Abstract.

In this study, Elaeonoma ultima sp. nov., a new species of oecophorid moth from the Ogasawara Islands, Japan is described. Photographs of adults, male and female genitalia, and aspects of this species’ biology, including adult behavior, egg-laying substrates, and hatched larval behavior, are shown. A phylogenetic tree and haplotype network were constructed based on the cytochrome c oxidase subunit I (COI-5P) DNA barcode sequences, including those of some Elaeonoma species related to this new species. In addition, the systematic position, geographical variation, habitat, and potential effects of alien predators on this new species are discussed.

Introduction

The Ogasawara Islands, an oceanic island archipelago, are located approximately 1,000 km south of mainland Japan in the Northwest Pacific Ocean. Only a limited number of plant and animal species have reached these islands and were subsequently isolated from the surrounding continents. These ancestors have evolved into new endemic species with approximately 40% plants, 30% insects, and 90% land snails within the islands (Toyoda 1981; Kato 1992; Tomiyama 1992; Shimizu 2003). Due to their unique and highly endemic biota, the Ogasawara Islands have been registered as a UNESCO World Natural Heritage Site (UNESCO World Heritage Centre 1992–2025; Government of Japan 2010). The Ogasawara Islands are the only places in Japan that belong to the Oceanian Region, and many insects and plants are found in this region, including species from Micronesian regions (Ito 1998; Takakuwa 2004; Yamada and Hirowatari 2007).

In Lepidoptera, many endemic species with various feeding habits have been described; for example, herbivores of endemic plants, Acidon sugii (Erebidae); detrivores of birds’ nests, Erechthias nidumicola (Tineidae); and the fungivore and detrivore, E. flavimacula (Watabiki et al. 2017; Park et al. 2025). Among herbivores, many endemic species are known to be mono- or oligophagous that feed on endemic plants (Takeuchi and Ohbayashi 2006). However, information on the detailed feeding habits of endemic fungivores and detrivores (bark or dead leaf feeders) is insufficient.

Kato (1992) provided a checklist of insect species from the Ogasawara Islands and was the first to list two species of the family Oecophoridae from the Chichijima and Anijima Islands. Takeuchi and Ohbayashi (2006) summarised moth species from the Ogasawara Islands and listed the above-mentioned species as “Oecophoridae Gen. et spp. (two species).” However, these species have not been identified yet.

Meyrick (1914) established the genus Elaeonoma based on the type species, Eulechria piodes Meyrick, 1902 from Queensland, Australia. In the same study, Meyrick (1914) transferred Eulechria autogramma Meyrick, 1905 from Ceylon and Eulechria chersodes Meyrick, 1914 from New Guinea to this genus and described E. nephelissa Meyrick, 1914 from Ceylon. There have been many taxonomic changes, however, and the concepts of Elaeonoma sensu Clarke (1984) and Common (2000) currently include four species: E. deltacostamela, E. adeneia, E. moira, and E. swezeyi, with the distribution in the areas of mainland Australia and the Micronesian Islands.

During microlepidopteran surveys of the Ogasawara Islands from 2022 to 2025, we collected unknown oecophorid moths from the Chichijima and Hahajima Islands and identified them as undescribed species of Elaeonoma. In this study, we describe it as a new species of Elaeonoma, provide some biological notes, and infer its phylogenetic relationships.

Materials and methods

Materials and observations

All examined specimens were collected using light traps (LT) at night and sweeping (SW) during the day on Chichijima and Hahajima Islands, both in the Ogasawara Islands. All specimens were deposited at the Entomological Laboratory of Kyushu University (ELKU). Morphological characteristics were observed using a Leica S9D stereoscopic microscope. The genitalia were dissected, stained with chlorazol black E and mounted in euparal. Terminology of morphology follows Scoble (1992) and Common (1994). A distribution map was created with QGIS 3.10.11 (QGIS 2020) using data from the Global Map Japan (Global Map Japan version 2.2 Vector data, https://www.gsi.go.jp/kankyochiri/gm_japan_e.html).

DNA analysis

Seventeen specimens were used for DNA analysis. Total DNA was extracted from the abdomen using a DNeasy Blood & Tissue Kit (Qiagen, Hilden, Germany), following the manufacturer’s instructions. We amplified the fragment of the partial mitochondrial COI gene using the primer set LepF1/LepR1 (Hebert et al. 2003) and the KOD One® PCR Master Mix -Blue- (TOYOBO CO., LTD., Osaka, Japan). Amplification conditions followed the manufacturer’s protocol, except for the annealing temperature, which was set to 50 °C. The amplicons were purified using the ExoSAP-IT Express (Thermo Fisher Scientific Inc., Massachusetts, USA). Cycle and bidirectional sequencing were conducted using premixed Sanger sequencing services (Eurofins Genomics, Tokyo, Japan).

The sequences obtained were manually aligned using MEGA 7.0.26 (Kumar et al. 2016). The sequence data were compared with the sequences registered in BOLD Systems, and similar species were searched using the BOLD identification engine (https://www.boldsystems.org/index.php/IDS_OpenIdEngine).

