Twenty-six new species of Hoploscopa (Lepidoptera, Crambidae) from South-East Asia revealed by morphology and DNA barcoding

Abstract Hoploscopa Meyrick (Lepidoptera: Crambidae) is a fern-feeding genus found in montane areas of South-East Asia and Melanesia, eastwards up to the Samoan Islands. It includes sixteen described species, with at least 70 further undescribed species known from scientific collections. An iterative approach including morphological and molecular characters was used in order to explore the diversity of Hoploscopa. The hitherto described species are revised, and descriptions authored by T. Léger and M. Nuss are provided for an additional 26 new species: H. agtuuganonensissp. nov., H. albipunctasp. nov., H. albomaculatasp. nov., H. anacanthasp. nov., H. boletasp. nov., H. cynodontasp. nov., H. danaoensissp. nov., H. gombongisp. nov., H. gracilissp. nov., H. ignitamaculaesp. nov., H. isarogensissp. nov., H. jubatasp. nov., H. kelamasp. nov., H. kinabaluensissp. nov., H. mallyisp. nov., H. marijoweissaesp. nov., H. matheaesp. nov., H. niveofasciasp. nov., H. pangrangoensissp. nov., H. parvimaculasp. nov., H. pseudometacrossasp. nov., H. sepanggisp. nov., H. sumatrensissp. nov., H. titikasp. nov., H. tonsepisp. nov., H. ypsilonsp. nov. Using a protocol specific for the amplification of DNA from old museum specimens, we recovered 101 COI barcodes for all but one of the newly described species, with 76 being barcode compliant (>487 bp). Species delimitation analyses suggest cryptic diversity, with six cases reflecting allopatric divergence, and two further cases found in sympatry.


Introduction
South-East Asia is home to a rich biodiversity encompassing three of the 25 world's biodiversity hotspots (Myers et al. 2000). Pyraloidea is one of the largest superfamilies of Lepidoptera in the Oriental region, with 3,771 known species (Heppner 1998). Much of its diversity for this region remains to be described in this group (Nuss 1998, Sutton et al. 2015. Hoploscopa Meyrick best illustrates this knowledge gap, with 16 species currently described and more than 70 species awaiting description (Robinson et al. 1994). Hoploscopa moths display elongated brown forewings often with median and postmedian diagonal pale yellow to red markings. The genus is distributed across the Oriental and Australasian regions, ranging from the North of Thailand eastwards to the Samoa Islands, but is virtually absent from the tropical rainforests of Northern Queensland, Australia. A recent study reported the larvae to feed on ferns (Mally et al. 2017). Nuss (1998) synonymised Syncrotaula Meyrick, a replacement name for Eudorina Snellen, with Hoploscopa, and transferred twelve species to it. The author also provided a checklist of the genus and described one new species. Robinson et al. (1994) erected the tribe Hoploscopini within Scopariinae for Hoploscopa (= Syncrotaula) and Perimeceta Turner. Nuss (1998) transferred the Hoploscopini to the Heliothelinae based on the shared inwardly directed spine of the corpus bursae in the female genitalia. Perimeceta, the only other genus included in the Hoploscopini, is distributed from Java to South Australia and its larvae also feed on ferns (Miller et al. 2015, Mally et al. 2017). The systematic placement of Hoploscopini is still a matter of debate, with some authors grouping them together with Heliothelini in the Heliothelinae (Nuss 1998), while others consider the Heliothelinae an ingroup of Scopariinae (Leraut 1980, Minet 1982, Munroe and Solis 1998. The first molecular-based phylogeny including representatives of Hoploscopa suggest it to represent a separate lineage outside of Scopariinae + Crambinae (Léger et al. 2019).
This paper aims at establishing an iterative approach following Yeates et al. (2011) for species discovery and description in Hoploscopa, serving as a first step of a comprehensive revision of the genus. With an iterative approach using morphology and COI barcode, we provide redescriptions for fifteen of the 16 described species and describe 26 new species. Incongruence between morphology and molecular datasets, as well as factors that possibly influenced the evolution of the group are discussed.

Material acquisition
Material collected on Borneo, Java, Fiji, the Malay Peninsula, North-Sulawesi, New Guinea, the Philippines, Samoa, Sumatra and Vanuatu was obtained from the Museum für Tierkunde Dresden (MTD), the British Natural History Museum, London (NHMUK), the United States National Museum, Washington (USNM), the Museum für Naturkunde Berlin (MFNB) and the Zoologische Staatsammlung München, Munich (ZSM). Examination of the type specimens of the previously described species, all deposited at the NHMUK, was done by TL during a visit at that institution. Specimens described from Sabah, Malaysia are stored at the Institute for Tropical Biology and Conservation, Borneensis, Kota Kinabalu, Malaysia (BORN).

Iterative approach
In order to test species hypotheses with two independent sets of characters, we followed the best practice outlined by Schlick-Steiner et al. (2010), here referred to as "iterative taxonomy" (Yeates et al. 2011). This workflow started by sorting undescribed material to morphospecies based primarily on wing pattern, followed by examination of characters in male genitalia that show most variation. Subsequently, for each morphospecies, specimens with slightly differing wing pattern and/or of each collecting locality were selected for amplification of the mitochondrial COI barcode. Finally, morphospecies with at least one obtained COI barcode sequence were considered for species description (with the exception of H. marijoweissae sp. nov.). If strong divergence was observed in the COI barcode within a morphospecies, a second more careful examination of the specimens was conducted in the search of diagnostic characters, and COI barcode was sequenced for all specimens of the series. Specimens in poor conditions (i.e., wing pattern faded away, abdomen missing) or new species without male specimens available were not further considered for species description.

Molecular work
One hundred and fifty-seven dried museum specimens were considered for DNA extraction. Assuming a higher degree of degradation due to the age of the dried specimens, DNA extraction and preparation of PCR samples for specimens collected before 1990 (referred to as "old sample") were performed in the clean-room facility of the SGN-SNSD-Mol-lab at MTD. The abdomen (one or two legs if the abdomen could not be used) was carefully removed with sterilised pincers for DNA extraction, following a non-destructive method (Knölke et al. 2004) with the NucleoSpin Tissue kit (Macherey-Nagel, Düren, Germany) according to the manufacturer's protocol. DNA concentration of old samples was measured with a NanoDropOne spectrophotometer. We followed a workflow previously used by other authors (Hebert et al. 2013, Mitchell 2015, Hundsdoerfer et al. 2017) by amplifying up to five short overlapping fragments covering the COI barcode part (see Suppl. material 1: Fig. S1). Primers, amplicon length and amplicon overlap are listed in Suppl. material 2: Table S1. In the first step (recently collected samples only), amplification of the whole COI barcode sequence (658 bp) was attempted with the primer pair Hy-bLCO/HybNancy, each flanked with universal primer tails ("Hyb-") facilitating sequencing (Wahlberg and Wheat 2008). If amplification was not successful, we proceeded in the second step with the amplification of fragments 1a (LepF1/K699) and 1b (f220/LepR1 or HybNancy). The latter reverse primer showed a slightly better performance with respect to the amplification of fragment 1b and was preferentially used. If amplification of fragments 1a or 1b in step two failed, we proceeded in step three with amplification of fragments A, B, C (covering fragment 1a) and C, D, E (covering fragment 1b). PCR-mix included 2 µl of BIORON complete buffer 10× incl. Mg 2+ (25 mM), 2 µl of each primer at 10pmol/µl, 0.4 µl of dNTP (each 10 mM), 0.2 µl of Taq polymerase BIORON (Bioron DFS Taq, Ludwigshafen, Germany) ,1 µl (recent samples) or 1 to 8 µl (old samples) of DNA (depending on the quantity of DNA measured) and filled up with water to final volume of 20 µl. Alternatively, the AccuStart II GelTrack PCR SuperMix (2×) (Quanta BioSciences, Beverly, USA) ready-to-use mix was used in the following PCR-mix: 6.5 µl AccuStart ready-to-use-mix, 1 µl of each primer, 1-8 µl DNA and filled up with water to final volume of 25 µl. Initial denaturation at 95 °C during 5 min was followed by 42 cycles of 30 sec denaturation at 95 °C, 40 sec hybridisation at 49 °C, 50 sec elongation at 72 °C, with a final elongation at 72 °C for 10 min. Amplification success was checked by electrophoresis on 1 or 2% agarose gels, subsequently stained with GelRed and visualised under UV light. Sequencing was performed by Macrogen (Netherlands) using the original PCR primers or the T7 and T3 sequencing primers (Wahlberg and Wheat 2008). Full COI-barcoding PCR products were sequenced in the forward direction only, while the shorter PCR products were sequenced in both directions.

Data analyses
Sequences were checked by eye and concatenated using PHYDE 0.9971 (Müller et al. 2005). In order to assess the amplification of the correct fragment, sequences were blasted with the blastn program against the nucleotide collection (nt), as provided by the National Center for Biotechnology Information (NCBI). In order to identify potential nuclear pseudogenes, sequences were searched for indels, internal stop codons and double peaks in electropherograms, as suggested in Song et al. (2008). Newly generated sequences (including ENA and BOLD accession numbers), as well as sequences retrieved from Barcode of Life Data System (BOLD, http://v4.boldsystems. org/) are reported in Suppl. material 3: Table S2. For the phylogenetic analyses, Parti-tionFinder2 (Lanfear et al. 2017) was used to determine the best partitioning scheme using the AICc model and the greedy search algorithm (Lanfear et al. 2012). Partition for each of the three codon positions was recovered as the best model and subsequently used for further analyses. Maximum Likelihood (ML) analysis was performed with RAxML (Stamatakis 2006) as implemented on the CIPRES portal (Miller et al. 2010), using the GTR+GAMMA substitution model and the best PartitionFinder scheme. Node support was estimated with 1000 thorough bootstrap replicates using the same algorithm.

Species delimitation
Two different methods were used to investigate species delimitation in our molecular dataset: Automatic Barcode Gap Discovery (ABGD) (Puillandre et al. 2012) and General Mixed Yule-coalescent method (GMYC) (Pons et al. 2006, Fujisawa andBarraclough 2013). These popular methods have been repeatedly used in DNA barcoding studies on Lepidoptera (Kekkonen and Hebert 2014, Dincă et al. 2015, Dumas et al. 2015, Nakahara et al. 2019). We noticed a detrimental effect of non-compliant barcode sequences (< 487 bp) on the estimation of species in densely sampled clades (n > 2), where these sequences were flagged as new species. Consequently, these sequences were removed to generate a COI_487 bp dataset. Distance-based analysis ABGD was calculated using the K80 Kimura distance model and X fixed to 0.5 on the dedicated platform (https://bioinfo.mnhn.fr/abi/public/abgd/abgdweb.html). Genetic distances are referred to as K2P-dist throughout the text.
The ultrametric tree required for the GMYC analysis was generated using BEAST 1.10.4 ). The COI dataset was partitioned after codon position accordingly to the best PartitionFinder2 scheme. An uncorrelated relaxed clock with lognormal distribution was used and the Speciation: Yule Process model was set as tree prior model and other parameters were left unchanged. MCMC chain was set to 40 million generations, sampled every 1000 th generation. Convergence was checked on Tracer ) and the first 4 million generations were discarded as burnin. Species delimitation analysis was performed using the GMYC method as implemented in the R-package SPLITS (Ezard et al. 2009, Fujisawa andBarraclough 2013). The single-threshold option was used as suggested in Fujisawa and Barraclough (2013).

