Elucidation of the Taxonomy of Three Problematic East Asian Nola Leach, 1815, with Description of a New Species (Lepidoptera, Nolidae, Nolinae)

: The paper discusses the taxonomy of three East Asian Nola Leach, 1815 taxa, N. costimacula Staudinger, 1887, N. japonibia (Strand, 1920) and N. innocua Butler, 1880. We describe a new species N. galliphaga sp. nov . from Japan and South Korea and synonymize N. japonibia syn. nov . with N. costimacula . Type specimens, additional adults, and their genitalia are illustrated.


Introduction
The genus Nola was established by Leach (1815), with the type species Phalaena cucullatella Linnaeus, 1758.The species of the genus are characterised by their small size, usually achromatic forewing, and trifine venation of the hindwing.The male genitalia of the Nola species are rather distinctive, and are characterised by the strongly reduced uncus, the bilobate valva, the usually simple harpe, the short, tubular aedeagus often bearing a large, curved cornutus.In the female genitalia, the ostium bursae is normally membranous, the ductus bursae is tubular, sometimes bulged, and the corpus bursae usually has an invaginated, somewhat thorn-shaped signum [1].
The genus is present in all continents (except for Antarctica) and especially highly diverse in the Indomalayan and Afrotropical biogeographic realms.Due to the vast number of taxar and the large proportion of cryptic species, the knowledge of the taxonomic content of the genus is far from complete and, based on our preliminary studies, a large number of taxa are undescribed, even in the most well-explored regions.In East Asia, 65 Nola species have been recorded to date, including the Russian Far East, Mongolia, China, Hong Kong, Taiwan, Korea, and Japan [2][3][4][5][6][7][8][9][10][11][12].The most comprehensive overview of the genus was published by Inoue [2], whose work has long been used as a basis of further studies on the East Asian Nolidae [3][4][5][12][13][14].However, this important monograph is not completely devoid of errors owing to some unexamined or overlooked historical primary type specimens deposited in European museums.The third and fourth authors of this paper had an opportunity to examine type specimens of East Asian Nola species housed in the NHMUK and MfN and this study allowed to recognise the misidentification of N. innocua by Inoue [2], whose erroneous concept was followed by Kononenko and Han [13] and Tshistjakov [4].László et al. [14] published the genital morphology of true N. innocua Butler, 1880/ for the first time, based on the dissection of the female holotype housed in the NHMUK and further topotypical male and female specimens accessed in the NHMUK and MWM/ZSM.Consequently, the species which had earlier been erroneously identified as N. innocua by Inoue [2] is in fact a species new to science and it is described in this paper as N. galliphaga sp.nov.
The taxonomy of N. japonibia (Strand, 1920) has also been a subject to several misinterpretations.The taxon was originally described as a subspecies of N. innocua, and it has later been treated as valid species by Inoue [15].The examinations of the female holotype of Celama innocua var.japonibia and female specimens of N. costimacula Staudinger, 1887 from various localities have confirmed the identity of the two taxa and thus here as a synonym of the latter: N. japonibia (Strand, 1920) is a junior synonym of N. costimacula Staudinger, 1887.
The taxonomy of N. japonibia (Strand, 1920) has also been a subject to several misinterpretations.The taxon was originally described as a subspecies of N. innocua, and it has later been treated as valid species by Inoue [15].The examinations of the female holotype of Celama innocua var.japonibia and female specimens of N. costimacula Staudinger, 1887 from various localities have confirmed the identity of the two taxa and thus here as a synonym of the latter: N. japonibia (Strand, 1920) is a junior synonym of N. costimacula Staudinger, 1887.