To construct a preliminary phylogenetic tree, we acquired sequences from BOLD, including all described Elaeonoma species and the similar genus Leimmatonca with registered COI barcode sequences longer than 500 bp. The sequence of Promalactis sakaiella Matsumura, 1931 (Oecophoridae) was used as an outgroup. A maximum likelihood (ML) phylogenetic tree was reconstructed using RAxML version 8.0.0 with an auto-substitution model selection and rapid bootstrap analysis with 1000 repetitions. The treefile was visualized using Figtree v1.4.4 (Rambaut 2018) to prepare the figure.

A haplotype network was constructed to clarify the inter-island genetic structures of the new species. Haplotype network analyses were conducted using the minimum spanning network algorithm of the Pop-ART software package (https://popart.maths.otago.ac.nz/), based on the COI barcode region. The unique haplotypes were determined using DnaSP v. 6.12.03 software (Rozas et al. 2017).

Results

Elaeonoma Meyrick, 1914

Elaeonoma Meyrick, 1914: 238; Meyrick 1922: 68; Turner 1935: 333; Gaede 1938: 115; Clarke 1963: 193; Clarke 1984: 146; Common 2000: 81.

Type species.

Eulechria deltacostamela Lower, 1896: 165.

= Eulechria piodes Meyrick, 1902: 148, synonymized by Turner (1935).

= Eulechria deltomela Meyrick, 1922: 72, synonymized by Gaede (1938).

Type locality.

Queensland (Australia).

Type depository.

Commonwealth Scientific and Industrial Research Organization (CSIRO).

Remarks.

The genus Elaeonoma Meyrick, 1914 (Gelechioidea: Oecophoridae) is characterised by a very short proboscis, stalked veins CuA1 and CuA2 of the forewing, lack of the distal process of the sacculus of the male genitalia, short apophysis posterior, very short ductus bursae, and small membranous corpus bursae of the female genitalia (Common 2000). This genus is placed in the Barea genus group and is related to Baioglossa Common, 2000 and Eremnozona Common, 2000 (Common 2000), but is distinguished by having developed gnathos and a relatively short cornutus of the male genitalia. The biology of Elaeonoma is poorly understood; however, some adults have been collected using light traps, and E. swezeyi has been reared on dead plant of Barleria sp. (Acanthaceae) (Clarke 1984; Common 2000).

Turner (1916) described two new species from Australia, and Turner (1918) described a new species from Lord Howe Island near the Australian Continent. Meyrick (1922) listed 12 species from Australia (8 spp.), Ceylon (2 spp.), and New Guinea (2 spp.) with some generic transfers. Turner (1935) reviewed the Australian species (including Tasmania Island) of this genus and listed 28 species, including 17 new species. Turner (1935) synonymized Elaeonoma piodes with Eulechria deltacostamela Lower, 1896, without any morphological evidence. Gaede (1938) listed 31 species in this genus. Bradley (1956) described one new species from Lord Howe Island. Clarke (1963) illustrated and described the venation of the right wing, markings of the left wing, and genitalia of the type specimens of three species, E. piodes (= E. deltacostamela), E. autogramma, and E. nephelissa. Clarke (1984) described three new species, E. adeneia, E. moira, and E. swezeyi from the Micronesian Islands. However, Common (2000) recognized that all Ceylonese and New Guinean species were misplaced in the genus Elaeonoma. Additionally, based on the genital morphological characteristics, Common (1997) and Common (2000) treated all Australian species as other genera or incertae sedis except the type species, E. deltacostamela. However, the three Micronesian species were not mentioned in either study.

Elaeonoma ultima sp. nov.

https://zoobank.org/21C7E95C-0E8D-4B10-BAB8-027C62927C57

Figs 1, 2, 3, 4, 5

Japanese name.

munin-hagane-maruha-kibaga.

Diagnosis.

Elaeonoma ultima sp. nov. is distinguished from other Elaeonoma species sensu Clarke (1984) and Common (2000) by the following characteristics: in the forewing, the cuneiform dark spot extending from 1/2 of the costal margin is absent; the ground colour is not entirely blackish brown; in the cell, the plical spot is present and is located at approximately the same level as the discal spot, but never basal to it. Male genitalia are similar to those of E. swezeyi with an pointed triangular uncus and a narrow dorsal and ventral apex of the valva. However, E. ultima sp. nov. is distinguished by the dorsal margin of the valva, which is convex at 2/3 of the costal margin (nearly straight to slightly concave in E. swezeyi). The female genitalia are similar to those of E. swezeyi with a long apophysis posterioris and lacking a signum. However, E. ultima sp. nov. is distinguished by its small weakly sclerotized lamella postvaginalis (strongly sclerotized plate in E. swezeyi).

Description.

Adult (Figs 1, 2, 3). Sexual dimorphism. Forewing length male 4.4–7.3 mm (n = 28; holotype: 5.0 mm), female 4.7–9.4 mm (n = 17). Wingspan male 9.6–16.0 mm (n = 28; holotype: 11.2 mm), female 10.8–21.0 mm (n = 17).

Figure 1.

Adults of Elaeonoma ultima sp. nov. from the Ogasawara Islands. A. Male, holotype, specimens no. ST580; B. Female, paratype, specimens no. ST675; C. Male, variation, specimens no. ST581; D. Female, variation, specimens no. ST674. Scale bars: 2.5 mm.

