A new species of Homalernis Meyrick, 1908 (Lepidoptera, Tortricidae, Tortricinae) represents the first record of the tribe Schoenotenini in Japan

. Homalernis fluctuosa Suzuki & Jinbo, sp. nov. , is described and illustrated from Honshu, Shikoku, Kyushu, Tsushima Island, Amamiooshima Island, and Okinawajima Island, Japan. This is not only the first re - cord of the genus Homalernis but also of the tribe Schoenotenini from warm temperate zones in the Palaearc-tic region. The association of males and females of the new species was confirmed based on the mitochondrial gene cytochrome oxidase submit 1 ( COI ). We discuss the taxonomic positions of two alleged Homalernis species from Malaysia and the taxonomic position of Homalernis within Schoenotenini.

Morphologically, Schoenotenini are considered a monophyletic group based on the following forewing characteristics: an M-stem vein that terminates between M 1 and M 2 , an almost equal arrangement of radial veins, and often with raised scales (Common 1965;Dugdale 1966;Horak 1998;Razowski 2008).In a molecular phylogenetic analysis of the Tortricidae, Schoenotenini were placed as a clade with Epitymbiini + Archipini or Phricanthini; however, neither hypothesis was well supported (Fagua et al. 2017).Meyrick (1908) described Homalernis semaphora Meyrick, 1908 from Assam (India), designating it as the type species of his new genus, which he assigned to Tortricinae.A decade later, Meyrick (1918) described a second species of the genus, H. arystis, also from Assam (India).Diakonoff (1939) initially placed Homalernis in the family Eucosmidae (currently regarded as Eucosmini of Olethreutinae).Subsequently, Diakonoff (1960) included the genus in the Schoenotenini without discussion, but he mentioned that the female genitalia closely resembled those of Metachorista, another member of this tribe.Common (1965) included Homalernis in Schoenotenini, remarking that it was closely related to the Proselena group of genera based on similarities of the female genitalia.
Currently, there are four described species assigned to Homalernis: H. arystis Meyrick, 1918, H. jeriau Razowski, 2012, H. mankoboi Razowski, 2012, and H. semaphora Meyrick, 1908(Gilligan et al. 2018).The type species of this genus, H. semaphora, was described allegedly based on a single male and a single female; however, Diakonoff (1939) found that the specimen reported as male was a female.Meyrick (1918) also described H. arystis based on a female specimen.In contrast, Razowski (2012) described both H. jeriau and H. mankoboi based on single males each and placed these two species in Homalernis based on similarities in appearance and wing venation.Therefore, no species in this genus has been described from both sexes, and the placement of the two species described based on only males needs confirmation.
While examining Tortricidae collected in Japan, we found many unidentified specimens of Homalernis.Based on morphological examination, we concluded that this was an undescribed species.In this study, we describe this species as new; it represents the first record of the tribe Schoenotenini and the genus Homalernis from Japan as well as from the Palaearctic region.In addition, we discuss the validity of the species assigned to the genus Homalernis.

Sampling and dissection
We examined specimens deposited in the following institutions: CBM Natural History Museum and Institute, Chiba.Chiba, Chiba Prefecture, Japan.NSMT National Museum of Nature and Science.Tsukuba, Ibaraki Prefecture, Japan.ELKU Entomological Laboratory, Kyushu University.Fukuoka, Fukuoka Prefecture, Japan.
Images of adults were obtained using a SONY α7R IV digital camera fitted with a CANON MP-E 65 mm macro lens.To examine the male and female genitalia, the abdomens of the specimens were removed and boiled in a 10% KOH solution for approximately 10 min.After washing with 70% ethanol, the genitalia were dissected in 70% ethanol, stained with Chlorazol Black E solution, and mounted in Euparal on glass slides.The specimens were dissected and examined under a Nikon SMZ1000 binocular microscope.Images of the genitalia were obtained using a CANON 90D digital camera attached to a Nikon ECLIPSE Ci stereoscopic microscope and processed using Adobe Photoshop 2023 and Illustrator 2023 software.To examine wing veins, wings were removed, descaled using tissue papers and tweezers, stained with acetocarmine solution for 24 h, and mounted in Euparal on glass slides.Wing veins were drawn with Adobe Illustrator 2023 based on the images obtained using a CANON 90D digital camera connected to a Nikon ECLIPSE Ci stereoscopic microscope.