Systematic treatment
Genitalia were mounted following the method of Robinson (1976). Photographs of the habitus of imagines were taken with an Olympus E-M1 with the Olympus M. Zuiko Digital ED 60 mm f/2.8 Macro Lens. Photographs of genitalia and measurements were performed on a Nikon Eclipse 90i at the MTD. Images were subsequently enhanced on Adobe Photoshop CS6 and illustrations plates were created on Adobe Illustrator CS6. Collecting data of holotypes was copied exactly as found on the labels, with vertical bars to mark line breaks. Abbreviations or translations are given in square brackets where judged meaningful. Paratype data are reported by country in alphabetical order, with information reported without indication of line change. Collecting localities are reported as written on labels. Dates and collectors' information were standardised and the latter placed in parentheses. The specimen depositories are reported with the use of the corresponding acronyms. GPS coordinates for localities were retrieved from the online platform geographic.org (https://geographic.org/geographic_names/) using the database of the National Geospatial-Intelligence Agency. Localities treated in this paper are reported in Table A1 in Appendix 1. Specimens whose conspecificity was uncertain were listed under "Other specimens examined". Nomenclature follows that of Landry (1995), except for the use of the term phallus (Kristensen 2003). The colours "bronze", "ochre", and "tawny" refer to the Wikipedia list of colours by shade for brown (https://en.wikipedia.org/wiki/Category:Shades_of_brown). Character description was facilitated by the use of Mesquite (Maddison and Maddison 2017) where each character was given a state, thereby allowing a better visualisation of character state variation as well as warranting consistency among species descriptions.

Molecular dataset
We obtained 101 COI barcodes ranging in length from 261 to 658 bp (mean length = 545 bp), among them 76 barcode "compliant sequences" (> 487 bp) according to Hebert et al. (2013). Ten non-compliant barcode sequences were kept in the final dataset for species with no or one barcode sequence. Fifteen further non-compliant barcode sequences were discarded from the final dataset (see Suppl. material 2: Tab. S1). We recovered the COI barcode for all but one (H. marijoweissae sp. nov.) species described in this paper and for seven of the sixteen previously described species. Twenty-five Hoploscopa sequences, as well as 13 sequences from the hypothesised sistergroup Perimeceta were retrieved from BOLD and added to our dataset, totalling 126 COI barcode sequences including nine of the described species. Eleven species were represented by only one barcode sequence. Recovered fragments and total sequence length are summarised in Suppl. material 3: Table S2. The oldest sample from which we could successfully amplify COI sequence data were collected in 1955 (63 years old).
The Maximum Likelihood analysis of the COI dataset is illustrated in Fig. 123, with bootstrap support (abbr. BS) over 50 displayed on nodes. Genetic distances (K2P-dist) between all COI barcodes are reported in Suppl. material 4: Table S3.

Species delimitation
Analysis of the morphology allowed delineation of 39 species. Both ABGD and GMYC analyses highlighted 48 molecular operational taxonomic units (MOTUs) in Hoploscopa species, splitting seven of the morphospecies into two MOTUs, and one into three MOTUs (H. danaoensis sp. nov.). Six of these splits reflect allopatric divergence. In each of the two species H. matheae sp. nov. and H. parvimacula sp. nov., there is one female from the Malay Peninsula recovered as a separate MOTU from other specimens from Borneo. In H. luteomacula, the two specimens from Borneo are recovered as a distinct MOTU to that of Sumatra. In H. albipuncta sp. nov., a female from Luzon (Philippines) was recovered in a MOTU distinct from that including the specimens from Borneo. Genetic differentiation is also observed among Philippines islands: specimens of H. danaoensis sp. nov. from Danao, Negros, and Mindanao represent three different MOTUs, while specimens of H. isarogensis sp. nov. from Luzon and Leyte are also recovered into two distinct MOTUs.
Sympatric divergence is observed in H. kinabaluensis sp. nov. and H. sumatrensis sp. nov. Hoploscopa kinabaluensis sp. nov. is split into two MOTUs representing sister clades and diverging from each other by 1.7-2.2%. In H. sumatrensis sp. nov., two separate MOTUs are recovered by the ML analysis in a clade with DNA samples BC_ MTD_LEP01421 (Hoploscopa sp. near sumatrensis) and BC_ZMBN_Lep00081 (larva of Hoploscopa sp.). The two H. sumatrensis MOTUs show a divergence of 4.1-6%. Second MOTU, represented by samples MTD8258 and BC_MTD_LEP01422, include only females and need examination of male specimens. Diagnosis. Hoploscopa displays brown to dark brown forewings, often bearing pale yellow-, yellow-, or red-coloured median and postmedian diagonal stripes. In male genitalia, the uncus is well developed, the gnathos forms a ribbon-like structure, often with a posterior projection, and the vinculum bears laterally a pair of coremata. In female genitalia, the antrum is short, often sclerotised, and the corpus bursae bears a thorn, often with a basal sclerotisation. Hoploscopa is morphologically very similar to Perimeceta. The forewings of Perimeceta are slightly larger, display a yellow to brown ground colour, with one basal elliptic and one postmedian lunule-shaped white spot.

Systematics
In male genitalia, Perimeceta shows a spade-shaped uncus, the gnathos arms connect shortly after arising and are expanded posteriorly into an elongated tip, the valva shows a conspicuously rounded ventral margin, and is apically narrowed into a tip, while the ventral margin is more or less straight in Hoploscopa. Perimeceta exhibits female genitalia similar to those of Hoploscopa with a slender ductus bursae and a rounded corpus bursa bearing a thorn. However, the latter displays a membranous antrum in female genitalia, while it is sclerotised in most species of Hoploscopa.
Description. Head. Antennae dorsally with pale yellow to brown scales. Ocelli absent. Frons slightly produced, rounded. Proboscis basally white to brown scaled. Maxillary palpi brown to dark brown, first segment pale yellow, inner side brown to pale yellow. Labial palpi porrect, 2-2.9 × diameter of compound eye, brown, white to pale yellow at base, inner side brown to pale yellow.
Thorax . Collar white to pale yellow. Frenulum simple in ♂, triple in ♀. Forewing length 7-13 mm, 2.4-2.9 × maximal width, females slightly larger than males. Wing venation (Fig. 2): R1 not connected to Sc; R2, R3 and R4 stalked together; R5 free, arising from upper corner of cell; M1, M2 not stalked; M3 arising from lower corner of cell; CuA1 arising below lower corner; CuA2 at distal 1/3 of cell; 1A+2A arising from cell base; Hindwing Sc+R1 connected to Rs at distal 1/3; M1 connected to Sc+R1 by short vein; cell closed; M2, M3 and CuA1 arising at lower angle of cell; CuA2 arising at middle of cell; 1A, 2A, 3A free. Forewing ground colour brown to dark brown, markings white, yellow or red colour. The pattern elements can be described as follows (Fig. 3): cf costal field stretched along costa up to postmedian area; bs basal streak small, weakly or not marked; bp basal patch often quadrangular, ill-defined; pds and dds proximal and distal discoidal stigmata along costal field; mds the median discoidal stigma trapezoid, with costal edge longer than dorsal one; mcp and mdp median cubital patch and median dorsal patch in line with median discoidal stigma, often forming together a Y-shape pattern; pmp postmedian patch roughly triangular, narrowing at costa; pml postmedian line running from costa to middle of dorsum, often barely marked, mostly visible close to costa; pma postmedian area often suffused to a various extend with white, yellow or red; sbl subterminal line running from costa close to apex towards dorsum, often incurved inwards at CuA2; sbf subterminal field more or less suffused with brown, red or yellow.  The margin is brown, in some species with spots, fringe unicoloured or chequered. Hindwing upper side pale yellow to pale brown; underside pale yellow, with brown markings on costa and subterminal line toward costa; males of some species with androconial organ on the upper side at dorsum, consisting of upright scales along CuA2 and 1A veins and a protruded margin between CuA2 and 1A, bearing a patch of greyish scales. Forelegs brown to dark brown. Midlegs with femur brown, tibia and tarsi often pale yellow, speckled with brown. Hindlegs pale yellow on inner side, pale yellow to brown on outer side, tarsi bronze to pale brown.
Abdomen. Pale brown to brown. In males, sternum A8 more or less broadly indented, in some species with short, rounded lateral projections (Figs 85,86).
Diagnosis. Hoploscopa albipuncta sp. nov. displays a basal white well-rounded spot on the forewing. In male genitalia, the dorsal margin of the valva is protruded, the juxta displays two conspicuous tips and the phallus bears an anvil-shaped cornutus. In female genitalia, the antrum is membranous and forms a rounded pouch.
Distribution. Known from the slopes of the Mount Kinabalu (4,095 m) on Borneo, at altitudes between 1,700 m and 2,000 m. DNA barcoding. Specimen MTD7430 from Luzon shows an K2P-distance of 4.5-4.9% with the two specimens from Borneo.
Etymology. The species name albipuncta refers to the Latin albus, white, and punctus, forming a point.
Distribution. Known from the Malay Peninsula, and Borneo (Brunei, Mount Kinabalu), at altitudes between 1,300 m and 1,600 m. DNA barcoding. Specimen MTD8244 from the Malay Peninsula shows an K2Pdistance of 2.2% with the two specimens from Borneo. It is recovered as a distinct MOTU in the species delimitation analyses. Specimens from Kinabalu and Tawau Hills share identical COI barcodes. No COI barcode was obtained for the specimen from Brunei (MTD8245).
Diagnosis. Hoploscopa sepanggi sp. nov. bears sulfurous to pale yellow patches on the forewing forming a roughly chequered pattern. In male genitalia, the gnathos projection is slender, ca. half the length of uncus, and the juxta distal half is narrow, with a notched apex. Similar species. Hoploscopa luteomacula. The postmedian cubital and subterminal patch are missing or reduced to traces in H. sepanggi sp. nov., while they are wellmarked, pale yellow in H. luteomacula. The fringes are brown, chequered with yellow in H. sepanggi sp. nov., while they are yellow in H. luteomacula.
Description. Head. Antennae dorsally striped with pale yellow and brown scales. Proboscis brown, speckled with pale yellow. Maxillary palpi brown, base and inner side pale yellow. Labial palpi brown, ventral base and inner side pale yellow. Thorax (Fig. 5). Thorax dorsally pale yellow, laterally brown. Collar pale yellow. Forewing length: 11-12.5 mm (♂), 11.5-13 mm (♀); forewing ground colour brown; basal patch and median discoidal stigma trapezoid, sulphur yellow with dark brown edges; basal and antemedian cubital patches abutting each other, quadrilateral, sulphur yellow with dark brown edges; costal field lightly marked with sulphur yellow; median dorsal patch oval, sulphur yellow; postmedian patch triangular, sulphur yellow, crossed on its middle with longitudinal brown streak; subterminal line reduced to small interspaced sulphur yellow dashes, with triangular sulphur costal spot; subterminal field broadly marked with pale yellow; fringe chequered pale yellow and brown. Hindwing pale brown. Forelegs dark brown. Midlegs brown, speckled with pale yellow. Hindlegs with femur brown; tibia pale yellow, speckled with brown, distally brown; tarsi bronze.
Distribution. Known from the slopes of the Mount Kinabalu on Borneo, at altitudes between 1,800 m and 2,000 m.
Etymology. This species is described in honour of the late Malaysian mountain guide Robbie Sepanggi who died on Mount Kinabalu during the 2015 Sabah earthquake while trying to save hikers. Diagnosis. This relatively small-sized species (forewing length 8-10 mm) shows brown forewings with red markings edged with pale yellow. The median discoidal stigma is oval, red, edged with pale yellow, median cubital patch is reduced to a small streak and median dorsal patch is absent. The postmedian patch consists of a mix of red and brown scales, at costa forming triangular pale yellow patch. Male genitalia are unique in the conspicuously incurved ventral margin of the valva, medially extending into a tip pointing ventrad. In female genitalia, the corpus bursae is small, globular, and bears a crest of sclerotised acanthae between thorn and corpus opening. Similar species. Hoploscopa isarogensis sp. nov.; to a lesser extent H. mallyi sp. nov., H. agtuuganonensis sp. nov., H. gracilis sp. nov., H. ignitamaculae sp. nov. Median cubital and dorsal patches form with the median discoidal stigma a disrupted pale yellow band (median dorsal patch not marked in H. mallyi sp. nov.), and postmedian pale yellow marking at costa is reduced to a blotch in above listed species. Hoploscopa gracilis sp. nov. and H. mallyi sp. nov. are slightly larger (9-12 mm). Comparison of the male genitalia allows unequivocal separation from these species. In female genitalia, H. isarogensis sp. nov. displays a narrower and slightly longer ductus bursae and a larger corpus bursae with large curved thorn, while it is small and straight in H. cynodonta sp. nov. Female genitalia of H. mallyi sp. nov., H. agtuuganonensis sp. nov., and H. gracilis sp. nov. share the broad ductus bursae, the small globular corpus bursae with H. cynodonta sp. nov. but have a longer ductus bursae, and the sclerotisation of corpus bursae is large and diffuse.