Figure 2.

Head of Elaeonoma ultima sp. nov. A. Male, labial palpus, lateral view, holotype, specimens no. ST580; B. Female, labial palpus, lateral view, paratype, specimens no. ST675; C. Male, proboscis, paratype, specimens no. ST796; D. Female, proboscis, paratype, specimens no. ST675. Red triangles point proboscis.

Figure 3.

Male wing venation of Elaeonoma ultima sp. nov., paratype, specimens no. ST586. Scale bar: 5.0 mm.

Male (Fig. 1A, C). Head: Frons and vertex yellowish brown tinged with dark brown. Labial palpus with second palpomere dark brown tinged with yellowish brown outer surface, yellowish brown inner surface; third palpomere dark brown tinged with yellowish brown. Antenna with scape yellowish brown ventrally; flagellomere dark brown; bipectinate trichoid sensillum, twice length of flagellomere. Proboscis very short.

Thorax: Fuscous with dark brown with yellowish brown scales dorsally. Tegula greyish brown. Fore and mid coxa to tibia dark brown with cream at middle and apically; fore- and mid-tarsomere dark brown basally and dorsally, cream ventroapically. Hind coxa to hind tibia dark brown tinged with cream; hind-tarsomere dark brown basally and dorsally, cream ventroapically. Forewing (Figs 1A, 3) lanceolate, ground color dark brown, tinged with yellowish brown; base to basal 2/5 yellowish brown tinged with dark brown; basal 2/5 dark brown band from costa to dorsum; discal and plical oval spots dark brown at basal 2/5; discocellular oval spot at basal 2/3; terminal dots dark brown along termen. Forewing venation with Sc, R1–3, R4+5, M1–3, CuA1–2, CuP and 1A+2A present; R1 from basal 2/3 of discal cell to basal 5/7 of costa; R4+R5 stalked from basal 1/2; R5 reached to apex; CuA1 and CuA2 not stalked; 1A+2A fused from basal 1/3 to end. Hindwing (Figs 1A, 3) dark brown, basal 1/3 greyish brown; fringe greyish brown. Hindwing venation with Sc, Rs, M1–3, CuA1–2, CuP, 1A+2A and 3A present; frenulum a single slender bristle.

Abdomen (Fig. 4A, B): Apodeme very short. Venula straight. Tergum deciduous spines present.

Figure 4.

Abdomen of Elaeonoma ultima sp. nov. A. Male, holotype, specimens no. ST580; B. Male, Hahajima island, specimens no. ST581; C. Female, paratype, specimens no. ST675; D. Female, Hahajima island, specimens no. ST674.

Genitalia (Fig. 5A, B): Symmetrical. Uncus triangular, apex acute and pointed. Gnathos fused, beak-shaped, twice length of uncus with denticulate protrusions dorsally; apex pointed with falcate acute projection. Tegumen trapezoidal, broad medially, lateral arm narrowed to blunt apex. Valva trapezoidal, parallel to basal 2/3, narrowed at apical 1/3; costa straight with setae; apex attenuate and blunt; sacculus slightly narrowed apically with setae; fusiform appendages from ventrobasal part of valva to middle costa, appendages slightly wrinkled and setose. Saccus semicircular, strongly sclerotized, 2/3 as long as uncus. Juxta rectangular, plate-like, sclerotized, bilobed basally, as long as uncus. Phallus sclerotized, 3/4 as long as valva, curved ventrally, widen to base; cornutus straight spine, 2/3 as long as phallus.

Figure 5.

Genitalia of Elaeonoma ultima sp. nov. A. Male, holotype, specimens no. ST580; B. Male, Hahajima island, specimens no. ST581; C. Female, paratype, specimens no. ST675; D. Female, Hahajima island, specimens no. ST674. Scale bars: 1.0 mm.

Female (Fig. 1B, D). Head: Frons and vertex greyish yellow tinged with brown. Labial palpus with second palpomere dark brown, tinged with cream outer surface, cream inner surface; third palpomere cream, tinged with dark brown. Antenna with scape greyish yellow, tinged with dark brown ventrally; flagellomere greyish yellow; sensillum very short. Proboscis very short.

Thorax: Fuscous with greyish yellow, with greyish brown scales dorsally. Tegula greyish brown. Fore and mid coxa to tibia dark brown with cream at middle and apically; fore- and mid-tarsomere dark brown basally and dorsally, cream ventroapically. Hind coxa to hind tibia cream tinged with dark brown; hind-tarsomere light brown basally and dorsally, cream ventroapically. Forewing (Fig. 1B) lanceolate, ground color greyish yellow, tinged with dark brown; base to basal 2/5 greyish yellow tinged with dark brown, costa dark brown; at basal 2/5 wide, dark brown band from costa to dorsum; discal and plical oval spots dark brown, distinct at basal 2/5; discocellular oval spot at basal 2/3; terminal dots dark brown along termen. Hindwing (Fig. 1B) greyish brown, basal 1/3 whitish brown; fringe greyish brown.

