceeding Hawaiian Entomological Society, 10 (3), 461 - 465.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","Johns, C. A., Moore, M. R. & Kawahara, A. Y. (2016) Molecular phylogeny, revised higher classification, and implications for conservation of endangered Hawaiian leaf-mining moths (Lepidoptera: Gracillariidae: Philodoria). Pacific Science, 70 (3), 361 - 372. https: // doi. org / 10.2984 / 70.3.7","ICZN (1999) International code of zoological nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 306 pp.","Swezey, O. H. (1934) New species of Hawaiian Lepidoptera. Proceedings of the Hawaiian Entomological Society, 8 (3), 523 - 525."]}

Philodoria molokaiensis Swezey, 1928 Figs. 8B, 26F, 41H, 42G, 43G, 56I, J, 62F–H, 73A, 80A, B, D, E. Philodoria molokaiensis Swezey, 1928: 188–189; Zimmerman 1978a: 709, figs. 464, 470, 478. Type locality. Kawela (Molokai). Type material. Lectotype ♂, Kawela, 3700ft, Molokai, 23.xii.1925, O.H. Swezey Collector, Lysimachia [hillebrandi var.], | Holotype | Philodoria molokaiensis Swezey Det. by O. H. Swezey| ♀ paratype & ♂ holotype abdomen sent ECZ 7?-17-61| Type no. 4271, slide no. Z-XII- 20-65 in BPBM (here designated). Paralectotype ♀, same data and locality as lectotype, Z-XII-19-62- 26 in BPBM. Described from two specimens. Swezey considered them as holotype and paratype, as indicated on their specimen labels. However, because the holotype was not specified in the original description, the so-labeled holotype and paratype are all to be considered syntypes under Article 73.2 of the Code (ICZN 1999), and any one is thus eligible for designation as lectotype under Article 74 of the Code (ICZN 1999). The syntype labeled ‘Holotype’ is here designated as lectotype (Fig. 8B), and the other syntype is the paralectotype. Addtional material. (2♂, 2♀). Molokai: 2♀, Kamakou, 16–20.i.2014 (stored), C.A. Johns leg., host: L. hillebrandi var., CJ229 / SK 668♀ (20.xii.2013 em., wing and abdomen), CJ239 / SK 669♀ (abdomen); 2♂, Kawela, 26.vii.2015 (stored), K. Bustamente leg., host: L. maxima, KMB 01, 20150317, CJ509 / SK667♂ (wing and abdomen), SK666♂ (abdomen); deposited in BPBM. Diagnosis. Forewing with two longitudinal streaks of ocherous-orange on distal area of thorax and the patagium is light gray instead of ocherous-orange as in P. lysimachiella Swezey (Swezey 1928) (Fig. 8B, 73A). See also diagnosis of P. lysimachiella. Redescription: Adult (Figs. 8B, 73A). Wingspan 9 mm in lectotype; forewing length 3.2 mm in paralectotype. Head light gray; frons white; maxillary palpus reduced; labial palpus white, slightly brownish externally on the base. Antenna light gray, white below on basal third, about 1.3x length of forewing. Thorax ocherous-orange on each side of disk, patagium light gray. Forewing shiny metallic bronze with ocherous-orange patches: a large oblique patch from base to 1/3, containing two longitudinal streaks; a median white one narrowing toward apex, another same length, on the dorsal margin; tf from costal 1/3 to dorsal 1/2; a large tp after the middle to costal 3/4, distinctly narrowing in the dorsum, extending to dorsal 2/3, containing white cs; one white band (w 1 ) on the middle of first bronze color band (bb 1 ) from the base via costal fold, curved at costal 1/4 to dorsal 1/2; others on costa to the middle of bb 2 and bb 3 ; apical portion with a fuscous patch extending toward the termen and apex with a bronze as with orange-ocherous color encroaching on the apex and a small white spot at tornus; cilia light gray, with a white spot at beginning of costal cilia, remainder of cilia with black scales at base. Hindwing light gray, slightly infuscated towards apex, cilia light gray. Accessible abdomen that we had in poor condition, pale brownish gray, preserved in 99% ethanol. Legs mostly gray, distal segment white. Male genitalia (Figs. 41H, 42G, 43G) (n=2). Capsule 970 µm. Tegumen 570 µm long and 0.9 x length of valva; valva 640–650 µm long, very broad and long apically extended and having a slender and very long dorsal process (Fig. 41H). Saccus short and thick, apically blunt (Fig. 42G). Phallus 710 µm long and nearly straight and gradually wider in phallobase (Fig. 43G). Female genitalia (Figs. 56I, J, 62F) (n=2). 1260 µm long. Ostium bursae large; antrum rather short; lamella antevaginalis 340 µm, trapezoid, large inflexed on the posterior margin. Corpus bursae 630 µm long; paired signa with a pair of large triangular-shaped spines. Distribution. Molokai (Swezey 1928). Host plants. Primulaceae: Lysimachia hillebrandii var. (Swezey 1928) and L. maxima (R. Knuth) H. St. John: new record. L. maxima is very rare plant, only two small populations remain on Molokai, listed as an endangered plant species in the United States. Biology. (Fig. 80A, B, D, E). Larvae form a slender linear mine on the adaxial leaf surface, and mines gradually expand as the larvae feed and grow (Fig. 80A, B). One mine per leaf. The late instar larva is pale yellow-green (Fig. 80D). Pupation occurs in the cocoon outside of the mine, usually on the abaxial leaf surface; a pupal cocoon is pale grayish ocherous and near ellipsoid (Fig. 80E). Parasitoids. Chalcidoidea spp. (including at least three species from CJ201), Euplemidae sp.: new record (Fig. 99E–L)., Published as part of Kobayashi, Shigeki, Johns, Chris A. & Kawahara, Akito Y., 2021, Revision of the Hawaiian endemic leaf-mining moth genus Philodoria Walsingham (Lepidoptera: Gracillariidae): its conservation status, host plants and descriptions of thirteen new species, pp. 1-175 in Zootaxa 4944 (1) on pages 34-36, DOI: 10.11646/zootaxa.4944.1.1, http://zenodo.org/record/4681813, {"references":["Swezey, O. H. (1928) Some new species of lepidopterous leaf-miners in Hawaii. Proceedings of the Hawaiian Entomological Society, 7 (1), 187 - 191.","Zimmerman, E. C. (1978 a) Gracillarioidea. In: Microlepidoptera. Part 1. Insects of Hawaii, Vol. 9. University of Hawaii Press, Honolulu, pp. 644 - 718.","ICZN (1999) International code of zoological nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 306 pp."]}

Kinematics of each individual used in the study.

Contains supplemental figures, tables, methods, results, and discussion

This file contains all supplemental text, figures, and tables.