Material and Methods
Specimens examined are preserved in the collections of the Hungarian Natural History Museum, Budapest, Hungary (HNHM), the Institute of Systematics and Evolution of Animals, Krakow, Poland (ISEZ), the Korean National Arboretum, Pocheon, Korea (KNAE), the Museum of Natural History, Berlin, Germany (MfN), the Museum Witt in the Bavarian State Collection of Zoology, Munich, Germany (MWM/ZSM), and the Natural History Museum, London, United Kingdom (NHMUK).Genitalia were dissected and examined using either a Leica EZ4 or a Nikon SMZ 745T stereomicroscope.Images of INU adults were taken by a Tucsen Dhyana 400DC digital camera attached to a Leica S6D stereomicroscope or Canon EOS 6D Mark II camera equipped with a canon Macro 100mm lens, with dome illuminator Leica LED5000 HDI.Photos of MWM/ZSM and NHMUK adults were taken using a Nikon D90 DSLR camera equipped with a Nikkor AF Micro 60 mm lens.Genitalia photographs were taken using either a Tucsen Dhyana 400DC digital camera mounted on a Leica S8AP0 stereomicroscope or a Nikon Eclipse 80i compound microscope connected to a Nikon DS-Fi1 digital camera.
The holotype label data have been transcribed verbatim in quotation marks and with "/" denoting a different label.
Host plant.Unknown.Remarks.This species was firstly reported from Korea by Oh [3] as N. japonibia.Kim et al. [18] reported N. costimacula in Korea without any examined materials or references.However, as a result of this present study, we found that N. japonibia is a junior synonym of N. costimacula based on the examination of type materials of both species (Figure 1a,b), therefore the name N. japonibia is to be replaced by N. costimacula in the check list of the Korean Nolinae.Tshistjakov [4] incorrectly referred costimacula as a subspecies of N. innocua and listed Taiwan as one of the countries where the taxon is distributed, mentioning, however, that only the nominotypical subspecies occurs in the island.In his erroneous taxonomic concept, it meant that N. innocua innocua occurs in Taiwan, which is correct, but mentioning it under the taxon costimacula is rather confusing as the latter is not a member of the Taiwanese fauna.
Distribution.Taiwan (endemic) [9].Host plant.Unknown.Remarks.Nola innocua was reported from Korea by Inoue [14]; however, this record proved to be based on a misidentification.The examinations of the holotype of N. innocua and further topotypical male and female specimens externally matching with the female primary type have revealed that ever since the publication of Moths of Japan [2], this species has been misidentified in the Far East.Consequently, the taxon which has long been treated as N. innocua is in fact an undescribed species, the description of which is given in this paper.Diagnosis.This new species is reminiscent of both N. innocua and N. costimacula, but it is distinguished by its markedly thicker and more evenly arched antemedial line, the smaller and more triangular medio-costal patch, the more sharply defined and more angled postmedial line of the forewing, and the small, rounded discal spot of the hindwing, which is dash-like in N. costimacula and absent in N. innocua.In addition, compared to N. innocua, N. galliphaga sp.nov. is considerably larger (the wingspan of the former species is 12-13 mm, that of the new species is 16-19 mm), while its ground color is darker than that of N. costimacula.Nevertheless, examination of the copulatory organ of these externally similar species provides the most reliable identification.Based on the configuration of the male genitalia, N. innocua belongs to a distinct lineage of Nola due to the highly modified, medially stalked, apically spatulate dorsal valval lobe and the massive, claw-like cornutus of the vesica excluding potential misidentification with any other Nola species.Compared the new species to N. costimacula, the differences between the two species' genital capsule are more subtle expressed by the somewhat shorter tegumen and the basally broader, more curved harpe of N. galliphaga.However, the configuration of the aedeagus is strikingly different in the two species, being slightly longer and narrower with a thin, strongly curved cornutus in N. galliphaga, while the cornutus of N. costimacula is markedly more robust and almost straight.In the female genitalia, the new species has a more heavily sclerotized, longer antrum, markedly shorter and narrower ductus bursae lacking a bulge, a larger, more elongate corpus bursae with more pronounced cervix, and a larger medial process of the thorn-shaped signum compared to those of N. costimacula.
Distribution.Korea, Japan.Host plant.The species-identified formerly erroneously as N. innocua-was observed feeding on the galls of Nipponaphis distyliicola Monzen and Monzenia globuli (Monzen) aphids (Hemiptera), on Distylium racemosum Siebold and Zuccarini (Hamamelidaceae) as well as on the leaves of the same plant [21].Two specimens reared from galls were traced in the Inoue collection housed in the NHMUK and their genital slides have proved their correct identity as N. galliphaga.
Remarks.Inoue [14] reported a single Korean female specimen from Andong, Gyeong--sangbuk-do province without providing exact label data.Purportedly, this specimen might have been dissected by Oh [3], but the specimen itself seems to have been lost; only its genitalia slide has been traced.Nevertheless, the genital preparation shows characters identical to other female specimens of the new species collected in Japan.Consequently, the single genitalia slide extant in KNAE is recognized here as a paratype specimen.The Korean population of the species is supposedly also connected to D. racemosum and the galls of the two aphids as reported by Ito and Hattori [21].Although these latter authors did not illustrate the moths, the result of present taxonomic study allowed us to infer that the species Ito and Hattori reported as N. innocua is undoubtedly N. galliphaga, as true N. innocua Butler is a species endemic to Taiwan.
Etymology.This species is named based on the peculiar feeding habit of the larva, making the new taxon a potential natural enemy of gall aphids.