Abdomen (Fig. 4C, D): Apodeme straight, slightly shorter than venula. Venula straight. Tergum deciduous spines present.

Genitalia (Fig. 5C, D): Papilla analis setose, elongated, rounded on posterior margin. Apophyses anteriores 5/8 length of apophyses posteriores. Eighth tergum sclerotized, trapezoidal, narrowed posteriorly, anterior middle convex, baculiform process elongated. Ostium bursae membranous. Antrum conical, short, narrowed anteriorly. Ductus seminalis elongated from anterior end of antrum. Ductus bursae membranous, slender, 2/3 length of apophyses posteriores, slightly widened to corpus bursae. Corpus bursae ovate, membranous, poorly developed, and small. Signum absent.

Variation: Slight and consistent differences were observed between the Chichijima (type locality) and Hahajima populations; these are treated as intraspecific variation and summarised below.

Male (Figs 1C, 4B, 5B). Forewing ground colour is often entirely dark brown in Hahajima populations. The abdominal apodeme of Hahajima populations is slightly longer than that of Chichijima populations. In the genitalia, the uncus of Hahajima populations is more obtuse triangular and wider at base than that of Chichijima populations; the attenuate apex of valva of Hahajima populations is slightly more elongated than that of Chichijima populations.

Female (Figs 1D, 4D, 5D). The costal base to basal 1/3 of forewing is often dark brown in Hahajima populations. The abdominal apodeme of Hahajima populations is slightly curved apically. In the genitalia, the anterior middle of eighth tergum is widened at base.

Type material.

Holotype. • ♂; [JPN: Tokyo-Pref.] Shigureyama, Chichijima-Is., Ogasawara-mura; 27.0629°N, 142.2214°E; alt. 252 m; 26.VI.2022; LT; S. Tomura leg.; specimens no. and sample ID for DNA analysis: ST580; deposited in ELKU.

Paratypes. [Chichijima Island] • 1♀, same locality, date, method and collector; specimens no. and sample ID: ST583; ELKU•1♀, same locality, date and method; S. Yagi, S. Tomura, M. Kimura leg.; ELKU•1♂3♀, same locality and method; 18.VI.2022; S. Tomura leg.; specimens no. and sample ID: ST678, ST679; ELKU•1♂, same locality date and method; T. Hirowatari, S. Yagi, S. Tomura, I. Ohshima, M. Kimura leg.; specimens no.: ST796; ELKU•2♂, same locality and method; 11.VI.2023; S. Yagi leg.; specimens no. ST726; ELKU•1♀, same locality, method and date; T. Hirowatari, S. Yagi, Y. Matsui, J.-H. Park, I. Kawashima, J. Hamaguchi, M. Kimura leg.; specimens no. and sample ID: ST758•1♂, Asahiyama, Ogasawara-mura; 27.0954°N, 142.2135°E; alt. 206 m; 26.VI.2022; SW; S. Tomura leg.; specimens no. and sample ID: ST677; ELKU•2♂, same locality and method; 18.VI.2022; S. Yagi leg.; specimens no. ST782; ELKU•2♂, same locality, date and method; T. Hirowatari leg.; specimens no. ST780; ELKU•1♀, Higashidaira, Ogasawara-mura; 27.0749°N, 142.2216°E; alt. 235 m; 18.VI.2022; S. Tomura leg.; specimens no. ST588; ELKU•1♂, same locality, date and method; T. Hirowatari leg.; specimens no. and sample ID: ST676; ELKU•1♂, same locality and method; 27.0739°N, 142.2223°E; alt. 239 m; 26.VI.2022; S. Tomura leg.; specimens no. ST586; ELKU•3♂, Mt. Chûô-san, Ogasawara Vill.; 19.VI.2023; SW; J.-H. Park leg.; specimens no. and sample ID: ST781; ELKU•3♂3♀, Kuwanokiyama, Ogasawara-mura; 27.0777°N, 142.2174°E; alt. 216 m; 19.VI.2022; LT; S. Tomura leg.; specimens no. and sample ID: ST582, ST587, ST675, ST795; ELKU•1♀, same locality, date and method; S. Yagi leg.; specimens no. ST779; ELKU.

Other materials. [Hahajima Island]•1♀, Sakaigatake, Ogasawara-mura; 26.6705°N, 142.1559°E; alt. 411 m; 22.VI.2022; S. Tomura leg.; specimens no. ST674; ELKU•1♂, same locality, 26.671°N, 142.155°E; 15.VII.2024; flying just after sunset; Y. Matsui leg.; ELKU•1♂, Kuwanokiyama; 26.6632°N, 142.1530°E; 13.VI.2025; LT; S. Yagi leg.; specimens no. and sample ID: ST784; ELKU•1♂, Mt. Chôki-yama, Ogasawara-mura; 26.6509°N, 142.1568°E; alt. 200 m; 15.VI.2023; LT; S. Yagi leg.; ELKU•1♀, same locality, date and method; Y. Matsui leg.; specimens no. and sample ID: ST786; ELKU•2♂, same locality, date and method; T. Hirowatari, S. Yagi, Y. Matsui, J-H. Park, I. Kawashima, J. Hamaguchi, M. Kimura leg.; ELKU•1♂, Funakiyama, Ogasawara-mura; 26.6511°N, 142.1692°E; alt. 271 m; 24.VI.2022; S. Tomura leg.; specimens no. and sample ID: ST581; ELKU•1♂, Funamidai, Ogasawara-mura; 21.VI.2022; S. Tomura leg.; specimens no.: ST673; ELKU•4♀, Nishiura; 26.6557°N, 142.1531°E; 10.VI.2025; LT; S. Yagi leg.; specimens no. and sample ID: ST785; ELKU•1♂, same locality, method and collector; 15.VI.2025; specimens no. and sample ID: ST783; ELKU•1♂, Minamizaki; 26.62°N, 142.18°E; 21.IX.2023; daytime search; Y. Matsui leg.; specimens no. ST757; ELKU.