Hoploscopa cynodonta
Description. Head. Antennae dorsally with pale yellow scales. Proboscis brown to pale brown. Maxillary palpi brown, base and inner side pale yellow. Labial palpi brown, ventrally pale yellow.
Abdomen. Male sternum A8 posterior margin forming two conspicuous triangular tips, with short, rounded lateral projections.
Distribution. Known on Borneo from Brunei to the Mount Kinabalu, the Mount Monkobo (1,759m) and the Tawau Hills in Sabah (Malaysia), at altitudes between 300 and 1,700 m.
Etymology. Formed by apposition of the Greek words cyno-, of the dog, and odous, tooth, referring to the conspicuous tooth-like extension on the ventral margin of the valva in male genitalia. Diagnosis. Hoploscopa parvimacula sp. nov. displays brown forewings with reduced pale yellow markings. The median discoidal stigma is crescent-shaped, and the median cubital patch forms a streak not connected to the median discoidal stigma. The postmedian patch and postmedian area are faintly marked with pale yellow. In male genitalia, the gnathos projection is distally spatula-shaped, reaching ca. 2/3 of the uncus length, and the juxta displays blunt lateral projections and a truncate apex. Similar species. Hoploscopa kinabaluensis sp. nov., H. pangrangoensis sp. nov. The forewings of H. kinabaluensis sp. nov. display more contrasting postmedian patch and subterminal line, and a subterminal field suffused with pale yellow. In H. pangrangoensis sp. nov., the median cubital patch is almost absent and the subterminal line is more strongly pronounced. In male genitalia, base of gnathos projection is narrower and the cornutus displays a narrow bump at apex in H. kinabaluensis sp. nov. The finger-like gnathos projection of H. pangrangoensis sp. nov. is narrower than that of H. parvimacula sp. nov. and reaches only half the uncus length. In female genitalia, ductus bursae of H. kinabaluensis sp. nov. is twice as long as in other species, with a loop on its middle and a large bow before corpus bursae. In H. pangrangoensis sp. nov., colliculum is membranous with longitudinal sclerotised lines, corpus bursae is larger and thorn is smaller.

Hoploscopa parvimacula
Description. Head. Antennae dorsally striped with brown and bronze scales. Proboscis pale yellow, speckled with pale brown. Maxillary palpi dark brown, base and inner side pale yellow. Labial palpi dark brown, ventro-basally pale yellow.
Thorax (Fig. 7). Collar pale yellow. Forewing length: 9-10 mm (♂ & ♀); forewing ground colour brown; basal dark brown dash, distally pale yellow; basal and distal discoidal stigmata dark brown; median patches Y-shaped, pale yellow, disrupted with brown at vein, edged basally and distally with dark brown; postmedian patch of a lighter brown, with pale yellow triangular costal spot distally edged with dark brown; subterminal line pale yellow, distally dark brown; fringe brown, with pale yellow dots. Hindwing pale brown. Forelegs brown. Midlegs with femur brown; tibia pale yellow, speckled with brown; tarsi bronze. Hindlegs with femur brown; tibia pale yellow, speckled with brown; tarsi pale yellow to pale brown.
Female genitalia (N = 2) (Fig. 91). Anterior apophyses with dorsal bump at posterior 1/3. Antrum sclerotisation twice as long as broad. Ductus bursae short and straight, slightly bent before corpus bursae. Corpus bursae pear-shaped, posterior half reticulated, anterior half membranous, with diffuse sclerotisation between thorn and corpus opening, medially with faintly marked sclerotised band. Thorn short and straight, with small dents pointing toward thorn apex.
Distribution. Known from Mount Kinabalu on Borneo, at altitudes between 1,550 and 1,950 m. DNA barcoding. Sample MTD8229 of H. parvimacula sp. nov. shows a K2P-dist of 3.1-3.4% with samples from Borneo and is recovered as separate MOTU.
Etymology. Refers to the Latin parvus, for small, and macula, for spot, referring to the Y-shaped median markings that are smaller than in other similar species.
Remarks. The specimen with DNA voucher MTD8229 belongs to a series of females from the Malay Peninsula, which are morphologically similar to H. parvimacula sp. nov. and deposited at NHMUK. Since no males of the same series were available for investigations, we restrained from drawing further conclusions here. Diagnosis. The forewing median markings form an interrupted pale yellow Y, and postmedian patch and subterminal line are pale yellow, well-marked. In male genitalia, the gnathos projection is ca. 2/3 of uncus length, spatula-shaped, broadening toward apex, and the cornutus displays a small but conspicuous bump. Female genitalia show a long ductus bursae broadly curved twice and a large pear-shaped corpus bursae. Similar species. Hoploscopa parvimacula sp. nov. (q.v.), H. brunnealis, H. danaoensis sp. nov., H. metacrossa. These species share similar forewing markings with H. kinabaluensis sp. nov. Male genitalia of these species show a gnathos projection larger at base, of constant width, reaching ca. half of uncus length, and a cornutus lacking the apical bump. In female genitalia, the shorter ductus bursae is nearly straight.

Hoploscopa kinabaluensis
Description. Head. Antennae dorsally striped with pale yellow and bronze scales. Proboscis brown speckled with pale yellow. Maxillary palpi brown, base and inner side pale yellow. Labial palpi brown, ventral base and inner side pale yellow.
Distribution. Known from the slopes of Mount Kinabalu and Mount Monkobo on Borneo, from altitudes between 975 m and 1,850 m. DNA barcoding. Two MOTUs diverging by 1.7-2.2% are recovered in H. kinabaluensis sp. nov. These two lineages are found in sympatry at Mount Kinabalu.
Etymology. The species name kinabaluensis refers to Mount Kinabalu on Borneo, where the species occurs.
Remarks. The two barcode lineages show no differences in wing pattern and female genitalia. Only minor differences are found in male genitalia, with the apex of the gnathos projection slightly narrower and the basal margin of the juxta slightly incurved in the second lineage. Since second MOTU is only represented by three specimens including only one male, further investigations with a greater number of specimens is needed. Nuss, 1998 Figs 9, 52, 93 Material examined. Holotype: ♂, with labels: "Holotypus"; "SUMATRA, Barat | N-Padangpanjang | Mt. Singgalang 2100m | 10-11.ii.1996, L[icht]F[ang] [light trap] | leg. A. Kallies"; "HOLOTYPE | Hoploscopa luteomacula | det. Nuss, 1996";"GU 744 | prep. Nuss 1996 Diagnosis. Hoploscopa luteomacula displays broad pale yellow patches in the forewing. In male genitalia, the uncus is rectangular with a truncate apex, the gnathos forms a tongue-shaped projection ca. 1/4 of the uncus length, the juxta is medially conspicuously narrowed and displays a duck-shaped apex.
Distribution. Known from Sumatra (Indonesia) at altitudes between 1,200 m and 2,100 m.
DNA barcoding. Specimens MTD8234 and MTD LEP3195 from Borneo show an K2P-distance of 2.5-4.4% with the specimen from Sumatra and are recovered as a distinct MOTU.

