Philodoria, 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 : 17-18

Key to adults

(Adults of Philodoria sp. 2 and P. sp. 3 are unknown and are thus excluded from this key.)

- Forewing shiny, metallic bronze with ocherous to orange patches ( Fig. 3A )....................................... A

- Forewing brown to fuscous with orange patches and a white spot near middle of wing ( Fig. 3B )....................... B

- Forewing brown with one to three costal white streaks and two to three dorsal white streaks ( Fig. 3C ).................. C

- Forewing white with costal ocherous patches and three dorsal ocherous patches ( Fig. 3D )............................ D

- Forewing brownish fuscous to black with white patches ( Figs. 12 , 13 )........................................... E

- Forewing leaden gray, externally with fuscous brown ( Fig. 7F )............................................ P. kolea

(A).

1 Basal patch (bp) absent; basal half of forewing leaden gray color ( Fig. 8C–F )............................... P. basalis

- bp present........................................................................................... 2

2 A bright orange transverse fascia (tf) at 3/ 4 in middle interrupted with blue patch; an orange medial transverse patch (tp), narrowing towards dorsum, ( Fig. 7C )............................................................ P. kauaulaensis

- An orange transverse patch beyond middle to costal 3/4, narrowing towards dorsum, extending to dorsal 2/3, with white costal spot ( Fig. 7A, B, D, E , 8A, B , 9 )......................................................................... 3

3 A black bp along costal fold; a fuscous patch near apex ( Figs. 7A, B ).................................. P. succedanea

- An orange bp along costal fold, fringed with blackish scales ( Fig. 7E ); a fuscous patch with dark orange scales at apex........................................................................................... P. auromagnifica

- An ocherous bp from near base of costal fold to dorsal 1/4 ( Fig. 9 )...................................................................... P. splendida ( Fig. 9A, B , 73B ), P. lama sp. n. ( Figs. 9C , 73E, F ), P. napaliensis sp. n. ( Fig. 9D ) § § It is difficult to identify three species: lama , napaliensis , and splendida from forewing pattern, see key to genitalia and diagnosis of species.

- An ocherous to oraenge bp containing two longitudinal streaks................................................. 4

4 An ocherous-orange tf from costal 1/3 to dorsal 1/2; bp from base to 1/3 ( Figs. 8B , 73A )................ P. molokaiensis

- tf absent............................................................................................ 5

5 bp from base to 1/2; bb completed ( Fig. 8A ); Oahu............................................. P. lysimachiella # 2

- Basal 1/4 forewing unknown, but orange patch reach 1/2; bb 2 interrupted at middle by an orange patch ( Fig. 26G ); Kauai................................................................................................. P. sp. A # P. sp. 7 has similar forewing pattern, but basal 1/4 of forewing missing and damaged.

(B).

1 ws at near middle of the wing, not reaching dorsal margin ( Fig. 3B )............................................. 2

- ws reaching dorsal margin ( Fig. 74B )..................................................................... 3

2 ws oval ( Fig. 10C ).................................................................................... 4

- A small oblque ws at near middle of the wing; Oahu ( Fig. 11A–D ).................................... P. pipturicola

3 ws large, about 3x in diameter than that of costalis ; w 2 widening toward the wing margin; Oahu ( Fig. 10B, C )... P. pipturiella

- White spot small; two white bands almost the same size; Oahu ( Fig. 10A )................................. P. costalis

4 ws almost erect line; Kauai ( Fig. 10F ).......................................................... P. micropetala

- ws bent toward apex; Hawaii ( Big Island ) (and Maui) ( Figs. 10D, E , 11E, F )...................................... 5

5 cs and w 1 rather narrowing; Hawaii ( Big Island ) and Maui ( Figs. 10D, E , 26M )............................. P. floscula

- cs and w 1 rather broad; Hawaii ( Big Island ) ( Figs. 11E, F , 26K, L , 74 )........................... P. haelaauensis sp. n.

(C).

1 A lemon-yellow streak from costal fold to 2/5, obliquely, at 3/5 to the middle ( Figs. 20C , 77A )................ P. hauicola

- Three white costal streaks (cs

1–3

) present; white streaks are very narrow ( Figs. 23A–C , 77E, F ).......... P. marginestrigata

- Two white costal streaks (cs 2 and cs 3) present ( Figs.14 , 15 ).................................................... 2

- One white streak (cs 3) present ( Figs.16–22 )................................................................. 4

2 A white longitudinal streak in basal half of wing ( Fig. 14A )......................................... P. dubauticola

1 2

- A broad white streak at costal fold; a transverse fascia at 3/4 (cs 3 + ds 3) ( Fig. 22A, B ).................. P. keaenisis sp. n.