Biology.

(Figs 6, 7) Adults of the new species were collected in June, July, and September, but not in March; therefore, this species is presumed to be uni- or bivoltine. During the day, adults were often collected by sweeping around mossed barks and bases of trees in the shade. At night, adults were collected using a light trap in the forest. When the collected adults were provided with cotton wool soaked in sugar water, they sucked the water by leaning forward and pressing their short proboscis against the surface. When the collected female adults were placed in a plastic case with dead leaves and bark covered with moss, they laid eggs in gaps between the bark and endemic moss, Leucobryum boninense Sull. & Lesq. Approximately two weeks later, the larvae hatched, spun thread and built a rough nest. A small amount of feces was also found, suggesting that this species feeds on moss and bark.

Figure 6.

Habitat and biology of Elaeonoma ultima sp. nov. A-B. Collecting sites of adults; C. Male adult attracted by light; D. Female adult attracted by light; E. Rearing adult with bark and dead leaf in a plastic case; F. Adult sucking sugar water; G. Female adult laying eggs into a gap in the bark; H. Egg on endemic moss, Leucobryum boninense Sull. & Lesq.

Figure 7.

Habitat and biology of Elaeonoma ultima sp. nov. A. Egg on moss; B. Hatched larva on endemic moss, Leucobryum boninense Sull. & Lesq.

Distribution.

(Fig. 8) Japan: Ogasawara Islands (Chichijima and Hahajima Islands).

Figure 8.

Distribution map of all known Elaeonoma species.

DNA data.

(Figs 9, 10, Table 1, Suppl. material 1) The DNA barcode sequences are deposited in the GenBank database under the accession numbers listed in Table 1 and uploaded to the BOLD system in the public dataset DS-OGAELAEO. The intra-island p-distances of E. ultima sp. nov. on Chichijima Is. are 0–3.33% (n = 11), those on Hahajima Is. are 0–2.48% (n = 6), and inter-island p-distances of E. ultima sp. nov. between the two islands are 4.29–6.03% (n = 17).

Figure 9.

Maximum likelihood tree of Elaeonoma ultima sp. nov. and related species based on the DNA barcode region. The labels provide data on Sample ID, species name, and island of collection. The number near the node is the bootstrap value. Branch lengths are proportional to genetic distances, and the scale bar indicates 0.2 substitutions/site.

Figure 10.

Haplotype network of Elaeonoma ultima sp. nov. constructed based on the COI barcode region. The sizes of the circles are proportionate to the numbers of specimens sharing a particular haplotype. The arrow marks and percentage values refer to the genetic distances between the haplotype groups.

Table 1.

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The list of Japanese specimens of Elaeonoma ultima sp. nov. used for DNA analysis.