Terminology
The terminology used is primarily that of Razowski (2008).Additional terms related to the male genitalia are shown in Fig. 2D.

DNA extraction, PCR amplification, and sequencing
Six adult specimens of Homalernis fluctuosa Suzuki and Jinbo, sp.nov.were selected for DNA analysis (Table 1).DNA was extracted from the abdomen of each moth using a DNeasy Blood and Tissue Kit (Qiagen, Netherlands), following the manufacturer's protocol.
To obtain partial sequences of the mitochondrial cytochrome oxidase subunit 1 (COI) gene (standard DNA barcode region), the sequences were amplified using the primers LCO1490 (GGTCAACAAAT-CATAAAGATATTGG) and HCO2198 (TAAACTTCAGGGTGACCAAAAAATCA) (Folmer et al. 1994).The PCR reaction mixture consisted of 5 µL of KOD One® PCR Master Mix -Blue-(TOYOBO, Japan), 0.3 µL (10 pmol/µL) of each of the forward and reverse primers, and 1.0 µL of template DNA, with Milli-Q water added to a final volume of 10 µL.PCR amplification was performed as follows: an initial denaturing at 98 °C for 10 s, followed by 35 cycles at 98 °C for 10 s, 50 °C for 5 s, and 68 °C for 5 s.The amplified products were purified using ExoSAP-IT™ Express (Thermo Fisher Scientific Inc., USA) and sequenced using Pre-mixed Sanger sequencing services (Azenta, USA).
Pairwise distances based on the K2P method were calculated using MEGA X software (Kumar et al. 2018) to quantify the genetic distance between males and females and intraspecific variation.