Hoploscopa obliqua
Diagnosis. The antennae striped with white and the antemedian oblique white streak extended distally into white suffusion in the forewing are unique to this species. In male genitalia, the gnathos projection forms a broad plate with a small apical tip. In female genitalia, the antrum is membranous, the ductus bursae is long and corpus bursae is densely covered with erect papillae on one half. Similar species. No similar species known. Description. Head. Antennae dorsally striped with brown and white. Proboscis brown. Maxillary palpi brown, base pale brown, apex white. Labial palpi brown with white apex. Thorax (Fig. 10). Collar white. Forewing length: 8-9 mm (♂ & ♀); forewing ground colour brown; basal area of a darker brown; antemedian oblique thick white streak running from dorsum to upper margin of cell, distally extended into white suffusion; postmedian patch reduced to costal white blotch; postmedian line of a darker brown at costa; barely marked subterminal white line incurved inwards at CuA2, running up to M2; fringes pale brown. Hindwing pale brown. Fore-and midlegs brown, tibia and tarsi segments distally white. Hindlegs brown.
Distribution. Known from the Papua (Indonesia) and Madang Provinces (Papua New Guinea) on New Guinea, at altitudes between 1,000 m and 1,700 m.
Phylogenetic relationships. Hoploscopa niveofascia sp. nov. is recovered as sister group in the ML analysis of the COI barcode (BS = 95).
Female genitalia Not known. Distribution. Known from Mount Susu (975 m) in the Morobe Province (Papua New Guinea).
Phylogenetic relationships. Hoploscopa obliqua is recovered as sister group in the ML analysis of the COI barcode (BS = 95).
Diagnosis. The well-marked median cubital and dorsal snow-white patches on the forewing segregate this species from its congeners. Median discoidal stigma is trapezoid, reddish brown, postmedian patch is reddish brown, distally edged by thin white streak. In male genitalia, the narrow uncus-tegumen connection and the broadly indented uncus apex is unique to this species. In female genitalia, papillae anales are thick, not connecting dorsally and ventrally, corpus bursae is small, globular, with a long straight thorn.
Similar species. No similar species known. Description. Head. Antennae dorsally brown. Proboscis pale brown. Maxillary palpi dark brown, base and inner side pale yellow to light brown. Labial palpi dark brown, ventro-basally pale yellow. Thorax (Fig. 12). Collar white. Forewing length 10-11 mm (♂), 9-10 mm (♀); forewing ground colour dark brown; cubital reddish brown fascia running from basal to postmedian area; basal and distal discoidal patches of a darker brown; rhombical reddish brown median discoidal stigma therebetween, basally and distally thinly edged with pale yellow; median cubital and dorsal patches white, elongated, slightly disrupted at 1A+2A; postmedian roughly quadrangular reddish brown patch, crossed with brown lines, with slender white streak abutting dorsally, running up to costa; postmedian line thin, marked on costal half; postmedian fascia white, speckled with brown; subterminal line thin, white, running more or less straight from dorsum distal 1/4 to apex; subterminal field faintly marked with reddish brown; fringes brown, with pale yellow dots. Hindwing pale yellow, slightly darker at apex. Legs brown, tibia distally pale yellow, tarsi brown to pale brown.
Distribution. Known from Yawan village (1,700 m) in the Eastern Highlands Province (Papua New Guinea).
Diagnosis. Hoploscopa tonsepi sp. nov. displays a triangular reddish brown median discoidal stigma with white edges, extending toward dorsum into narrow white cubital streak. In male genitalia, the two-armed uncus with two small tips on its middle is unique to this species. In female genitalia, the absence of posterior apophyses, the short ductus bursae and the small corpus bursae without thorn are atypical for the genus and observed only in this species.
Similar species. No similar species known. Description. Head. Antennae dorsally striped with brown and bronze scales. Proboscis pale yellow, brown at base. Maxillary palpi dark brown, base and inner side pale yellow to light brown. Labial palpi dark brown, ventro-basally pale yellow.
Thorax (Fig. 13). Collar pale yellow. Forewing length: 9-10 mm (♂ & ♀); forewing ground colour brown, broadly suffused with reddish brown; basal oblique thin white streak; cubital reddish brown fascia running from basal to postmedian area; basal and distal discoidal patches of a darker brown, basally and distally thinly edged with white; median discoidal stigma reddish brown, with V-shaped white edge, together with narrow white cubital streak forming a Y; postmedian patch roughly triangular, reddish brown, crossed with brown lines, abutted with pale yellow blotch at costa; postmedian line broad, marked on costal half; postmedian suffusion faintly marked, pale yellow; subterminal line pale yellow, diffuse, running more or less straight from dorsum distal 1/4 to apex; subterminal field marked with reddish brown; fringes brown, with pale yellow dots. Hindwing pale brown, darker at apex. Forelegs dark brown; tibia dark brown, distally pale yellow; tarsi pale yellow. Midlegs with femur dark brown; tibia pale yellow, speckled with brown, distally brown; tarsi pale yellow. Hindlegs brown; tibia and tarsi segments distally pale yellow.
Female genitalia (N = 1) (Fig. 101) Diagnosis. This species is recognisable by the brown forewings broadly suffused with copper-coloured scales giving them a shiny appearance. Median cubital patch is white, discoidal stigma and postmedian patch are yellow, the latter marked with white toward dorsum and at costa. In male genitalia, uncus is elongated, with a truncate apex, and gnathos projection is ca. half the length of uncus, with truncate apex. In female genitalia, antrum is barely sclerotised, ductus bursae is long, slender and straight and corpus bursae is large, globular, with a small slightly curved thorn.
Similar species. No similar species known. Description. Head. Antennae dorsally bronze. Proboscis pale yellow. Maxillary palpi brown, inner side pale brown. Labial palpi brown, ventrally pale yellow, inner side pale brown.
Thorax (Fig. 14). Collar pale yellow. Forewing length 11 mm (♂), 12 mm (♀); forewing ground colour brown; broad basal yellow band stretching from subdorsum to costa, with crossing veins copper-coloured, edged with copper; costal field copper; median discoidal stigma yellow with copper edges, costally filled with copper, forming together with cubital trapezoid white patch a canine tooth shape; postmedian patch broad, yellow, basally encroached with copper, crossing veins copper-coloured, with white blotch at costa; postmedian line broad, marked on costal half; broad copper postmedian fascia with crossing veins brown; subterminal field marked with a mix of yellow and copper scales; fringe brown, with white dots. Hindwing white to pale brown. Forelegs brown. Midlegs with femur brown; tibia pale yellow, apically brown; tarsi pale yellow to pale brown. Hindlegs with femur brown; tibia pale yellow, speckled with brown; tarsi pale brown.
Distribution. Known from Mount Yamin (Indonesia: Papua), and Mount Kaindi (Papua New Guinea: Morobe Province), at altitudes between 1,700 m and 2,350 m.
Etymology. Dedicated to the late Marijo Weiss, a close friend of Théo Léger's family.
Remarks. The specimen from Mount Kaindi is very similar to those from Mount Yamin but its postmedian fascia of the forewing is yellow and copper instead of dark copper, and uncus and gnathos projection are slightly longer in male genitalia. Diagnosis. Hoploscopa titika sp. nov. is recognisable by its well-marked white quadrangular median cubital spot on the forewing. In male genitalia, the gnathos projection is reduced to a small ridge, and the dorsal margin of the valva is rounded, broadly sclerotised. Female genitalia not known.

Hoploscopa titika
Similar species. No similar species known. Description. Head. Antennae dorsally with pale yellow to bronze scales. Proboscis white. Maxillary palpi brown, inner side white to pale brown. Labial palpi brown, ventro-basally white.
Thorax (Fig. 15). Collar white. Forewing length: 9 mm; forewing ground colour brown; basal patch extending from dorsum to subcosta, red, with small basal pale yellow spot; median discoidal stigma triangular, pale yellow, filled with red scales, with white trapezoid cubital patch abutting dorsally; postmedian patch roughly quadrangular, pale yellow, filled with red scales; subterminal line diffuse, white; subterminal field marked with red and pale yellow; fringe missing on specimen. Forelegs lost. Mid-and hindlegs white to pale yellow.