Species name	Sample ID	sex	location	collecting date	GenBank accession number	BIN	primer sets for PCR
Elaeonoma ultima sp. nov.	ST781	male	Ogasawara-mura, Chichijima Is., Mt. Chûô-san	19.VI.2023	PZ539381	BOLD:AHN1451	LepF1/LepR1
Elaeonoma ultima sp. nov.	ST580	male	Ogasawara-mura, Chichijima Is., Shigureyama	26.VI.2022	PZ539382	BOLD:AHN1451	LepF1/LepR1
Elaeonoma ultima sp. nov.	ST677	male	Ogasawara-mura Chichijima Is., Asahiyama	26.VI.2022	PZ539383	BOLD:AHN1451	LepF1/LepR1
Elaeonoma ultima sp. nov.	ST676	male	Ogasawara-mura Chichijima Is., Higashidaira	18.VI.2022	PZ539384	BOLD:AHN1451	LepF1/LepR1
Elaeonoma ultima sp. nov.	ST673	male	Ogasawara-mura, Hahajima Is., Funamidai	21.VI.2022	PZ539385	BOLD:AHM4183	LepF1/LepR1
Elaeonoma ultima sp. nov.	ST783	male	Ogasawara-mura, Hahajima Is., Nishiura	15.VI.2025	PZ539386	BOLD:AHM4183	LepF1/LepR1
Elaeonoma ultima sp. nov.	ST784	male	Ogasawara-mura, Hahajima Is., Kuwanokiyama	13.VI.2025	PZ539387	BOLD:AHM4183	LepF1/LepR1
Elaeonoma ultima sp. nov.	ST581	male	Ogasawara-mura, Hahajima Is., Funakiyama	24.VI.2022	PZ539388	BOLD:AHM4183	LepF1/LepR1
Elaeonoma ultima sp. nov.	ST583	female	Ogasawara-mura, Chichijima Is., Shigureyama	26.VI.2022	PZ539389	BOLD:AHN1451	LepF1/LepR1
Elaeonoma ultima sp. nov.	ST582	female	Ogasawara-mura, Chichijima Is., Kuwanokiyama	19.VI.2022	PZ539390	BOLD:AHN1451	LepF1/LepR1
Elaeonoma ultima sp. nov.	ST679	female	Ogasawara-mura, Chichijima Is., Shigureyama	18.VI.2022	PZ539391	BOLD:AHN1451	LepF1/LepR1
Elaeonoma ultima sp. nov.	ST678	female	Ogasawara-mura, Chichijima Is., Shigureyama	18.VI.2022	PZ539392	BOLD:AHN1451	LepF1/LepR1
Elaeonoma ultima sp. nov.	ST758	female	Ogasawara-mura, Chichijima Is., Shigureyama	11.VI.2023	PZ539393	BOLD:AHN1452	LepF1/LepR1
Elaeonoma ultima sp. nov.	ST675	female	Ogasawara-mura, Chichijima Is., Kuwanokiyama	19.VI.2022	PZ539394	BOLD:AHN1452	LepF1/LepR1
Elaeonoma ultima sp. nov.	ST786	female	Ogasawara-mura, Hahajima Is., Mt. Chôki-yama	15.VI.2023	PZ539395	BOLD:AHM4184	LepF1/LepR1
Elaeonoma ultima sp. nov.	ST780	male	Ogasawara-mura Chichijima Is., Asahiyama	18.VI.2022	PZ539396	BOLD:AHN7240	LepF1/LepR1
Elaeonoma ultima sp. nov.	ST785	female	Ogasawara-mura, Hahajima Is., Nishiura	10.VI.2025	PZ539397	BOLD:AHM4184	LepF1/LepR1

The DNA barcode of E. ultima sp. nov. (Sample ID: ST673) is closest to that of E. adeneia (BIN, BOLD:ADZ8644, Sample ID: CCDB-30823-C06, Process ID: LNAUW4108-18), according to the BOLD identification engine, with 8.73% pairwise divergence.

According to the phylogenetic tree constructed based on the COI barcode region, E. ultima was recovered as sister to E. adeneia, although the bootstrap support value was low (59%). The Chichijima population of E. ultima sp. nov. formed a monophyletic clade (bootstrap value = 84%). In contrast, the Hahajima specimens were not monophyletic and appeared as two lineages outside the Chichijima clade. According to the haplotype network constructed based on the COI barcode region, the Chichijima population of E. ultima is clearly separated from that of the Hahajima population. An association between the collection site and haplotypes within each island was not clearly observed.

Etymology.

The species epithet ultima is derived from the Latin word “ultimus” meaning “furthest.” This name reflects that the type locality of the Ogasawara Islands, which are located at the eastern end of Japan and are the outermost limit of the known distribution range of this genus.

Remarks.

Kato (1992) recorded two species of “Oecophoridae gen. sp.” from Chichijima and Anijima Islands. However, these were the unidentified species of the family Stathmopodidae (Kato, personal communication). It is presumed that this is due to the historical taxonomic treatments in which stathmopodid moths were regarded as a subfamily “Stathmopodinae” within Oecophoridae until recent years (e.g., Hodges 1998; Wang 2006). Recently, Stathmopodidae is clearly distinguished from Oecophoridae based on the morphological characteristics and molecular phylogenetic analysis (Heikkilä et al. 2014). According to the current classification, this study is the first confirmed record of the Oecophoridae from the Ogasawara Islands. Clarke (1984) mentioned that the Syûti Issiki’s specimen collection of the Smithsonian Museum contains some Gelechioidea of northern Micronesian Islands, but specimens and associated data from the Bonin (= Ogasawara) Islands were not confirmed.

Common (1997, 2000) treated most Australian Elaeonoma species as incertae sedis or the following other genera: Barea Walker, 1864, Chezala Walker, 1864, Diaphanta Common, 2000, Eulechria Meyrick, 1883, Ioptera Common, 1997, Ischnomorpha Common, 1996, Leimmatonca Common, 2000, Leipochlida Common, 2000, Micramicta Common, 2000, Neosigala Common, 2000, Orescoa Turner, 1927, Stenoptena Common, 2000, and Tanycaula Common, 1997. This new species is distinguished from these genera by the following morphological characteristics: short proboscis; bifid scale-crests on the thorax and forewing are absent; the forewing is not narrowly elongated and veins CuA1 and CuA2 are not strongly curved; the uncus is defined and rostral; the gnathos is not deeply concave dorsally, not strongly sclerotized, and does not have an inflated apex; the apex of the valva does not have a curved projection; the sacculus does not have an up-curved process and apex does not have a sclerotized distal process; apophysis posterior is not extremely long; and ductus bursae is not sclerotized.

Discussion

Systematic position of Elaeonoma ultima sp. nov.