Discussion
The detailed morphological analyses of the primary types and additional specimens of three East-Asian Nola species have shed light on long-standing taxonomic misinterpretations of Nola costimacula, N. innocua and N. japonibia.The species, which has long been treated as N. innocua, has proved to be an undescribed species, described here as N. galliphaga, showing a closer kinship to N. costimacula.The true Nola innocua has highly distinctive genitalia features without any known closer relative and has hitherto been only recorded from Taiwan suggesting that the species is endemic to the island.The new species has been reported to feed on galls of two aphid species (Nipponaphis distyliicola Monzen and Monzenia globuli (Monzen)), and also on their host plant Distylium race-

Discussion
The detailed morphological analyses of the primary types and additional specimens of three East-Asian Nola species have shed light on long-standing taxonomic misinterpretations of Nola costimacula, N. innocua and N. japonibia.The species, which has long been treated as N. innocua, has proved to be an undescribed species, described here as N. galliphaga, showing a closer kinship to N. costimacula.The true Nola innocua has highly distinctive genitalia features without any known closer relative and has hitherto been only recorded from Taiwan suggesting that the species is endemic to the island.The new species has been reported to feed on galls of two aphid species (Nipponaphis distyliicola Monzen and Monzenia globuli (Monzen)), and also on their host plant Distylium racemosum Siebold and Zuccarini.Consequently, this new species could potentially be a natural enemy of some aphid pests in forests.Nevertheless, those full of morphologic evidence, the additional molecular study will need in near future.Many recent molecular studies revealed numerous cryptic and misidentified species.In this reason, molecular study for those of this species, and also other nolid species are necessary.However, in this study, the authors focused on the morphologies, that those species have noticeable characteristics.

Conclusions
The results of this study demonstrate that in-depth revisional research is still essential to clarify the identity and distribution of species of poorly studied Lepidoptera groups such as Nolinae.This current study clarified the erroneous taxonomy of three East Asian Nola taxa resulting in the discovery of a new species occurring in Japan and Korea.Similar studies are still required to elucidate further taxonomic problems in the Nolinae of the Far East and even discoveries of further new species cannot be ruled out.
It would also be important to obtain further information of the biology of these predominantly arboreal species which could bring surprising discoveries regarding their potential economic utilization.In our study, the newly described Nola species has been revealed as a potential natural enemy of gall-inducing aphids as it was reported by Ito and Hattori [21] although they attributed the observed behavior to the misidentified N. innocua.Behavioral studies and breeding experiments addressed to the early stages of noline moths may reveal further species with similar tendencies, rendering them potential natural enemies of forest pests.