Hoploscopa pangrangoensis
Diagnosis. The forewing markings of H. pangrangoensis sp. nov. are reduced to a small crescent-shaped median discoidal stigma; median cubital and dorsal patches are not marked, and postmedian patch is reduced to a blotch at costa. In male genitalia, the uncus is medially widened and the gnathos shows a finger-like projection ca. half the length of uncus. In female genitalia, the thin longitudinal sclerotised lines of the antrum are unique to this species. Similar species. Hoploscopa parvimacula sp. nov. (q.v.), H. sumatrensis sp. nov. Hoploscopa sumatrensis sp. nov. shares with H. pangrangoensis sp. nov. the crescentshaped pale yellow median discoidal stigma on the forewing and the small median cubital patch. However, median discoidal stigma and postmedian patch are filled with reddish brown in H. sumatrensis sp. nov. In male genitalia, the gnathos projection reaches 4/5 of the uncus length in H. sumatrensis sp. nov. In female genitalia, antrum of H. sumatrensis sp. nov. is completely sclerotised.
Description. Head. Antennae dorsally striped with brown and bronze scales. Proboscis pale brown. Maxillary palpi brown, ventro-basally pale yellow. Labial palpi brown, base and inner side pale yellow. Thorax (Fig. 16). Collar pale yellow. Forewing length: 10-11 mm (♂ & ♀); forewing ground colour brown; basal dark brown dash distally pale yellow; small pale yellow spot at base of cell; basal and distal discoidal patches dark brown; median discoidal stigma crescent-shaped, white to pale yellow; median cubital pale yellow blotch in some specimens; postmedian pale yellow streak at costa, distally with dark brown streak; postmedian area speckled with pale yellow; subterminal line pale yellow, diffuse; fringes brown, with pale yellow dots. Hindwing pale brown. Forelegs brown, tarsi pale brown. Midlegs with femur brown; tibia brown speckled with pale yellow; tarsi pale yellow speckled with bronze. Hindlegs with femur brown; tibia pale yellow, speckled with pale brown; tarsi pale yellow.
Distribution. Known from the slopes of Mount Pangrango (3019 m) on Java (Indonesia), at an altitude of 1,625 m.
Etymology. Named after Mount Pangrango, a dormant stratovolcano on Java, where the specimens were collected. Diagnosis. Hoploscopa isarogensis sp. nov. is a relatively small brown-winged species (forewing length = 7-9.5 mm) with red markings edged yellow. Median discoidal stigma, cubital and dorsal patches form together a disrupted band, and postmedian patch is red with a pale yellow blotch at costa. In male genitalia, the apex of the uncus is duck beak-shaped, with a small marked bump on its ventral side, and gnathos projection is ca. 1/3 of uncus length, with an indented apex. In female genitalia, the corpus bursae displays a clearly delimited sclerotisation between the thorn and the corpus opening, and bears a long, thick, curved thorn. Similar species. Hoploscopa cynodonta sp. nov. (q.v.), H. ignitamaculae sp. nov. (q.v.), H. agtuuganonensis sp. nov. The forewings of H. agtuuganonensis sp. nov. display a trapezoid median discoidal stigma, and median markings are only slightly disrupted, while they are well separated in H. isarogensis sp. nov. In male genitalia, gnathos projection reaches 2/3 of uncus length and has a truncate apex. In female genitalia, ductus bursae is long, broad, with one loop, while it is shorter, slender and straight in H. isarogensis sp. nov., and the small corpus bursae displays one plump thorn with a small sclerotisation at its base.
Thorax (Fig. 17). Collar white. Forewing length: 7-8 mm (♂), 8-9.5 mm (♀); forewing ground colour brown; broad basal patch pale yellow to yellow, crossed by longitudinal reddish brown fascia running up to postmedian area; costal field reddish brown; median discoidal stigma rhomboid to elliptic, pale yellow, filled with reddish brown; cubital and dorsal pale yellow patches forming with median discoidal stigma an oblique band disrupted at veins; postmedian patch roughly triangular, reddish brown, more or less speckled with pale yellow, with pale yellow blotch at costa; subterminal line pale yellow, conspicuously incurved inwards at CuA2; subterminal field marked with reddish brown; fringe brown, with pale yellow dots. Hindwing pale brown. Forelegs brown. Midlegs brown, inner side pale yellow. Hindlegs brown, tibia marked with pale yellow.
Female genitalia (N = 6) (Fig. 100). Anterior apophyses with small dorsal bump at posterior 1/3. Antrum sclerotisation short, ca. as wide as long. Ductus bursae relatively short, nearly straight. Corpus bursae globular and reticulated but in some specimens with a rounded pouch-like extension, with well-marked sclerotisation between thorn and ductus bursae opening, medially with faintly sclerotised band. Thorn sabrelike, curved, with small dents pointing toward thorn base.
Distribution. Known from the Luzon and Mindoro islands (Philippines), between sea level and 1,150 m.
DNA-barcoding. Specimens from Luzon and Leyte show an K2P-distance of 3.4-3.9% and are recovered in two MOTUs.
Etymology. Named after Mount Isarog (1966 m), a volcano on Luzon Island (Philippines) where the species is found.
Remarks. Minor differences were observed in male and female genitalia of the specimens from Leyte. However, with only one male from Leyte on hand, we refrained from drawing further conclusions on its species status here. Diagnosis. Hoploscopa ypsilon sp. nov. displays well-marked white coloured markings on the forewing. Median discoidal stigma and cubital patch are Y-shaped, postmedian patch is triangular and subterminal line is conspicuously incurved inwards at CuA2. In male genitalia, the gnathos forms a short, broadly rounded projection. Similar species. Hoploscopa danaoensis sp. nov. This species displays pale yellow markings on the forewing, the postmedian area is more or less broadly suffused with pale yellow, with subterminal line only slightly incurved inwards at CuA2. In male genitalia, H. danaoensis sp. nov. has a tongue-shaped projection ca. half the uncus length. Female genitalia are very similar to those of H. danaoensis sp. nov. and cannot be confidently separated.
Etymology. The species name ypsilon, for the Greek letter Y, refers to the shape of the median markings on the forewing. Description. Head. Antennae dorsally with brown scales. Proboscis pale brown, of a darker brown at base. Maxillary palpi brown, basally pale yellow, inner side pale brown. Labial palpi brown, ventro-basally pale yellow, inner side speckled with pale yellow.
Distribution. Known from Lake Danao (650 m Thorax (Fig. 20). Collar white. Forewing length: 10 mm; forewing ground colour brown; basal longitudinal red fascia dorsally edged with pale yellow, running up to postmedian area; costal field reddish brown; median discoidal stigma v-shaped, white, forming oblique streak extending down to dorsal area; postmedian patch reddish brown speckled with white, with costal white blotch; postmedian line marked on costal half; postmedian suffusion white; subterminal line white; subterminal field broadly marked with reddish brown; margin brown, chequered with pale yellow. Hindwing pale brown. Fore-and midlegs brown to bronze. Hindlegs with femur bronze, tibia pale yellow speckled with bronze, tarsi bronze.
Distribution. Recorded from West Java (Indonesia), at an altitude of 1,300 m. Remarks. Nuss (1998)  Diagnosis. Hoploscopa brunnealis displays a forewing pattern very similar to that of H. danaoensis sp. nov., with a median pale yellow Y, a postmedian pale yellow patch invaded with brown or tawny and a pale yellow subterminal line. In female genitalia, the antrum sclerotisation is short, the ductus bursae is short, straight, and the corpus bursae is pear-shaped, with a small straight thorn. Male genitalia are not known.
Description. Head. Antennae dorsally with bronze scales. Proboscis pale brown to pale yellow. Maxillary palpi brown, base and inner side pale yellow. Labial palpi brown, ventral base and inner side pale yellow. Thorax (Fig. 21). Forewing length: 10 mm; ground colour brown; basal dash dark brown distally pale yellow; costal field tawny; median discoidal stigma and median cubital patch forming together a pale yellow-coloured Y, with basal and distal edges dark brown; postmedian patch tawny speckled with pale yellow, distally pale yellow, with costal pale yellow blotch; postmedian area suffused with pale yellow; subterminal line pale yellow; fringes brown, with pale yellow dots.
Distribution. Recorded from West Java (Indonesia) at an altitude of 1,300 m. Remarks. Nuss (1998)  Diagnosis. The large oblique elliptic white patch at base of the forewing, the median trapezoid and postmedian semi-elliptic tawny patches are unique to H. ocellata. In female genitalia, the long, wrinkled and multicoiled ductus bursae, the ovoid corpus bursae bearing long acanthae, and the long glabrous thorn unequivocally segregates H. ocellata from other Hoploscopa species. Male unknown.
Male genitalia. Not known.
Distribution. Recorded from Bacan island in the Moluccas (Indonesia). Remarks. Nuss (1998)   Diagnosis. The white-coloured basal patch and median discoidal stigma as well as the oblique postmedian line allows separation of H. quadripuncta from its congeneric species. In male genitalia, uncus apex is cuneate and gnathos projection is ogiveshaped. In female genitalia, the thorn of the corpus bursae is long, slender, curved, and shows a well-marked rounded sclerotisation at its base.
Female genitalia (Fig. 105). Anterior apophyses with tip pointing upward at posterior 1/3. Antrum with a narrow weakly sclerotised ring. Ductus bursae of medium length, straight. Corpus bursae ovoid, posterior half reticulated, anterior half membranous, sclerotised between thorn and corpus opening, medially with faintly sclerotised band. Thorn long, curved, with small dents pointing toward thorn base on posterior edge, basally with conspicuous outwardly directed plump extension.
Distribution. Recorded from the Oetakwa River (Papua: Indonesia) in New Guinea, also known from the Oro Province (Papua New Guinea) and the Moluccas (Indonesia), at altitudes between 500 m and 1,000 m.
Remarks. Nuss (1998) transferred this species from Eudorina to Hoploscopa. Other specimens examined from the Moluccas and Papua New Guinea display the same pattern as the holotype from Western New Guinea, however a small difference in the shape of the gnathos projection is observed: it is pointed in the holotype, while it is rounded in the other specimens. Unfortunately, no molecular investigations were done on the holotype. Diagnosis. The forewings of H. semifascia display an antemedian dark brown fascia edged with pale yellow and a marked postmedian pale yellow patch. Median markings are completely lacking. In female genitalia, the thorn is long, thin, curved in distal half, with a small, well-marked sclerotisation at its base. Male unknown.

Hoploscopa semifascia (Hampson, 1919)
Similar species. No similar species known. Description. Head. Antennae dorsally striped with bronze and brown scales. Proboscis pale yellow. Maxillary palpi brown, base and inner side pale yellow. Labial palpi brown, base pale yellow, inner side pale yellow to pale brown. Thorax (Fig. 24). Collar white. Forewing length: 7 mm; forewing ground colour brown; antemedian broad transversal dark brown fascia, basal edge pale yellow, incurved inwardly on costal half, distal edge oblique, pale yellow; postmedian elongated pale yellow patch with dark brown edges; fringes brown, with pale yellow spots. Hind-wing pale brown. Forelegs brown. Midlegs brown; tarsi pale brown. Hindlegs with femur brown; tibia pale yellow, speckled with brown; tarsi pale brown.
Distribution. Recorded from Fak-Fak (Papua, Indonesia), also known from the Southern Highlands and the Morobe Province (Papua New Guinea) at altitudes between 550 m and 1,400 m.
DNA barcoding. The K2P-distance between specimens MTD LEP3158 from the Morobe Province and MTD8242 from the Southern Highlands (Papua New Guinea) is 0.5%.
Remarks. Nuss (1998)  Diagnosis. Hoploscopa subvariegata displays four roughly triangular yellow spots filled with reddish brown on forewing costa. In male genitalia, uncus is long and slender with truncate apex, gnathos projection is tongue-shaped, reaching 1/3 of uncus length, and dorsal margin of the valva is conspicuously protruded dorsad. In female genitalia, ductus bursae is long and straight, corpus bursae is large, globular and bears a small, slightly curved thorn with leaf-shaped sclerotisation at its base. Similar species. Hoploscopa persimilis. The latter species lacks the subterminal triangular yellow blotch at forewing costa. In the male genitalia of H. persimilis, uncus apex is duck-shaped, gnathos projection forms a small triangular tip, and valva dorsal margin is not protruded. Female of H. persimilis is not known.
Distribution. Recorded from the Angabunga River in the Central Province (Papua New Guinea), also known from Mount Kaindi, Morobe Province, at altitudes between 2,000 m and 2,360 m.
Remarks. Nuss (1998)  Diagnosis. Hoploscopa persimilis displays three nearly triangular yellow spots filled with reddish brown on the costa of the forewing. In male genitalia, uncus is long, slender, with duck-shaped apex, gnathos is projected into a small, triangular pointed tip, and valva dorsal margin is conspicuously convex. Female genitalia not known. Similar species. Hoploscopa subvariegata (q.v.). Description. Head. Not examined. Thorax (Fig. 26). Collar pale yellow. Forewing length: 9 mm; forewing ground colour brown; basal patch elongated, reddish brown, basally crossed by transverse pale yellow streak; costal field reddish brown; median discoidal stigma trapezoid, pale yellow, with basal and distal edges reddish brown; postmedian patch triangular, pale yellow and reddish brown, distally with costal pale yellow blotch; subterminal field tawny; fringes brown with pale yellow spots. Hindwing pale yellow. Legs missing or badly preserved.
Female genitalia. Not known. Distribution. Described from the Oetakwa River on New Guinea (Indonesia: Papua), at an altitude of 1,000 m.
Remarks. Nuss (1998)  Diagnosis. Hoploscopa diffusa displays a brown forewing with postmedian patch, part of postmedian area and subterminal line white coloured, and fringes marked with white dots. In male genitalia, gnathos projection is triangular with rounded tip, valva dorsal margin is conspicuously protruded and juxta is tongueshaped. Female unknown.
Thorax (Fig. 30). Collar white. Forewing length: 9 mm; forewing ground colour dark brown; small white dots scattered in basal and median area; postmedian patch white, filled with brown, with white costal blotch; postmedian line marked on costal half; postmedian area medially suffused with white scales; subterminal line white, shifted distally at M1; margin with white and black spots; fringe brown, with white dots; hindwing pale brown. Fore-and midlegs brown; tibia and tarsi segments distally white. Hindlegs similar, of a lighter brown.
Female genitalia. Not known.

