Philodoria marginestrigata ( Walsingham, 1907 )

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, Zootaxa 4944 (1), pp. 1-175 : 101-103

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.

Figures

FIGURE 23. Adults of Philodoria species. A P. marginestrigata (Walsingham, 1907) paralectotype male (BPBM 34197), B P. marginestrigata (Walsingham, 1907) Lectotype female, left wings, C Same specimen, right wings, D P. lipochaetaella (Swezey, 1940) paralectotype male, one of four cotypes on the same mount (upper left specimen on mount), E Lectotype male (upper right specimen on mount), F Paralectotype female (lower left specimen on mount), G Paralectotype (sex unknown, lower right specimen on mount). Scale bar 1 mm.

FIGURE 27. Forewing pattern variation in Philodoria. Specimens from recent collection efforts that were preserved in ethanol or RNA stabilization reagent, originally for molecular studies. A P. dubauticola, male (CJ402/SK812), B P. dubautiella, male CJ524a/SK686), C P. hibiscella, female (CJ399/SK723), D Same species, male (CJ089/SK724), lateral view, E Same species, dorsal view, F Philodoria sp., male, host: Dubautia plantaginea, Maui (CJ352c/SK770), G Philodoria sp., female (CJ500), host: Dubautia plantaginea, Oahu, H P. naenaeiella (CJ188), abdomen missing, I P. neraudicola, female (CJ336/SK838), host: Pipturus albidus, J Same species, female (CJ369), host: Pipturus sp., K P. obamaorum, female (CJ423/SK844), L Same species, left forewing, M Same species, female (CJ403a/SK735), N P. touchardiella, female (CJ401a/SK764), O Same species, male (CJ401b/SK765), P P. marginestrigata, male (CJ165/SK760), host: Lipochaeta rockii, Q Same species, male (CJ187/SK771), host: Abutilon menziesii, R Same species, male (CJ379c), host: Sida fallax. Scale bar 1 mm.

FIGURE 37. Illustrations of the head of Philodoria. A P. basalis, female, Hawaii (Big Island) (SKH09-1, SK615), host: Metrosideros polymorpha, B P. lysimachiella, male, Oahu (CJ530a/SK749), host: Lysimachia hillebrandii, C P. marginestrigata (sex unknown), Molokai (CJ186), host: Abutilon grandiflorum, D P. hauicola, female, Oahu (SKH08-9/SK750), host: Hibiscus tiliaceus. ey: eye, lp: labial palpus, mp: maxillary palpus, sc: scape, pd: pedicel, pf: pilifer, pr: proboscis.

FIGURE 40. Genitalia of Philodoria. A–D: Male genitalia of P. marginestrigata, Kauai (CJ521/SK784), host: Sida fallax, E–H: Female genitalia of P. basalis, Hawaii (Big Island [SKH09-1]), host: Metrosideros polymorpha, E–G: SK617, H: SK616. A Phallus, lateral view, B Genitalia except right valva and phallus, ventral view, C Right valva, inside view, D Dorsal flap, inside view, E, G Female genitalia, lateral view, F, H Same, ventral view. aa: apophysis anterioris, an: antrum, ap: apophysis posterioris, cb: corpus bursae, co: coecum, cr: cornuti, db: ductus bursae, de: ductus ejaculatorius, ds: ductus seminalis, la: lamella antevaginalis, ob: ostium bursae, ph: phallus, phb: phallobase, sa: saccus, si: signum, ta: tuba analis, te: tegumen, va: valva, vi: vinculum.

FIGURE 50. Valva of Philodoria, inner view. A, C, E–J, L: Left valva, B, D, K, M: Right valva. A, B P. neraudicola, paralectotype, Oahu, SK707, C, D P. obamaorum, paratype, Hawaii (Big Island), SK709, E P. urerana, Oahu, paralectotype, SK671, F P. hauicola, Oahu, SK694, G P. keahii, holotype, Oahu, SK758, H P. touchardiella Maui, SK643, I P. platyphylliella, holotype, Maui, SK833, J, K P. marginestrigata, Oahu, SK662, host: unknown, L Same species, Molokai, SK771, host: Abutilon menziesii, M Same species, Kauai, SK784, host: Sida, fallax.

FIGURE 51. Genital capsule of Philodoria, excluding right portion of male genitalia, ventral view. A P. neraudicola, paralectotype, Oahu, SK707, B P. obamaorum, paratype, Hawaii (Big Island), SK709, C P. urerana, paralectotype, Oahu, SK671, D P. hauicola, Oahu, SK694, E P. keahii, holotype, Oahu, SK758, F P. touchardiella Maui, SK643, G P. platyphylliella, holotype, Maui, SK833, H P. marginestrigata, Oahu, SK662 host: unknown.

FIGURE 52. Phallus of Philodoria, lateral view.A P. neraudicola, paralectotype, Oahu, SK707, B P. obamaorum, paratype, Hawaii (Big Island), SK709, C P. urerana, paralectotype, Oahu, SK671, D P. hauicola, Oahu, SK695, E P. keahii, holotype, Oahu, SK758, F P. touchardiella Maui, SK643, G P. platyphylliella, holotype, Maui, SK833, H P. marginestrigata, Oahu, PHIL0029 host: Abutilon incanum.

FIGURE 60. Female genitalia of Philodoria. A P. marginestrigata, SK665, B Same species, SK772, host: Abutilon menziesi, C Same species, SK787, host: Sida fallax, D P. sciallactis, paralectotype, SK713, E P. ureraella, paralectotype, SK677, F P. opuhe, paratype, SK675.

FIGURE 77. Resting posture of adult Philodoria. A P. hauicola, Oahu, CJ270, host: Hibiscus tilaceus, B–D P. hibiscella, Oahu, CJ238, host: Hibisucs arnottianus, E P. marginestrigata, Molokai, CJ150, host: Lipochaeta rockii, F Same species, Molokai. CJ168/169, host: Sida fallax.

FIGURE 93. Mines of Philodoria species with thier Malvaceae host plants, Oahu. A, B, D, E: P. hibiscella, host: Hibiscus arnottianus, C P. marginestrigata, host: Sida fallax, F P. hauicola, host: Hibiscus tilaceus, D Mine of late instar larva, E Cocoon on the adaxial side of leaf, F Old mine.

FIGURE 108. Parasitoid wasps (Chalcidoidea) that emerged from Philodoria marginestrigata on Sida fallax, Kaena point, Oahu. A CJ137-1, B CJ137-2, C CJ137-3, D CJ137-4, E CJ137-5, F CJ137-6, G CJ137-7, H CJ137-8, I CJ137-9, J CJ137-10, K CJ137-11.

FIGURE 109. Parasitoid wasps that emerged from Philodoria marginestrigata on Sida fallax. A: Chalcidoidea, B–M: Eulophidae. A CJ055 Maui, B, C CJ056 Maui, D, E CJ193, Maui, F, G CJ205, Maui, H, I CJ378, Lanai, J, K CJ407-1, Hawaii (Big Island), L, M CJ407-2.