Diagnosis.
Homalernis is characterised by the following features: hindwing veins R s and M 1 widely separate, M 3 and CuA 1 short stalked at the base (Fig. 4B); male genitalia with a deeply bifid uncus and a valva with both the costa and sacculus extended into projections (Fig. 2A, C) and female genitalia with a pair of wedge-shaped signa in the corpus bursae (Fig. 3A, B).
Homalernis is similar to Diactenis Meyrick, 1907 in wing venation with all veins free in the forewing, the M-stem well-developed; and the hindwing with a very slender median cell (Fig. 4A).The two genera also share a bifid uncus in the male genitalia.The female genitalia of Diactenis are easily distinguished by the absence of a distinct signum in the corpus bursae.The female genitalia of Homalernis shares with Syncratus Common, 1965 a twisted ductus bursae and a pair of similar signa, although the signum is a round plate with a blade-like process in Syncratus.Metachorista Meyrick, 1938 shares with Homalernis a valva with the costa and sacculus ending in distal projections in the male genitalia; and a similar shaped signum in the female genitalia, However, Metachorista has a hindwing with R s and M 1 stalked; prominent hami in the male genitalia; and only a single signum in the female genitalia.
Redescription. Adult (Fig. 1).Head coarsely and densely covered with scales.Parietal portion to frons densely covered with thick scales.Antennae approximately 1/2 length of forewing, with short sensory setae, scape basally covered with scales covering frons.Labial palpus approximately 1.5 times length of eye diameter, second palpomere broadening towards tip, apex covered with coarse hair-like scales.Forewing costa slightly curved basally, slightly concave medially and gently curved toward apex, apex roundish pointed and projected, termen strongly oblique, lightly curved, dorsum straight, except for curved basal part; patches of raised scales tufts at end of cell.Hindwing: slender and long, rather sparsely scaled except along veins, especially on underside, approximately 3 times as long as wide; costa curved basally, slightly concave medially, and straight and slightly oblique in outer half, apex rounded, termen rather straight and oblique below apex, dorsum rounded.
Venation (Fig. 4): Forewing: discal cell approximately 3/5 length of forewing, narrow at base and widening distally, slightly curved to dorsum, all veins separate except distal half of 1A+2A, R 1 to R 4 , R 5 to M 2 nearly equidistant; M 3 and CuA 1 slightly curved basally, CuA 1 from angle of cell, CuA 2 from 3/5 of cell, 1A+2A with basal fork.Hindwing: cell approximately 2/5 length of hindwing, slender and elongated, slightly broadened from base to middle, apical half tapering distally; R s and M 1 separate and remote at base, M 1 to M 3 equidistant, M 3 and CuA 1 stalked at base, from apex of cell, CuA 2 from distal 1/8 of cell, slightly sinuate to termen.
Distribution.India, Japan (new record), ?Malaysia.Diagnosis.Homalernis fluctuosa, sp.nov., is most similar to H. arystis in characteristics of the forewings and female genitalia.The two species can be distinguished based on the following: in H. fluctuosa, the apical third of the forewing has three distinct narrow fasciae, and the hindwing is approximately 3/4 the length of the forewing, whereas in H. arystis, the apical third of the forewings has scattered dark scales and the hindwing is approximately 2/3 the length of the forewing.In addition, the pair of signa of the female genitalia of H. fluctuosa is narrower and less sclerotised than those in H. arystis.
Thorax: Dorsum brownish white; tegula white.Forewing length 6 mm in holotype, 4-7 mm in paratypes (n = 62).Forewing; ground colour greyish white, with scattered pale brown scales, small dark brown spots along wing margin; blackish-brown dorsal blotch (DB) at 1/3 of dorsum in females, linear in males, subtriangular with round corners in females; black spot at 1/3 from base between dorsal blotch and costa; median fascia reduced to a series of pale brown spots, mixed with black scales in costal, central and dorsal portions, extending from just beyond middle of costa to 3/5 of tornus, costal portion broad, central portion extend along outer edge of cell, all scales not raised; subterminal fascia distinct, extending from 1/4 of costa to 1/4 of dorsum, gently convex, with black scales at middle, and costal and dorsal edges; subapical fascia thin, sometimes indistinct, extending from 6/7 of costa to middle of termen, nearly straight; preapical fascia wider than subapical fascia, extending from before apex to 1/3 of termen, nearly straight, costal and dorsal edges with black scales; cilia concolorous with ground colour, gradually becoming longer from apex to distal part of dorsum.Hindwing 4/5 length of forewing, with base to basal 2/3 slightly concave toward distal area of costa, basal 2/3 of costa to apex slightly curved, apex strongly curved and pointed, termen somewhat strongly curved, dorsum strongly curved; ground colour very pale ochre, rather sparsely scaled except along veins; cilia ochrous white, dorsal cilia becoming longer towards base of hindwing.

DNA sequence data.
The sequences data of the six examined specimens were deposited in GenBank and BOLD.For more details, see Table 1.
Etymology.The specific name fluctuosa refers to the three narrow fasciae in the apical third of the forewing that appear as ripples (Latin: flucticulus).
Remarks.The DNA barcodes of the individual collected from Okinawajima Island were 1.56-2.38%distant from those of individuals collected in Honshu, Kyushu, and Tsushima Island (Table 2).In addition, the forewing length of individuals collected from Okinawajima Island were slightly shorter than those of individuals collected from other regions.However, in the absence of other morphological differences, we provisionally assign the specimen to Homalernis fluctuosa.