Distribution. Recorded from Fergusson Island (Papua New Guinea).
Remarks. Nuss (1998)  Diagnosis. The large median dorsal patch with costal margin rounded on the forewing is unique to H. triangulifera. In male genitalia, the gnathos shows no distal projection, and the saccus is broad, quadrangular. In female genitalia, ductus bursae is short and slender, corpus bursae is large, globular, and bears a long curved thorn.
Similar species. No similar species known. Description. Head. Antennae dorsally pale yellow. Proboscis brown. Maxillary palpi brown, inner side pale brown. Labial palpi brown, ventral base pale yellow, inner side pale brown. Thorax (Fig. 31). Thorax brown, dorsally pale yellow. Collar pale yellow. Forewing length: 7-8 mm (♂), 8-9 mm (♀); forewing ground colour brown; basal and distal discoidal stigmata dark brown; median discoidal stigma forming roughly defined white X; median cubital patch elliptic, dark brown with white edges; large median dark brown patch on dorsum with rounded pale yellow costal margin; postmedian patch triangular, dark brown; postmedian line pale yellow, marked on costal half; pale yellow and brown postmedian fascia on costal half in some specimens, edged distally by white subterminal line; fringes brown with white spots. Hindwing pale brown. Forelegs bronze, tarsi speckled with pale yellow. Midlegs with femur bronze; tibia brown, distally pale yellow; tarsi bronze. Hindlegs with femur bronze; tibia brown, basally and distally pale yellow; tarsi bronze.
Distribution. Known from the Madang, the Northern and the Milne Bay Provinces (Papua New Guinea) at altitudes of ca. 1,600 m.
Remarks. Nuss (1998) transferred this species from Eudorina to Hoploscopa. The handwritten label from Hampson stipulates that the type is a male, while the abdomen dissected shows it is a female. The abdomen was re-affixed to the specimen, suggesting that either the sex wasn't identified properly by Hampson or that the wrong abdomen was reaffixed to the specimen. Diagnosis. The forewings of H. astrapias display a median cubital triangular white patch and a bean-shaped postmedian patch red with yellow edges, distally with thick white streak. In male genitalia, the uncus is slender, elongated, the gnathos is reduced to a ring without posterior projection and the valva is slender, with an evenly rounded apex. In female genitalia, the corpus bursae is large, spherical, and bears a large, straight, glabrous thorn. Similar species. Hoploscopa anamesa, H. nauticorum but the latter can be separated from H. astrapias and H. anamesa based on the forewing: median markings form an elongated white streak running down to dorsal area, disrupted at veins (forming roughly triangular white patch not extending beyond CuA2 in the two other species), and the postmedian patch is quadrangular. In male genitalia, the bristles at uncus apex observed in H. nauticorum are absent or reduced in H. astrapias and H. anamesa, and the valva dorsal margin is strongly produced dorsad on basal half, with a more pointed apex in H. nauticorum (dorsal margin slightly convex in two other species, apex evenly rounded). Forewing and male genitalia of H. astrapias and H. anamesa do not provide unambiguous diagnostic characters to separate them. Median cubital patch of H. anamesa is slightly thicker than that of H. astrapias in specimens examined. Female genitalia allow clear segregation of these two species: antrum is membranous in H. anamesa, while it is lightly sclerotised, twice as long as broad in H. astrapias, and ductus bursae is long, gently coiled twice in H. anamesa, while it is short and broadly curved in H. astrapias.

Hoploscopa astrapias
Description. Head. Antennae dorsally with brown scales. Proboscis white to pale yellow. Maxillary palpi brown, base and inner side pale brown. Labial palpi brown, ventro-basally pale yellow to white. Thorax (Fig. 27). Collar white. Forewing length: 10-11 mm (♂ & ♀); forewing ground colour brown; basal yellow streak along 1A+2A, abutted with cubital reddish brown fascia running up to median area, disrupted by median cubital patch; costal field reddish brown; median discoidal stigma trapezoid, reddish brown, edged basally and distally with yellow, median cubital patch triangular, snow white, together with median discoidal stigma forming a canine tooth shape; post-median patch bean-shaped, reddish brown with yellow edges, with thick snow white streak abutting dorsally; postmedian area suffused with reddish brown; subterminal line white, not reaching dorsum; subterminal field broadly marked with reddish brown; fringes brown. Hindwing pale yellow, bronze toward distal margin. Forelegs brown. Midlegs brown to bronze; tibia medially white. Hindlegs brown to bronze, tibia base dorsally pale yellow.
Distribution. Known from the island of Viti Levu (Fiji), at altitudes between 0 and 800 m.
Phylogenetic relationships. Hoploscopa astrapias, H. anamesa, and H. nauticorum are recovered together in the ML analysis of the COI barcode (BS = 61). This topology is congruent with the morphology of these three species: the forewings display a snow white transversal median line and postmedian streak; in male genitalia, the uncus is elongated, slender, the gnathos is reduced to a sclerotised band without posterior projection, the phallus bears a large, flattened, spatula-shaped cornutus; in female genitalia, the corpus bursae is globular and bears a long and slender straight thorn. Within this clade, a close relationship between H. astrapias and H. anamesa is supported by both morphology and molecular data (BS = 78). Diagnosis. Hoploscopa anamesa (Fig. 28) display forewing and male genitalia similar to those of H. astrapias. Female genitalia differ from those of H. astrapias in following characters: the antrum is not sclerotised, and the ductus bursae is long and coiled. Corpus bursae and thorn similar to H. astrapias. Similar species. Hoploscopa astrapias and H. nauticorum (q.v.). Description. Head. Antennae dorsally with brown scales. Proboscis white to pale yellow. Maxillary palpi brown, base and inner side pale brown. Labial palpi brown, ventro-basally pale yellow to white.
Distribution. Known from the Fiji (see Remarks), the Aneityum and Tanna islands on Vanuatu, at an altitude of ca. 350 m.
Remarks. Nuss (1998)  Thorax (Fig. 29). Collar pale brown. Forewing length: 10-11 mm; forewing ground colour dark brown; basal yellow streak along 1A + 2A, abutted to cubital red fascia running up to postmedian area, disrupted by median cubital patch; small reddish blotch at base of cell, with minute snow white dot; median discoidal stigma trapezoid, red, basally and distally edged with yellow, with snow white streak abutting dorsally, running down to dorsal area, disrupted at veins; postmedian patch quadrangular, red, dorso-distally with thick snow white streak; subterminal line white, indented inwardly at CuA2; subterminal field red; fringe brown. Hindwing pale yellow, bronze toward distal margin. Forelegs brown, tarsi speckled with pale yellow. Midlegs brown; tibia base speckled with pale yellow. Hindlegs brown.
Female genitalia. Not investigated. Distribution. Known from the island of Upolu on Samoa, at an altitude of ca. 600 m. Phylogenetic relationships. Hoploscopa nauticorum is recovered as sister to H. astrapias and H. anamesa in the ML analysis of the COI barcode (BS = 61). See H. astrapias for further remarks.
Diagnosis. Hoploscopa anacantha sp. nov. displays dark brown forewings with basal, median and postmedian white streak-like markings. In male genitalia, the gnathos projection forms a conspicuous triangle reaching half the length of the uncus. In female genitalia, the elongated bulging sclerotisation without thorn on corpus bursae is unique to this species.
Distribution. Known from North Sulawesi (Indonesia) at altitudes between 1,000 m and 1,200 m.
Phylogenetic relationships. This species is recovered sister to another similar but yet undescribed Hoploscopa species (DNA voucher MTD8238) from North Sulawesi (BS=67).
Etymology. From the Greek a-, an-, without, and acantha, spine, referring to the absence of thorn on the corpus bursae of female genitalia. This name was suggested by Francesca Vegliante. Diagnosis. Hoploscopa kelama sp. nov. displays basal and distal discoidal dark brown spots edged with pale yellow, as well as a pale yellow cubital double line on the forewing. In female genitalia, the very small thorn located at the corpus bursae opening is unique to this species. Similar species. Hoploscopa boleta sp. nov., H. pseudometacrossa sp. nov. Both species display light brown median discoidal stigma and postmedian patch and lack the double pale yellow cubital line observed in H. kelama sp. nov. Examination of genitalia allows unambiguous separation of these species from H. kelama sp. nov.: male genitalia of both species show a slender uncus bearing thick setae, ventrally marked with three small ridges, and female genitalia show a larger thorn located on posterior half of corpus bursae.
Thorax (Fig. 33). Collar white. Forewing length: 8-9.5 mm (♂ & ♀); forewing ground colour brown to dark brown; basal dash black at base, distally pale yellow; basal discoidal patch crescent-shaped, pale yellow, filled with dark brown; distal discoidal patch dark brown, quadrangular, with basal and distal edges pale yellow; double cubital line running from median area distally to tornus, pale yellow; postmedian costal blotch small, pale yellow; subterminal line pale yellow, barely or not marked on its middle fringes brown, with pale yellow dots. Hindwing pale brown. Forelegs brown. Mid-and hindlegs with femur brown; tibia dark brown, distally pale yellow; tarsi brown.
Distribution. Known from North Sulawesi (Indonesia) at altitudes between 1,000 m and 1,200 m.
Phylogenetic relationships. The ML analysis of the COI barcode recovered H. kelama sp. nov. in a clade with H. albomaculata sp. nov. and H. ignitamaculae sp. nov., but without significant support (BS = 30). These three species vary greatly in their wing pattern but show similarities in the morphology of the genitalia: the uncus apex is duck-shaped, with a small tip on its middle in male genitalia; in female genitalia, the ductus bursae is long and broad, the corpus bursae is globular, with an elongated, wellmarked sclerotisation. Hoploscopa kelama sp. nov. and H. ignitamaculae sp. nov. share a small-sized thorn on corpus bursae. Hoploscopa anacantha sp. nov. shares a wing pattern similar to that of H. albomaculata sp. nov., and displays an elongated, marked sclerotisation like in the other three species above, although without thorn.
Etymology. The species epithet kelama comes from the Indonesian "kelam" meaning dark, referring to the dark patterns of the forewing. Diagnosis. The forewings of H. ignitamaculae sp. nov. display marked basal, median and postmedian reddish orange markings edged with yellow; postmedian patch with undulated basal margin. In male genitalia, the uncus is medially widened, its apex is duck-shaped, ventrally with small bump, and the gnathos is projected into a short, pointed tip. In female genitalia, the well-defined elongated sclerotisation of the corpus bursae with the small slightly curved thorn on its middle is unique to this species.