Common (2000) included the genus Elaeonoma in the Barea genus group and mentioned “fusiform appendages” of the male genitalia as a distinctive character of this group. This characteristic was also observed in E. ultima sp. nov. and Micronesian species, which suggests that these species belong to the same species group. Moreover, Common (1994, 2000) treated most species recently assigned to Elaeonoma as belonging to different genera based on their morphological characteristics. This new species was not assigned to any of these genera but agree with the diagnostic characteristics of Elaeonoma. However, some morphological differences were observed between the type species, E. deltacostamela of Australia and Elaeonoma species of the Micronesian Islands. For example, the forewing veins CuA1 and CuA2 are stalked/not stalked; the uncus is rostral/triangular; the apophyses anteriores are very short/long, respectively. Morphological similarities indicate that E. ultima sp. nov. is closely related to the Micronesian species. In the present study, we tentatively placed them in Elaeonoma, but the four species from the Micronesia and Ogasawara Islands need to be reexamined in the future.

Based on the fact that the apex of the valva in the male genital structure is acute, E. ultima sp. nov. is considered to be most closely related to E. swezeyi from Guam Island, which is the closest to the Ogasawara Islands among the four Micronesian species. In the phylogenetic tree, E. adeneia has been shown to be closely related; however this is presumably due to incomplete lineage sorting.

Geographic patterns and morphological variation

Oceanic islands often promote allopatric speciation through inter-island dispersal followed by geographic isolation. In Hawaii, which is also oceanic islands, the endemic snail-killer moth Hyposmocoma (Cosmopterigidae) migrated to each island resulting in allopatric speciation due to geographic isolation (Rubinoff 2008). In Marquesas Islands, flightless weevil Rhyncogonus species (Coleoptera, Curculionidae) and some snails and plants dispersed and speciated, and they have been discussed in the context of the stepping-stone hypothesis (Hembry 2018). The distribution range of the genus Elaeonoma includes the Oceanian and Australasian Regions, and not the Palaearctic and Oriental Regions. Therefore, the ancestor of this new species is assumed to have migrated northward to the Ogasawara Islands from the Oceanian Islands. The route of the arrival of the ancestors of Elaeonoma is unclear. It is presumed to be dispersed by the stepping-stone dispersal rather than the radial dispersal scenario, judging from the morphological similarities between species from geographically close islands, including the pairs E. ultima sp. nov. and E. swezeyi, and E. adeneia and E. moira. However, this hypothesis requires verification of the phylogenetic relationships within the genus based on additional molecular markers.

In the Ogasawara Islands, genetic distances between island populations have been confirmed for some moth species, even when clear morphological differentiation was not confirmed (Yagi et al. 2024; Park et al. 2025). In E. ultima sp. nov. as well, molecular analyses supported the inter-island differentiation. Additionally, according to the abdominal and male genital morphological characteristics, slight but consistent differences were observed between the Chichijima and Hahajima populations. These results suggested that E. ultima sp. nov. is genetically and morphologically isolated between the two islands and is undergoing allopatric speciation on the two islands. In several Gelechioidea species, a high intraspecific DNA divergence in the COI barcode region (approximately 2–4%) accompanied by slight morphological variations has been reported (Kim et al. 2020). E. ultima sp. nov. exhibits large genetic variations, both inter- and intra-island. Given its allopatric distribution and the presence of both morphological and mitochondrial divergence, the inter-island variation of this species appears to be broadly consistent with the subspecies framework proposed by Braby et al. (2012). However, discordance between mitochondrial and nuclear markers has been reported (Hinojosa et al. 2019), indicating that the current DNA-based evidence remains insufficient. Furthermore, intra-island genetic diversity is high and two genetically distinct clades emerged on Hahajima; however, no corresponding morphological differences were observed. Therefore, we avoided separating species and subspecies solely on inter-island basis before accounting for genetic and morphological variations more precisely.

Habitats and potential effects of invasive alien predators

The Ogasawara Islands contain several types of forests with different tree heights, precipitation, humidity, and other conditions, and their species compositions and ecosystems differ (Yoshida et al. 2002; Shimizu 2003). Adults of E. ultima sp. nov. were found in a variety of environments, including the subtropical dry scrub of Chichijima, the moist forest of Hahajima, and the coastal forests. However, in Chichijima, adults were not confirmed on Mt. Ogami-yama and Mikazuki-yama, which are near extremely dry settlements. Therefore, such environments are presumed to be unsuitable for habitation.

On rearing, E. ultima sp. nov. lays eggs on moss and bark, and in the wild, adults have been collected in dimly lit areas with mossy bark near the ground, so it is likely that the larvae prefer such environments in the wild as well. Considering that E. ultima sp. nov. lays their eggs on bark in multiple conditions and that the congener E. swezeyi emerges from the dead leaves of herbaceous plants (Clarke 1984), it is likely that their host preference is not very specific. Another moth known to live in a similar habitat on the Ogasawara Islands is the bryophyte-feeder endemic moth, Glaucocharis triocellaris (Crambidae) (Matsui et al. 2024). Some adult moths, including the detrivore tineid moth Erechthias species, were collected from gaps in the dead leaves of Livistona chinensis var. boninensis Becc. (Park et al. 2025), but Glaucocharis triocellaris, which is similar to E. ultima sp. nov., was collected by sweeping the forest floor and rarely collected by sweeping the leaves of L. chinensis. On the other hand, Erechthias, whose larvae feed on detritus and associated fungi and have a feeding habit similar to that of Oecophoridae, was frequently collected by sweeping the leaves of L. chinensis and rarely by sweeping the forest floor. Thus, the habitat of this new species is estimated to be closer to that of Glaucocharis, which inhabits moist environments with mosses, than to that of Erechthias, which lives in dry decaying plant matter.