Discussion
Homalernis fluctuosa, sp.nov., is the first species of the genus for which both sexes are known.The other known species of this genus, H. semaphora and H. arystis, were described by Meyrick based only on females, and H. jeriau and H. mankoboi were described by Razowski based only on males.We place H. fluctuosa, sp.nov., in Homalernis based on a comparison of its external characteristics and genitalia with those of H. semaphora, the type-species.The association of males and females of H. fluctuosa, sp.nov., was confirmed by molecular data (i.e., barcodes), which showed a genetic distance of 0.15% between females and males from the same locality (Hiroshima Prefecture, Akioota-Chou) (Table 2).A comparison of the male genitalia among several related schoenotenine genera revealed the following putative synapomorphies of Homalernis: uncus narrow and deeply bifurcated with a short fused basal part; gnathos with five short, pointed spines; and valva membranous and short, with a dorsoposterior projection and saccular projection 0.3-0.5 times the length of the valva.External features and the male genitalia of H. fluctuosa, sp.nov., differ significantly from those of the two species described by Razowski (2012).Hence, the generic and tribal position of those two may need to be re-evaluated.Moreover, their narrow and densely scaled hindwings are not similar to any other schoenotenine; they lack any raised scales on the forewing; and they lack the sparse scaling on the hindwing, with scales concentrated on the wing veins.Their male genitalia have a distinctly petiolate uncus, a broad median gnathos arm, and a valva with only one long ventral spine-shaped process, somewhat reminiscent of Tortricini.Although wing venation may be critical for their correct placement, Razowski (2012) simply states 'a somewhat simplified venation'.All known Schoenotenini have at least a trace of the forewing M-stem ending between M 1 and M 2 , rather than between M 2 and M 3 as in all other tortricids that possess an M-stem present.
The discovery of both sexes of a species of Homalernis provides an opportunity to re-examine its relationships to allied genera.In previous studies, relationships of Homalernis have focused on wing venation and female genitalia (Diakonoff 1960;Common 1965).Diakonoff (1960) stated that the wing venation of Homalernis is similar to that of Diactenis, and that the female genitalia are similar to those of Metachorista.However, these morphological characteristics were not described in detail.From our study, the following characteristics were found to agree with Diakonoff's assessment: in the forewing of Homalernis and Diactenis the M-stem originates from the basal 1/3 of the median cell, whereas in Metachorista the M-stem originates from approximately the middle of the cell; the female genitalia of Homalernis and Metachorista share the round basal plate of their signa, whereas Diactenis lacks a signum.Common (1965) considered Homalernis closely related to Syncratus, based on the twisted ductus bursae and the signa represented by a pair of plates with a blade-like projection in the female genitalia.Common (1965) also stated that Homalernis and Syncratus differ in the length of the basal fork of 1A+2A in the forewing, and that veins R s and M 1 of the hindwing extend nearly parallel and are widely separated at the base in Homalernis, whereas they are stalked in Syncratus.
Based on a comparison of the male genitalia of Homalernis and three genera previously considered closely related to this genus (i.e., Diactenis, Metachorista, and Syncratus), we conclude the following.All four genera share an elongated, tapering tip of the phallus; Homalernis shares a bifurcated uncus with Diactenis; and Homalernis shares elongated dorsoposterior and saccular projections reaching the distal edge of the valva with Metachorista.These morphological features suggest that Homalernis could be close to Diactenis and Metachorista.The very short stalked, apically bifurcate uncus and dorsoposterior and saccular projections, which are elongated far beyond the distal edge of the valva, are considered synapomorphies of Homalernis.A comprehensive phylogenetic analysis of Schoenotenini using DNA data is required to determine the details regarding the phylogeny and evolution of this tribe.
Schoenotenini are widely distributed from India to New Zealand (Horak 1998) and have been recorded in relatively high-altitude areas in the Northern Hemisphere.Several species (e.g.Homalernis semaphora, H. arystis, and Diactenis bidentifera Meyrick, 1928) were described from Assam (India), while D. youngi Razowski, 2000 was described from Taiwan.Komai and Nasu (2011) mentioned that at least one undescribed species is known from the Ryukyu Islands, Japan; however, its details remain unknown.Homalernis (excluding H. jeriau and H. mankoboi) is known only from India (Assam) and Japan, and H. fluctuosa is the most northerly species in the Schoenotenini.It seems likely that Homalernis and other schoenotenine species will be found in the northern Oriental and southern Palaearctic areas.

Table 1 .
Sample information of Homalernis fluctuosa used for DNA analyses.

Table 2 .
Intraspecific K2P pairwise distances between DNA barcode sequences of Homalernis fluctuosa.GenBank accession numbers are appended to each sample.