Philodoria basalis 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 : 36-38

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).

Figures

FIGURE 8. Adults of Philodoria species. A P. lysimachiella Swezey, 1928 lectotype male, B P. molokaiensis Swezey, 1928 lectotype male, C P. basalis Walsingham, 1907 lectotype male, D Paralectotype male, E Female, Hawaii (Big Island), F Male, Hawaii (Big Island). Scale bar 1 mm.

FIGURE 26. Forewing pattern variation in Philodoria. Specimens from recent collection efforts that were preserved in ethanol or RNA stabilization reagent, originally for molecular studies. Wings were placed without mountant under a coverslip. A P. succedanea (sex unknown, CJ419), B P. auromagnifica, male (CJ148/SK689), C P. basalis, female (CJ410), D P. splendida, female (CJ488a), host: Syzygium sp., E P. splendida (sex unknown, CJ488b), host: Syzygium sp., F P. molokaiensis, male (CJ509b/ SK667), G P. sp. A nr. P. lysimachiella, female (CJ346a/SK753), H P. lysimachiella, female (CJ530), lateral view, I P. pipturiella, female (CJ529), J Same species, male (CJ529), K P. haelaauensis, female (CJ209), L Same species, female (CJ216), M P. floscula, female (CJ426/SK845), N P. wilkesiella, male (CJ009/SK649), O P. epibathra, male (CJ506a/SK678), P Same species, female (CJ506b/SK681). Arrows point to diagnostic characters of species: black patch at base (A), basal fascia (D), transverse fasciae (tf) (F, G), white spot (ws) (I, J), white patch (K–M). Scale bar 1 mm.

FIGURE 34. Adult female head of five species of Philodoria, showing their maxillary palpus. A P. succedanea, B, C P. kolea, paratype, D P. basalis, E P. dubauticola, F P. hauicola. lp: labial palpus, pf: pilifer, pr: proboscis. Arrows point to the maxillary palpus. Scale bar 0.1 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 41. Male valva of Philodoria, inner view. A–F, H–J, L, M: Left valva, G, K, N: Right valva. A P. succedanea, Maui, SK641, B P. auromagnifica, Hawaii (Big Island), SK800, C P. auromagnifica, Kauai, SK689, D P. kolea, holotype, Hawaii (Big Island), SK851, E P. lysimachiella, Oahu, SK656, F, G P. sp. 7, Kauai, SK754, H P. molokaiensis, Molokai, SK666, I P. basalis, Hawaii (Big Island), SK614, J P. splendida, Maui, SK696, K Same species, Oahu, SK747, host Syzygium sp., L P. lama, holotype, Maui SK744, M P. napaliensis, paratype, Kauai, SK767, N Same as L, paratype, Kauai, SK745.

FIGURE 42. Genital capsule of Philodoria, excluding right portion of male genitalia, ventral view. A P. succedanea, Maui, SK641, B P. auromagnifica, Hawaii (Big Island), SK800, C P. auromagnifica, Kauai, SK689, D P. kolea, holotype, Hawaii (Big Island), SK851, E P. lysimachiella, Oahu, SK656, F P. sp. 7, Kauai, SK754, G P. molokaiensis, Molokai, SK666, H P. basalis, Hawaii (Big Island), SK614, I P. splendida, Maui, SK696, J Same species, Oahu, SK747, host Syzygium sp., K P. lama, holotype, Maui, SK744, L P. napaliensis, paratype, Kauai, SK745. Abbreviation: ta: tuba analis.

FIGURE 43. Phallus of Philodoria, lateral view. A P. succedanea, Maui, SK641, B P. auromagnifica, Hawaii (Big Island), SK800, C P. auromagnifica, Kauai, SK689, D P. kolea, holotype, Hawaii (Big Island), SK851, E P. lysimachiella, Oahu, SK656, F P. sp. 7, Kauai, SK755, G P. molokaiensis, Molokai, SK666, H P. basalis, Hawaii (Big Island), SK614, I P. splendida, Maui, SK696, J Same species, Oahu, SK747, host Syzygium sp., K P. lama, holotype, Maui, SK744, L P. napaliensis, paratype, Kauai, SK767.

FIGURE 61. Images of genitalia slides prepared by E. C. Zimmerman. A–D: P. auromagnifica, male, Oahu, Z-XII-20-62-5, E–G: P. auromagnifica, female, Oahu, Z-XII-20-62-6, H: P. basalis, female, Hawaii (Big Island), Z-XII-20-62-4, I–N: P. costalis, paralectotype male, Oahu, Z-XII-20-62-2. A, M Phallus, lateral view, B Entire genitalia except phallus, ventral view, C Coremata, D Abdominal segments II–III, E Abdominal segments II–VI, F Tergum II, G, H Entire female genitalia, I Abdominal segments II–V, J Right valva, K Genital capsule, ventral view, L Left valva, N Abdominal segment VIII with dorsal flap and coremata.

FIGURE 81. Philodoria basalis and its host plant, Metrosideros polymorpha. A, B, D–M: Hawaii (Big Island), C: Kauai. A Leaf mine of late instar larva, B Fallen cocoons in rolled leaf disc “tacos”, C Fallen leaf and leaf mine with cocoon, D Cocoon and mine, E, F Old mine, G Mine made by late instar larva, H, I Late instar larva, J Final instar larva cutting out disc to create oval “taco” shelter with silk, K, L Cocoon and mine showing the “taco” cut out on the leaf, M Cocoon “taco”, similar to B.