Philodoria succedanea Walsingham, 1907

Kobayashi, Shigeki, Johns, Chris A., Lopez-Vaamonde, Carlos, Camiel Doorenweerd,, Kawakita, Atsushi, Ohshima, Issei, Lees, David, 2018, Hawaiian Philodoria (Lepidoptera, Gracillariidae, Ornixolinae) leaf mining moths on Myrsine (Primulaceae): two new species and biological data, ZooKeys 773, pp. 109-141: 109

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Philodoria succedanea Walsingham, 1907


Philodoria succedanea Walsingham, 1907  Figs 2 A–D, 5 A–D, 6 A–D, 7E, F, 8A, 9, 14A

Philodoria succedanea  Walsingham, 1907: 717-718; pl. 25, fig. 19.

Philodoria (Philodoria) succedanea  Walsingham, 1907: Zimmerman 1978: 718, figs 433, 435, 467, 472.

Type locality.

Olinda, Haleakala (Maui).

Type material.

Lectotype ♀, Olinda, 4000 ft., Haleakala, MAUI, Hawaiian Is. iv.1894, Perkins. 26695 [Walsingham specimen number]|PHILODORIA SUCCEDANEA Wlsm. Fn. Hawaii. I TYPE ♀ descr. figd.|Walsingham Collection. 1910-427.|NHMUK010305341 (here designated).

Paralectotypes 17 (2♂ 1♀ 14 unsexed; NHMUK ones are all from above Walsingham accession and ‘PARATYPE’ below is short for 'PHILODORIA SUCCEDANEA Wlsm. PARATYPE’ as printed on large black-margined labels, with the 5-digit Walsingham specimen numbers whose first digit is ‘2’ borne on the locality label): 1 ♂, Haleakula 4000 ft. MAUI, Hawaiian Is. V. 1896|Perkins. 28505|PHILODORIA SUCCEDANEA Wlsm. Fn. Hawaii. I TYPE ♂|BM ♂ Genitalia slide no. 2755|NHMUK010305341. 1♀ 2 unsexed: Haleakala, 5000ft, MAUI, Hawaiian Is., v.1896, Perkins. 28355|PARATYPE 3/17|NHMUK010862804|; 28230|PARATYPE 4/17♀| BPBM 34324; 28236|PARATYPE 5/17| BPBM 34321. 4 unsexed: Haleakula -4000 ft. Maui, v. 1896, Perkins. 28492|PARATYPE8/17; 28493|PARATYPE 9/17| BPBM 34320|; 28494|PARATYPE10/17|NHMUK010862806; 28495|PARATYPE 11/17|NHMUK010862807.

1 ♂ 7 unsexed, same data and locality as lectotype: 26696|NHMUK010862803; 26661|PARATYPE1/17|BPBM34325; 26667|PARATYPE2/17| BPBM 34222; 28511|PARATYPE12/17|NHMUK010862808; 28512|PARATYPE13/17|NHMUK010862809; 28513|PARATYPE14/17|NHMUK010862810; 28514|PARATYPE15/17♂|NHMUK010862811; 28552|PARATYPE 16/17| BPBM 34323.

This species was described from 19 specimens: 'type ♀ (26695); ♂ (28505)' and 17 ‘paratypes’ from Kauai and Haleakala, Maui. This seems to indicate that Lord Walsingham considered them as holotype, allotype, and paratypes, as indicated on their specimen labels. But as a holotype was not specified in the description, the so-labelled types and paratypes are all to be considered syntypes under the present Code, Article 73.2 ( ICZN 1999), and any one is thus eligible for designation as lectotype. The syntype 'type ♀ (26695)', which Walsingham listed first and figured, is here designated as lectotype (Fig. 2A). The remaining syntypes are therefore paralectotypes.

Additional material.

32 (11♂ 15♀ 6 unsexed).

Adults: Oahu Is.: 2♀, Mt. Kaala, 18.ii.1923, Swezey coll., host: " Suttonia  " (= Myrsine  ), SK797♀, 798♀ in USNM; 3♂ 3♀, Palikea, 28.iii.2016 stored in 99 % ethanol (stored), K. Bustamente leg., host: M. lessertiana  , 10.xi.2015, CJ-526, CJ-531, SK639♂, SK803♀, 804♀, 807♂, 808♂ in BPBM.