- Not so, ds 1 present or brown in basal half.................................................................. 3

3 ds absent; cs rather short, not reaching the middle of wing ( Figs. 14F , 15A–E )..................... P. alakaiensis sp. n.

- ds 1 present; cs 2 reaching the middle of wing ( Figs. 14B–D , 15F, G )............... P. dubautiella , P. limahuliensis sp. n. ** ** P. limahuliensis has more dark color forewing.

4 Three white dorsal streaks (ds

1–3) present respectvely......................................................... 5

- ds

2 connecting basal patch on dorsum (ds

1)................................................................. 8

- ds 1 absent; a small white costal spot at near base; blue to fuscous patch at apical portion ( Figs. 22D , 75D–G ).............................................................................................. P. platyphylliella sp. n.

5 ds

1 is small white patch ( Fig. 16C, D ); ds

3 rather small.............................................. P. naenaeiella

- ds 1 is triangular-shaped streak ( Fig. 21E ); ds 3 rather small (See also key of P. nigrella )................... P. touchardiella

- ds 1 reaching the middle of wing.......................................................................... 6

6 Base to apex of ds 1 and ds 2 are almost same in width; ds 3 is triangular-shaped white patch ( Fig. 19C–E ) P. obamaorum sp. n.

- ds 1, 2 are slightly widend at dorsum; ds 3 is nearly triangular-shaped white patch ( Fig. 19A, B ).............. P. neraudicola

- ds 1–3 are widened at dorsum ( Figs. 16A, B , 17 , 18 , 75B, C , 77B–D , 78E, F )....................................... 7

7 Rather narrowing ds:

- Apical portion orange with fuscous scales at center and a large as; ocherous color forewing ( Figs. 16A, B , 77B–D )................................................................................................... P. hibiscella

- Apical portion pale lemon-yellow with a small as; brounish fuscous to brown color forewing ( Figs. 20A, B , 78E, F )................................................................................................... P. urerana

- Rather wider ds from dorsum to the middle; apical portion with blue patch ( Figs. 17 , 18 , 75B, C ).................................................................................. P. hesperomanniella sp. n., P. knudseniiella sp. n.

8 ds 1 is narrowing white band ( Figs. 20F , 76 )................................................................. 9

- ds 1 is weakly triangular ( Fig. 21B, D ).................................................................... 10

9 A fuscous patch at dorsum 3/4; dark brown color forewing ( Fig. 76 )........................................ P. sp. 1

- A fuscous patch at 3/4, not reaching dorsal margin; ocherous color forewing ( Fig. 20F )................... P. keahii sp. n.

10 Apical portion fuscous; bronze brown color forewing ( Fig. 21A, B )................................. P. funkae sp. n.

- Apical portion light ocherous brown, but damaged; light ocherous brown color forewing ( Fig. 21D )......... P. pittosporella

(D).

1. A fuscous costal line from base to about 1/2 ( Fig. 25C, D ).......................................... P. opuhe sp. n.

- cp 1 absent ( Figs. 25A, B , 78D )................................................................... P. ureraella

- cp 1 present; dp 1 indistinct, connecting mp (ds 2 connecting ds 1) ( Figs. 23D–G , 24 , 78A–C )........................... 2

2 dp 2 and dp 3 not meeting..................... P. lipochaetaella ( Figs. 23D–G , 24A ), P. sciallactis ( Figs. 24B–G , 78A–C )

- dp 2 and dp 3 meeting at median line ( Fig. 24H )......................................................... P. sp. B

(E).

1 An oval white spot about the middle of wing ( Fig. 12A–C ).................................................... 2

- An oval white spot absent ( Figs. 12D , 13 ).................................................................. 3

2 A longitudinal white streak from base to 1/3; an oblique white costal spot at 2/3, recurves to the costa ( Fig. 12A–C )................................................................................................. P. pipturiana

- A white costal spot at 2/3, not curve ( Zimmerman 1978a: fig. 467)....................................... P. spilota

3 Two or three costal spots ( Fig. 12D )............................................................ P. wilkesiella

- A costal streak (cs 3) at 3/4 ( Fig.13 )....................................................................... 4

4 Dorsal streaks (ds 1–3) rather wider at dorsal base; ds 1 indistinct or along dorsal margin ( Fig. 13A–C ).......... P. epibathra