Similar species.
Hoploscopa isarogensis sp. nov. Forewing markings of H. isarogensis sp. nov. tend more to red. Postmedian patch is triangular with a straight basal margin, and subterminal line is often markedly pale yellow. Examination of genitalia allow unambiguous separation of these species.
Description. Head. Antennae dorsally with bronze scales. Proboscis white to pale yellow. Maxillary palpi brown, basally pale yellow. Labial palpi brown, ventral base and inner side pale yellow.
Female genitalia (N = 2) (Fig. 113). Anterior apophyses with dorsal bump at posterior 1/3. Antrum sclerotisation short, as long as wide. Ductus bursae of medium length, broad, bent at anterior and posterior 1/4. Corpus bursae globular, posterior half reticulated, anterior half with erected papillae, with elongated sclerotised area running from corpus opening to its middle. Thorn on middle of sclerotisation, very small, slightly curved, with dents pointing toward thorn base.
Distribution. Known from North Sulawesi (Indonesia) at altitudes between 300 m and 1,150 m. Phylogenetic relationships. See H. kelama sp. nov. Etymology. From the Latin ignitus, set on fire, and macula, spot or blot, in reference to the red and yellow spots of the forewing.
Remarks. Specimen MTD8238 shows a forewing pattern resembling that of H. ignitamaculae sp. nov., but with markings reduced. However, the thorn is absent in female genitalia.    Diagnosis. Hoploscopa albomaculata sp. nov. is unique by virtue of its median trapezoid and postmedian streak-like white patches on the forewing. In male genitalia, the gnathos projection is reduced to a small triangular tip, and the juxta is elongated, slender, with a deeply indented apex. In female genitalia, ductus bursae is long, broad and curvy, and corpus bursae displays well-delimited sclerotised band and a straight thorn.
Similar species. No similar species known. Description. Head. Antennae dorsally with brown scales. Proboscis pale yellow to pale brown. Maxillary palpi brown, base and inner side pale yellow. Labial palpi brown, ventral base and inner side pale yellow.
Diagnosis. The forewings of H. sumatrensis sp. nov. display a pale yellow crescentshaped median discoidal stigma filled with reddish brown, as well as a postmedian area broadly suffused with pale yellow. In male genitalia, the gnathos projection is slender, ca. 4/5 of uncus length, with a tongue-shaped apex, and the juxta has a rounded base and a notched apex. In female genitalia, the short and slender ductus bursae bent before corpus and the pear-shaped corpus bursae with small straight thorn resemble those of other Hoploscopa species, e.g., H. danaoensis sp. nov. and H. parvimacula sp. nov. Similar species. Hoploscopa pangrangoensis sp. nov. (q.v.). Description. Head. Antennae dorsally with brown scales. Proboscis pale yellow to brown. Maxillary palpi brown, base and inner side pale yellow. Labial palpi brown, ventral base and inner side pale yellow.
Distribution. Known from North Sumatra (Indonesia). DNA barcoding. Two MOTUs are found in morphologically identical specimens from the same locality. The K2P-distance between the two MOTUs is 4.1-6%. The first MOTU, which is the one of the type material, shows an intraspecific variation of 0.7%. The second MOTU is found in three females (samples BC MTD LEP01422, MTD8258 and MTD8261) and shows an intraspecific divergence of 0.6%.
Etymology. Named after the island of Sumatra where the species is encountered. Remarks. Future examination of male specimens from the second lineage will help to determine whether or not it represents a separate species.   Diagnosis. The forewings of H. mallyi sp. nov. display pale yellow crescent-shaped median discoidal stigma filled with red, together with median cubital patch forming a Y. In male genitalia, the gnathos projection is long, slender, reaching circa 2/3 of the uncus length, and the juxta is ogive-shaped, with a slightly indented apex. In female genitalia, the ductus bursae is long, broad, curved twice, and the corpus bursae is small, globular, with one short curved thorn.

Hoploscopa mallyi
Similar species. Hoploscopa agtuuganonensis sp. nov., H. gracilis sp. nov. In H. agtuuganonensis sp. nov., median discoidal stigma together with cubital and dorsal patches forms a band progressively narrowing toward dorsum (median cubital patch reduced to small streak, median dorsal patch absent in H. mallyi sp. nov.). Hoploscopa gracilis sp. nov. displays wing pattern similar to H. mallyi sp. nov. and can be best separated by examination of genitalia: the uncus and valva are slenderer, the valva apex is pointed (rounded in H. mallyi sp. nov.) and the juxta displays a broad duck beak-shaped apex. In female genitalia of H. gracilis sp. nov., the ductus bursae shows a narrow loop at posterior 1/3, and the thorn is less curved.
Description. Head. Antennae dorsally with brown scales. Proboscis brown to pale brown scaled. Maxillary palpi brown, base and inner side pale yellow. Labial palpi brown, ventral base and inner side pale yellow.
Thorax (Fig. 37). Collar pale yellow. Forewing length: 10-12 mm (♂), 11-12 mm (♀); forewing ground colour brown; basal longitudinal red streak basally edged with pale yellow; costal field red; median discoidal stigma crescent-shaped, pale yellow, filled with red; median cubital patch streak-like, pale yellow, together with discoidal stigma forming a Y; postmedian patch triangular, red, basal edge partially pale yellow, with pale yellow blotch at costa; postmedian area suffused with pale yellow and iridescent scales; subterminal field marked with red; fringes brown with pale yellow dots. Hindwing pale brown. Forelegs brown. Midlegs brown; femur and tibia distally pale yellow. Hindlegs with femur brown; tibia pale yellow speckled with brown; tarsi pale yellow to pale brown.
Distribution. Known from the slopes of Mount Kinabalu on Borneo, at altitudes between 1,600 m and 2,000 m.
Phylogenetic relationships. This species is recovered in a clade with H. agtuuganonensis sp. nov. and H. gracilis sp. nov. in the ML analysis of the COI barcode (BS = 92). The morphology of these species is in agreement with the molecular find-ings: in male genitalia, gnathos projection is long and slender; in female genitalia, the antrum sclerotisation is short, the ductus bursae is relatively broad, looped or with conspicuous curves, and the corpus bursae is small, globular.

Hoploscopa gracilis
Diagnosis. Hoploscopa gracilis sp. nov. displays pale yellow crescent-shaped median discoidal stigma filled with reddish brown, together with median cubital patch forming a Y. In male genitalia, uncus is long and slender, valva is slender with pointed apex, and juxta displays a broad duck beak-shaped apex. In female genitalia, the ductus bursae is long, forming one loop on its middle, and the corpus bursae is small, globular, with one short slightly curved thorn. Similar species. Hoploscopa agtuuganonensis sp. nov., H. mallyi sp. nov. (q.v.). Median markings of the forewing form a band progressively narrowing toward dorsum in H. agtuuganonensis sp. nov. In male genitalia, uncus of H. agtuuganonensis sp. nov. is larger, valva has a rounded apex and juxta apex is tongue-shaped. Female genitalia of H. agtuuganonensis sp. nov. are very similar to those of H. gracilis sp. nov. but differ by a more marked corpus sclerotisation and a thicker thorn.
Description. Head. Antennae dorsally ochreous to brown. Proboscis pale yellow. Maxillary palpi brown, base and inner side pale yellow. Labial palpi brown, ventral base and inner side pale yellow. Thorax (Fig. 38). Collar pale yellow. Forewing length: 9 mm (♂), 11 mm (♀); forewing ground colour brown; broad elongated basal patch reddish brown with basal edge pale yellow, distally running up to median cubital patch; costal field reddish brown; median discoidal stigma crescent-shaped, pale yellow, filled with reddish brown; median cubital patch streak-like, pale yellow, disrupted at 1A+2A, together with median discoidal stigma forming a Y; postmedian triangular patch reddish brown, speckled with pale yellow, with pale yellow blotch at costa; postmedian area suffused with a mix of pale yellow and iridescent scales; subterminal field marked with reddish brown; fringes brown, with pale yellow dots. Hindwing pale yellow. Forelegs brown. Midlegs brown; tibia distally pale yellow. Hindlegs brown, tibia pale yellow speckled with brown.
Distribution. Known from North Sumatra, at altitudes between ca. 1,200 m to 1,900 m.
DNA barcoding. The species shows an intraspecific variation of 0.9% Phylogenetic relationships. See H. mallyi sp. nov. Etymology. From the Latin gracilis, slender, narrow, referring to the slender shape of the uncus, the gnathos projection, and the valva in male genitalia. Diagnosis. In H. agtuuganonensis sp. nov., median markings of the forewing form together a pale yellow band progressively narrowing toward dorsum, faintly disrupted with red at veins. In male genitalia, uncus is long, slender, narrowed at apical 1/4 and gnathos projection is long and slender. In female genitalia, ductus bursae is long, broad, with one loop, and corpus bursae is small, globular, with one short plump thorn. Similar species. Hoploscopa isarogensis sp. nov. (q.v.), H. mallyi sp. nov. (q.v.), H. gracilis sp. nov. (q.v.).