Molokai Is.: 1♀, Kainalu [Kainalu Forest, South East Molokai Forest Reserve], 27?vii.1927, Philodoria auromagnifica  Walsingham Det. by O.H. Swezey, 34145 in BPBM; 1 unsexed, 4000 ft Molokai P. 2.02 Philodoria succedanea  Wals. 1/1 E. Meyrick det. in Meyrick coll., in NHMUK.

Lanai Is.: 2♂, 2750 ft, Munro Trail, 2.x.1976, K. & E. Sattler BM1976-605, BMNH(E)1621676 and BMNH(E)1621677, Philodoria  sp. 8 (Lanai) Sattler coll. D.C. Lees Sep 2016.

Maui Is., in BPBM: 1♂ 6♀ 1 unsexed, below Eke, 17&21.v.2013 (stored), C.A. Johns leg., host: Myrsine  sp., 24.iv.2013, CJ-136, 141, SK799♀; 1♂, Waikamoi, 24.v.2016 (stored), C.A. Johns leg., Spring.2016, CJ-539, SK641♂.

Hawaii Is., host: M. lessertiana  in BPBM: 1♂ 1♀, Hawai’i Volcanoes National Park, Hawaii, A. Kawakita leg., “Leaf-dropper”, 25.iv.2016 (larva), SK624♀, SK625♂; 2♂ 1♀ 1 unsexed, Same locality, 17&21.v.2016 em., A.Y. Kawahara leg., 29.iv.2016(Cocoon & larva), SKH-10, SK801♂, SKH-13, SK633♂. 1♀, 3800 ft, N. Kohala, Distr. Kohala Mts, Puu Laalaau area, 14-17.vii.1976, K. & E. Sattler, BM1976-605, BMNH(E)1621089, Philodoria  sp. 9 (Hawaii) Sattler coll. D.C. Lees Sep 2016, 1621676 in NHMUK; 1♂, same data as last specimen, BMNH(E)1621090, Philodoria  sp. 9 (Hawaii) Sattler coll. D.C. Lees Sep 2016; 1 unsexed, Kohala Watershed Partnership, (stored), C.A. Johns leg., host: Myrsine sandwicensis  , 18.v.2015, CJ-419 in BPBM.

Larvae: 2 unsexed, Kokee, Kauai Is., 16& (stored), C.A. Johns leg., host: M. knudsenii (larva), CJ-433, 442 in FLMNH.


This species is very similar to P. auromagnifica  feeding on the same hostplant, Myrsine  , but is recognizable by the rather bright orange patches and black triangular shaped basal patch in the forewing (Table 4; Figs 2 A–D, 5 A–D); in the male genitalia by the rather broad valva, slender and long saccus curving toward dorsal side (Fig. 6 A–C); in the female genitalia by signa with slender and long spines (Fig. 7E, F).


Adult (Figs 2 A–D, 5 A–D, 8A). Wingspan 9-10 mm in type series; forewing length 4 mm in 'TYPE ♂ (28505)' (fig. 2B), 3.6-3.8 mm in paralectotypes. Head bronze; frons white; maxillary palpus reduced; labial palpus bronze grey, with dark brown scales at apex. Antenna shiny tawny fuscous. Thorax bronze.

Forewing shiny, metallic bronze with bright orange-ochreous patches: a black triangular basal patch along the costal fold (Figs 5A, C, 8A); an oblique transverse fascia before the middle of wing, bordered with black scales; a large transverse patch after the middle to costal 3/4, distinctly narrowing in the dorsum, extending to dorsal 2/3, containing white costal spot; one white color band on the middle of the first bronze color band, others on both extremities of second and third bands; a fuscous patch extending toward the termen and apex with a black apical spot, sometimes with orange-ochreous color encroaching on the apical part; cilia tawny, with two metallic silver basal lines, one at the apical cilia, another from termen to tornus. Hindwing dark tawny; cilia tawny. Abdomen tawny above, silvery beneath. Legs tawny, with silvery spurs and slightly paler tarsi.