- ds 1–3 rather narrow; ds 1 reaching the middle of wing ( Fig. 13D, E )........................................ P. nigrella

- ds 1–3 rather narrow; ds 1 is very small white patch ( Zimmerman, 1978a: fig. 466; Fig. 22C )................ P. nigrelloides

Figures

FIGURE 3. Nomenclature adopted in this study for the characterization of the four major Philodoria forewing pattern types (A-D). After Kobayashi et al. (2018). A tf and tp present, B wp present at the middle, tf absent and tp obscure, C cs 1~3 and three ds present, D cp and dp present and cs and ds indistinct. a, b, c: white line on costal cilia, as: apical spot, bb(1–3): bronze color band, bp: basal patch, bl(1–2): basal line, cl: costal line, cs (1–3): costal streak, cp: costal patch, ds(1–3): dorsal streak, dp(1–3): dorsal patch, op: orange patch, mp: median patch, tf: transverse fascia, tp: transverse patch, us: uplight spot, w: white color band, (1–5) ws: white spot.

FIGURE 7. Adults of Philodoria species. A P. succedanea Walsingham, 1907 lectotype female, B Paralectotype male, C P. kauaulaensis Kobayashi, Johns & Kawahara, 2018 holotype female, D Philodoria sp. nr. P. auromagnifica male Kauai, E P. auromagnifica Walsingham, 1907 holotype male, F P. kolea Kobayashi, Johns & Kawahara, 2018 holotype male. Scale bar 1 mm.

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 9. Adults of Philodoria species. A P. splendida Walsingham, 1907 lectotype male, B Female, Oahu, C P. lama sp. n. holotype male, D P. napaliensis sp. n. holotype (sex unknown). Scale bar 1 mm.

FIGURE 10. Adults of Pipturus feeding Philodoria species. A P. costalis Swezey, 1934 lectotype male, B P. pipturiella Swezey, 1923 lectotype male, one of two cotypes (upper specimen) on the same mount, C Paralectotype male on the same mount as of lectotype (lower specimen), D P. floscula Walsingham, 1907 lectotype female, E Paralectotype male, F P. micropetala Walsingham, 1907 holotype female. Scale bar 1 mm.

FIGURE 11. Adults of Pipturus feeding Philodoria species. A P. pipturicola Swezey, 1915 paralectotype male, one of four cotypes (upper left specimen) on the same mount, B Lectotype male (upper right specimen on mount), C Paralectotype female (lower left specimen on mount), D Paralectotype female (lower right specimen on mount), E P. haelaauensis sp. n. male, F Paratype female. Scale bar 1 mm.

FIGURE 12. Adults of Philodoria species. A P. pipturiana Swezey, 1923 lectotype male, left lateral view, B Same right lateral view, C Same dorsal view, D P. wilkesiella Swezey, 1940 lectotype male. Scale bar 1 mm.

FIGURE 13. Adults of Philodoria species. A P. epibathra (Walsingham, 1907) holotype female, B Same, left forewing, host Dubautia sp., CJ492, Molokai, C Same, right forewing, D P. nigrella (Walsingham, 1907) lectotype male, E Paralectotype male. Scale bar 1 mm.

FIGURE 14. Adults of Philodoria species that have two costal streaks (cs1 and cs2). A P. dubauticola Swezey, 1940 lectotype female, B P. dubautiella (Swezey, 1913) lectotype male one of two cotypes on the same mount (upper specimen), C Paralecto- type female (lower specimen), D Female, Oahu, E P. sp. nr. P. dubautiella male, Kauai, F P. alakaiensis sp. n. holotype male. Scale bar 1 mm.

FIGURE 15. Adults of Philodoria species that have two costal streaks (cs1 and cs2). A Forewings of P. alakaiensis sp. n. holotype male, B Paratype male, C Same as B, forewings, D Paratype male (BMNH(E)1621107) in NHMUK, E Paratype female

FIGURE 16. Adults of Philodoria species. A P. hibiscella (Swezey, 1913) lectotype male, one of two cotypes on the same mount (upper specimen), B Paralectotype female (lower specimen on mount), C P. naenaeiella (Swezey, 1940) lectotype male, one of two cotypes on the same mount (upper specimen), D Paralectotype female (below holotype specimen on mount). Scale bar 1 mm.

FIGURE 17. Adults of Philodoria knudseniiella sp. n. A holotype lateral view (CJ480/SK862), B Same, dorsal view, C Same, right wings, D Paratype female right wings (CJ480/SK863), E Paratype female (34241 BPBM slide Z-I-10-66-A). Scale bar 1 mm.