Many adults have been confirmed on the Chichijima Island, and the population size is thought to be large. In contrast, fewer individuals were obtained from Hahajima, despite the generally more mesic forest environment of Hahajima compared to the drier vegetation of Chichijima (Shimizu 2003). This pattern suggests that island-scale humidity alone may not determine local abundance. Furthermore, given the genetic differentiation observed between populations on the two islands, it is possible that ecological differentiation has also occurred, potentially leading to differences in habitat use, microhabitat preference, or life-history traits between the islands. However, these differences require further investigation.

Elaeonoma ultima sp. nov. is only confirmed in the largest islands (Chichijima and Hahajima Islands) and has not been found in many surveys on other islands (Mukojima Island, Anijima Island, or Meijima Island). The reason that it is confirmed only on these two islands may be explained by the law of “island species-area relationships,” which states that the larger the island, the greater the number of species that inhabit it (MacArthur and Wilson 1963). Rather than arriving by chance on only two islands, it is more likely that this species has failed to establish itself on other drier small islands or has already become extinct.

On the Ogasawara Islands, many alien plants and animals have been introduced and gone wild, causing a variety of adverse effects on the ecosystem (Shimizu 2003). In particular, the predation pressure of generalist predators has a serious impact on small endemic insects and arthropods, such as the green anole, Anolis carolinensis Voigt, 1832 (Reptilia), the toad, Bufo marinus (Linnaeus, 1758) (Amphibia), and the land nemertean, Geonemertes sp. (Nemertea) (Shimizu 2003; Yago 2014; Yoshino 2025). In fact, the Ogasawara Holly Blue, Celastrina ogasawaraensis (Lycaenidae) has not been reported in the wild since 2020, suggesting that the species may already be extinct, with invasive predators such as the green anole considered as a major contributing factor (Nakahama et al. 2024). Many species are known to be extinct from the inhabited islands, where many alien predators live, and are only found on nearby islands where predators do not live (Karube 2014; Hisasue and Hideshima 2025). Because many individuals of this new species were confirmed on inhabited islands where other taxa have become extinct or drastically decreased in number, it is assumed that this species is relatively tolerant to predation pressure. However, as the habitat of this species’ larvae and adults, on or inside the moss, is an area that predators can easily access, it is possible that the current situation has already seen a decline in the population due to predation pressure. These two islands, the only distribution areas for this species, are inhabited by alien predators and there are no refugia on the nearby islands, unlike in some other endemic species. If the impact of these predators increases in the future, the risk of extinction may increase rapidly. Although the extent of predation pressure cannot be quantified because the prey behavior of green anole and other predators on this moth has not been observed, it is necessary to closely monitor the future stability of this species’ population.

Acknowledgements

We sincerely thank Lauri Kaila, Francisco Urra and the anonymous reviewer for their critical comments on this manuscript. We are deeply indebted to Masaaki Kimura (Okinawa Prefecture, Japan), Ikumi Kawashima, Jinhyeong Park, Junpei Hamaguchi (ELKU), Yuki Matsui (National Agriculture and Food Research Organization), Toru Miyamoto (School of Science, University of Tokyo), and Issei Ohshima (Kyoto Prefectural University) for their cooperation in collecting samples, and Johei Oku (Oita Prefecture) for providing references. We are grateful to the Nature Conservation Office of Ogasawara, Ministry of the Environment, Japan, and the Kanto Regional Forest Office, Forestry Agency, Japan, for allowing us to conduct field surveys in the nature reserves of the Ogasawara Islands, Japan.

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Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Artificial Intelligence (AI) use

The authors accept full responsibility for the content of the manuscript, including the disclosure of any use of AI.

No AI tools were used in the preparation of this manuscript.

Funding

This research was supported by the Environment Research and Technology Development Fund (JPMEERF20224201 and JPMEERF20254RA1) of the Environmental Restoration and Conservation Agency, provided by the Ministry of the Environment of Japan to ST, TH, and SY, and to ST, JSPS KAKENHI (Grant numbers JP21J22192 and JP23K19301).

Author contributions

All authors have contributed equally.

Author ORCIDs

S. Tomura https://orcid.org/0000-0002-5413-1220

S. Yagi https://orcid.org/0000-0002-4261-1219

T. Hirowatari https://orcid.org/0000-0002-6839-2229

Data availability

All of the data that support the findings of this study are available in the main text or Supplementary Information.

Supplementary material

Supplementary material 1

Interspecific and Intraspecific pairwise p-distances in the COI barcode region of the genus Elaeonoma

Shunsuke Tomura, Sadahisa Yagi, Toshiya Hirowatari

Data type: xlsx

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

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