Male genitalia (Fig. 6 A–D) (n = 7). Capsule 960 µm. Uncus absent. Tegumen 570-580 µm long, 1.2 –1.3× length of valva with series of long hairs at lateral side of base (Fig. 6C). 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. Valva broad, 430 µm in length covered with fine setae distally, and having a short dorsal process (Fig. 6A).Vinculum U-shaped; saccus 250 µm long, slender , curved toward dorsal side (Fig. 6B, C). Phallus 720 µm long, tubular and long about 1.2 –1.3× length of valva, sinuous in lateral view with two series of minute spiniform cornuti in vesica; coecum slightly curved toward inner side (Fig. 6D).

Female genitalia (Fig. 7E, F) (n = 7). Ostium bursae rather small, opening at the middle of 7th abdominal segment; antrum cup-shaped with slender a pair of lateral lobes; ductus bursae slender, tubular, extremity connected to antrum very slender and membranous, curved inside of body, and middle part weakly sclerotized and plate-shape; end of the ductus bursae broad; inception of ductus seminalis on the posterior part of ductus bursae. Corpus bursae pyriform, anterior end weakly sclerotized; some lines consisting of wrinkles running longitudinally, some sclerotized; paired signa with a pair of long slender spines.


Kauai, Oahu and Lanai: new record, Maui (Walsingham 1907), Molokai and Hawaii (Big Island) ( Zimmerman 1978).

Host plants.

Primulaceae  : Myrsine sandwicensis  A. DC., M. lessertiana  A. DC. ( Johns et al. 2016), Myrsine  sp. ( Zimmerman 1978). Myrsine linearifolia  Hosaka and M. knudsenii  (Rock) Hosaka are new host records (see Remarks).


(Figs 8A, 9, 14A). The larvae mine the adaxial side of leaves of Myrsine  species, forming a long linear mine (Fig. 9B, G, H). The mine is at first tornus-shaped (Fig. 9C, D, I, J) and the larva broaches the mid vein towards the petiole of the leaf, forming a straight mine; the vein mine and surrounding pattern are red in coloration (Fig. 9F, H) and later instars leave the mid vein usually near the base of the leaf, gradually expanding as they feed and grow forming a full-depth mine (Fig. 9E, F). There were usually one to two mines per leaf (Fig. 9B, G, M). The pupal cocoon is situated outside of the mine, usually on the leaf surface, and also on the woody tissue of the host plant with leaf mines and larvae. At Hawai’i Volcanoes National Park, larvae were collected from leaves that had fallen to the ground and reared to adulthood (Fig. 9A). The adult has been observed during the day (Maui and Hawaii Island), resting on the upper leaf surface of the host plant (Fig. 8A).

DNA barcoding.

BIN BOLD:ADF5435. The two specimens sequenced for COI, one from Maui and one from Hawaii, have identical DNA barcode sequences. The p-distance to the nearest neighbor, P. kauaulaensis  , is 6.63%.


We identified two adult moths (Coll ID CJ-144 / GenBank accession no. ID KT982414 and CJ-145) as P. succedanea  , based on the presence of a basal black patch on forewing, from which whole bodies were sacrificed for molecular analysis ( Johns et al. 2016; Figs 6O, 12). Zimmerman (1978) did not recognize Walsingham’s (1907) Kauai record of this species because Walsingham had only one specimen at hand, which was in poor condition (specimen data: 1 ♂, Mts [which Mts not further specified], 3-4000 ft., Kauai, vi. 1894 Perkins.27297| PARATYPE 17/17 (?)|'NOT succedanea  Det. by E. C. Zimmerman|NHMUK010862812). We could not find the specimen from Kauai. However, we found Myrsine knudsenii  (Endangered, IUCN) leaves with mines with active larvae from Kokee, Kauai Is. (CJ-433, 442), which were similar in appearance to mines of P. succedanea  on M. lessertiana  . Judging from these data, we consider the larval mines on M. knudsenii  were made by P. succedanea  . We also collected active Philodoria  leaf mines from M. linearifolia  (Endangered, IUCN) at the same location as M. knudsenii  , but were unable to rear adult moths. It is thus possible that P. succedanea  also mines M. linearifolia  , but this needs to be further examined.