FIGURE 18. Adults of Philodoria hesperomanniella sp. n. A Wings of holotype male, B Same, head and thorax, dorsal view, C Same, head lateral view, D Paratype female, Maui, collected by O. H. Swezey, E Paratype female, Oahu (BPBM no.34224). Scale bar 1 mm.

FIGURE 19. Adults of Philodoria neraudicola (Swezey, 1920) and P. obamaorum sp. n. A P. neraudicola lectotype male, B Forewings of lectotype female, host Pipturus albidus, Maui, C P. obamaorum holotype male, D Paratype female, E Female (CLV017). Scale bar 1 mm.

FIGURE 20. Adults of Philodoria species. A P. urerana (Swezey, 1915) lectotype female, one of two cotypes on the same mount (upper specimen), B Paralectotype (sex unknown, lower specimen on same mount as A), C P. hauicola (Swezey, 1910) lectotype male, D P. keahii sp. n. holotype male, E Wings of holotype, F Left forewing of paratype female. Scale bar 1 mm.

FIGURE 21. Adults of Philodoria species. A Wings of P. funkae sp n., holotype female, B Right forewing of holotype shown in A, C Wings of paratype female, D P. pittosporella (Swezey, 1928) holotype male, E P. touchardiella (Swezey, 1928) lectotype female. Scale bar 1 mm.

FIGURE 22. Adults of Philodoria species. A P. keaensis sp. n. holotype female, lateral view, B Forewings of holotype, C P. nigrelloides (Swezey, 1946) holotype female, D Forewings of P. platyphylliella sp. n. holotype male. Scale bar 1 mm.

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 24. Adults of Philodoria species. A P. lipochaetaella, specimens originally described as Parectopa lipochaetae Swezey, 1946, syntype male B P. sciallactis (Meyrick, 1928) lectotype male C Paralectotype male D Paralectotype female E Male left forewing of specimen CJ164, host: Melanthera kamolensis, Maui F Same specimen, right forewing G Male right forewing of specimen CJ189, host: Melanthera kamolensis, Maui H Philodoria sp. B nr. P. sciallactis (specimen CJ483, sex unknown) right forewing, host: Lipochaeta acris, Kauai. Scale bar 1 mm.

FIGURE 25. Adults of Philodoria ureraella (Swezey, 1915) and P. opuhe sp. n. A P. ureraella paralectotype (sex unknown) one of four cotypes on the same mount (upper left specimen on mount), B lectotype male (lower left specimen on mount), C P. opuhe holotype male (upper right specimen on mount), D P. opuhe paratype (sex unknown, lower right specimen on mount), E Types of P. ureraella and P. opihe on type mount 213 (BPBM) (= four cotypes of ‘P. ureraella’ by Swezey). 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 73. Resting posture of adult Philodoria and Caloptilia. A P. molokaiensis, Molokai, CJ229, host: Lysimachia hillebrandii, B P. splendida, Oahu, CJ319, host: Metrosideros polymorpha, C Same species, Oahu, D Caloptilia mabaella, Lanai, CJ333, host: Diospyros sandwicensis, E P. lama, Maui, CJ190, F Same species, Maui, CJ194.

FIGURE 74. Resting posture of adult P. haelaauensis, male, Makawao Forest Reserve, Maui, CJ227/PHIL0019, host: Pipturus rockii. A lateral view, B dorsolateral view.

FIGURE 75. Resting posture of adult Philodoria. A Philodoria sp. nr. P dubauticola, Maui, CJ315, host: Dubautia platyphylla, B P. hesperomanniella, Maui, CJ367, host: Hesperomannia arborescens, C Same moth species and host plant, West Maui, CJ362, D, E P. platyphylliella, Maui, CJ368, host: Dubautia platyphylla, F, G Same moth species and host plant, male, Maui, CJ358.

FIGURE 76. Resting posture of adult Philodoria sp. 1 on Dubautia menzesii, Maui (CJ334). A dorsolateral view, B dorsal view, C lateral view, D lateral view with wings held open.

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 78. Resting posture of adult Philodoria on their larval host A P. sciallactis, Maui, CJ195, host: Melanthera kamolensis, B, C P. sciallactis, Maui, CJ164, 189, host: M. kamolensis, D P. ureraella, Maui, CJ316, host: Urera glabra, E, F P. urerana, Oahu, CJ307, host: U. glabra.