Hoploscopa agtuuganonensis
Description. Head. Antennae dorsally striped with brown and pale yellow scales. Proboscis pale yellow to pale brown. Maxillary palpi brown, base and inner side pale yellow. Labial palpi brown, ventral base and inner side pale yellow.
Thorax (Fig. 39). Collar pale yellow to white. Forewing length: 8-9 mm (♂), 9 mm (♀); forewing ground colour brown; basal reddish brown quadrangular patch basally edged with pale yellow, crossed by longitudinal pale yellow streak, with median cubital and dorsal patches abutting dorsally; costal field reddish brown; median discoidal stigma trapezoid, pale yellow, filled with reddish brown, basal and distal edges marked with reddish brown, together with two pale yellow cubital patches forming a broad band narrowing toward dorsum; postmedian patch triangular, reddish brown, speckled with pale yellow, with pale yellow blotch at costa; postmedian area suffused with pale yellow; subterminal line pale yellow; subterminal field reddish brown; fringes brown, with large pale yellow spots. Hindwing pale brown. Fore-and midlegs brown. Hindlegs with femur brown; tibia pale yellow speckled with brown; tarsi brown.
Female genitalia (N = 3) (Fig. 118). Anterior apophyses with dorsal tip at basal 1/3. Antrum sclerotisation short, ca. as long as broad. Ductus bursae long, broad, with one loop. Corpus bursae small, globular, reticulated, with roughly defined sclerotisation between thorn and corpus opening. Thorn plump, straight, with small dents pointing toward thorn base, basally with small outwardly projected extension. Diagnosis. The forewings of H. boleta sp. nov. display median discoidal stigma and postmedian patch of a lighter brown, the latter with a well-marked pale yellow blotch at costa. In male genitalia, the conspicuous basal lateral projections of the juxta resemble the shape of a mushroom. Female genitalia are somewhat similar to those of other Hoploscopa species, e.g., H. parvimacula sp. nov., and H. danaoensis sp. nov., with a short straight ductus bursae, a pear-shaped corpus bursae, and a small straight thorn.
Similar species. Hoploscopa pseudometacrossa sp. nov., to a lesser extent H. kelama sp. nov. (q.v.). In H. pseudometacrossa sp. nov., forewing median discoidal stigma and postmedian patch are barely marked, and postmedian pale yellow blotch at costa is much smaller. In male genitalia, gnathos of H. pseudometacrossa sp. nov. displays a thumb-like projection, juxta shows less prominent lateral projections and an indented apex. In female genitalia, corpus bursae is larger than that of H. boleta sp. nov.
Female genitalia (N = 2) (Fig. 119). Anterior apophyses with dorsal bump at posterior 1/3. Antrum sclerotisation twice as long as broad. Ductus bursae short, nearly straight. Corpus bursae globular, posterior half reticulated, medially with erect acanthae, anterior half membranous, with sclerotisation between thorn and corpus opening, medially with a sclerotised band. Thorn straight, with small dents pointing toward thorn base. Diagnosis. The forewings of H. pseudometacrossa sp. nov. display barely marked median discoidal stigma and postmedian patch, while basal and distal discoidal patches are dark brown. Male hindwing displays an androconial organ on the dorsum. In male genitalia, the gnathos shows a short, thumb-like projection, and the juxta is slender, with notched apex. In female genitalia, the ductus bursae is relatively short-sized, the corpus bursae is large, spherical, with a small straight thorn. Similar species. Hoploscopa boleta sp. nov. (q.v.), H. metacrossa, H. kelama sp. nov. (q.v.). Forewing median discoidal stigma forms a pale yellow Y with median cubital patch in H. metacrossa. In male genitalia, gnathos projection of H. metacrossa is thinner, valva is larger, and juxta apex is rounded. Female genitalia are very similar, but the antrum sclerotisation is shorter, and the corpus bursae smaller in H. metacrossa.
Description. Head. Antennae dorsally with bronze to brown scales. Proboscis pale yellow to brown. Maxillary palpi brown, base and inner side pale yellow. Labial palpi brown, ventro-basally pale yellow. Thorax (Fig. 41). Collar pale yellow. Forewing length: 9-10 mm (♂ & ♀); forewing ground colour brown; basal dash dark brown, distally pale yellow; basal and distal discoidal stigma quadrangular, dark brown; median discoidal stigma faded, basally and distally thinly edged with pale yellow; postmedian patch faded, with distal edge dark brown, at costa with pale yellow blotch; postmedian line marked on costal half; postmedian area suffused with pale yellow near costa; subterminal line pale yellow; fringe brown, with pale yellow dots. Hindwing pale brown; in males, presence of an androconial organ on the dorsum of the hindwing. Forelegs brown. Mid-and hindlegs with femur brown, tibia brown speckled with pale yellow, tarsi bronze.
Diagnosis. The forewings of H. metacrossa display Y-shaped median and a postmedian pale yellow patches observed in several other Hoploscopa species (e.g., H. danaoensis sp. nov.). Male hindwing displays an androconial organ on the dorsum. The strongly sclerotised tympanic drum in tympanal organs of males is only observed in this species. In male genitalia, gnathos is projected into a small, slender, tongueshaped tip, and valva is broad, with a rounded ventral margin. In female genitalia, antrum sclerotisation is as long as wide, ductus bursae is short, straight and the corpus bursae is large.
Similar species. Hoploscopa brunnealis, H. danaoensis sp. nov., H. kinabaluensis sp. nov. (q.v.), H. pseudometacrossa sp. nov. (q.v.). Males of H. metacrossa are easily separated from similar species (except H. pseudometacrossa sp. nov.) by the presence of scent organs on the hindwing, the sclerotised tympanic drum, and the characteristic spatulate uncus apex with ventral ridges. Female genitalia of the four above-mentioned species are similar, but ductus bursae is bent before corpus bursae in these species (nearly straight in H. metacrossa) and corpus bursae are smaller than that of H. metacrossa.
Description. Head. Antennae dorsally brown. Proboscis pale yellow to brown. Maxillary palpi brown, base and inner side pale yellow. Labial palpi brown, ventral base and inner side pale yellow.
Female genitalia (N = 3) (Fig. 121). Anterior apophyses with dorsal bump at posterior 1/3. Antrum sclerotisation short, as long as wide. Ductus bursae short, nearly straight. Corpus bursae spherical, reticulated, with sclerotisation between thorn and corpus opening and faintly marked sclerotisation medially. Thorn straight, with small dents pointing toward thorn base, basally with small outwardly projected extension. Diagnosis. This species shows a strong sexual dimorphism. In males, median and postmedian markings are white, and postmedian area is broadly suffused with white. In females, basal patch, median discoidal stigma and postmedian patch are reddish orange, and subterminal area is orange. Male hindwing displays an androconial organ on the dorsum. In male genitalia, gnathos projection is slender, ca. 1/3 of the uncus length, and juxta displays two conspicuous lateral projections at base, and a deeply indented apex. In female genitalia, the four loops formed by the ductus bursae is unique to this species.
Similar species. No similar species known. Description. Head. Antennae dorsally brown. Proboscis brown. Maxillary palpi brown, base and inner side pale yellow. Labial palpi brown, ventral base and inner side pale yellow.
Thorax (Figs 2,43,44). Collar brown. Forewing length: 9-10 mm (♂ & ♀); forewing ♂ ground colour brown; basal narrow white streak, distally marked with yellow and orange; median discoidal stigma white, filled with few red and yellow scales, together with cubital and dorsal patch forming a white Y; postmedian patch brown speckled with red, more or less marked, distally with costal white streak; postmedian line marked on costal half; postmedian area with broad white suffusion; subterminal line conspicuously incurved inward at CuA2, white; fringe chequered brown and pale yellow; forewing ♀ ( Fig. 44) ground colour brown; yellow basal patch crossed by orange streak; median trapezoid white patch, filled to a various extend with yellow and red scales, together with cubital and dorsal white patches forming a Y; postmedian patch roughly oval, reddish or-ange, basally edged with yellow, distally with costal white streak; postmedian line marked on costal half; postmedian area faintly speckled with white between postmedian patch and subterminal line; subterminal line white, incurved inwards at CuA2; subterminal field marked with reddish orange and yellow; fringe brown with pale yellow dots. Hindwing pale yellow; in males, presence of an androconial organ on the dorsum of the hindwing. Forelegs bronze. Midlegs with femur brown; tibia brown, distally pale yellow; tarsi bronze. Hindlegs with femur brown; tibia pale yellow, speckled with brown; tarsi bronze.
Abdomen. Male tergite I, II partially sclerotised; tergite III sclerotised, with two shallow depressions on each side of middle, bearing patches of modified scales. Sternum A8 posterior margin straight.
Female genitalia (N = 3) (Fig. 122). Anterior apophyses with dorsal bump at posterior 1/3. Antrum sclerotisation short. Ductus bursae very long, forming four conspicuous loops bent before corpus opening. Corpus bursae small, globular, reticulated, medially with light broad sclerotised patch, with weak rounded sclerotisation at corpus opening. Thorn gently curved, with small dents pointing toward thorn base, basally with small outwardly projected extension. Remarks. This species is the only one known from the genus to exhibit a pronounced sexual dimorphism.

Misplaced species
Hoploscopa mediobrunnea (De Joannis, 1929) from Vietnam has been provisionally placed in Hoploscopa by Nuss (1998). He justified his choice as follows: ""Since Eudorina Snellen is a homonym of Eudorina Ehrenberg, 1832 (Protozoa), I place E. mediobrunnea here preliminarily in Hoploscopa. With the white discocellular stigma and the white fasciata adjacent to the median space, the wing pattern of this species look similar to the Musotiminae genus Uthinia. In contrast to Uthinia, H. mediobrunnea shows porrect labial palpi, a long ductus bursae, and the ductus seminalis originates near the antrum. The corpus bursae shows three round signa, with inwardly directed tiny spines arising from their edges." (Nuss 1998). Its wing shape and the female genitalia are different from all Hoploscopa species we have examined to date, and we therefore assume that H. mediobrunnea belongs to a different genus.   Figure 123. RaxML analysis of the COI barcode dataset partitionned after codon positions. Node support estimated with 1000 thorough bootstrap replicates using the GTR+GAMMA substitution model. Columns on the right display species delimitation inferred from the morphology (red), the ABGD (pink), and GMYC (green) methods.

Discussion
Our iterative taxonomic revision resulted in the discovery and description of 26 new species as well as the redescription of 15 species. Thirty of 38 morphospecies were concordant with species delimitation based on the COI barcode. Distinct genetic divergence (i.e., more than 2%) among populations of different islands, with minor morphological differences in wing pattern or male genitalia, suggests reproductive isolation through allopatric effects. In H. matheae sp. nov. and H. parvimacula sp. nov., specimens from Borneo and from Malaysia Peninsula form two distinct MOTUs that deserve closer examination. Cryptic divergence between Borneo and Malayan Peninsula has been found in other studies, e.g., ants (Quek et al. 2007, Feldhaar et al. 2010, bats (Francis et al. 2010), and birds (Lohman et al. 2011). Similarly, specimens of H. danaoensis sp. nov. and H. isarogensis sp. nov. from different Philippines islands display distinct genetic differentiation suggesting allopatric differentiation. Cryptic diversity among populations of different Philippines islands has also been reported in birds (Lohman et al. 2010). We found specimens from Eastern New Guinea resembling the type specimens of H. quadripuncta and H. semifascia from Western New Guinea, but displaying minor differences in male genitalia in each case. This suggests potential east-west differentiation along the Cordillera, as observed in birds (Joseph et al. 2001). Molecular investigations of specimens from Western New Guinea should be performed to test this hypothesis. High DNA barcode divergence in geographically distant species reveals the need for a better sampling with higher geographical coverage to investigate both intraspecific and interspecific variation (Bergsten et al. 2012, Talavera et al. 2013. Two cases of high genetic divergence found in sympatric populations deserve closer attention. The first case involves H. sumatrensis sp. nov. from north Sumatra, where two mitochondrial lineages differ by 4.1-6%. In the second case, a divergence of 1.7-2.2% is observed in sympatric specimens of H. kinabaluensis sp. nov. on Mount Kinabalu. Closer examination of the morphology did not reveal any differences in either case. Inclusion of nuclear markers is needed in order to test if these mitochondrial lineages reflect different species or cases of introgression, symbiont infection, or maternal lineage sorting (Funk and Omland 2003, Hurst and Jiggins 2005, Harrison and Larson 2014. Notably, introgression has been reported in Lepidoptera (Zakharov et al. 2009;Cong et al. 2017), including Crambidae (Mally et al. 2018).
The investigated material more than doubled the number of Hoploscopa species. With 41 species now described, we still have not reached the saturation phase in species discovery for this group. For example, morphology of the female genitalia and molecular data support specimens MTD8238 and MTD8243 as different species, for which male specimens remain to be discovered. Similarly, molecular data and images of the habitus support specimen USNM_ENT_00739239 as a new but yet undescribed species. Unfortunately, this specimen could not be located (Scott Miller, pers. comm.). Additionally, the nine cases of morphospecies split into two or more MOTUs suggest putative new cryptic species and require further examination. In the NHMUK collection, we estimate at least a further 30 species await description. The presumed sister-group Perimeceta is probably less diverse, with four described species to date and an estimated twelve undescribed species in the collections of the NHMUK.
Hoploscopa is predominantly found in montane wet forests of South-East Asia at altitudes above 1,000 m, while species from higher latitudes tend to stretch their altitudinal distribution down to the lowlands, e.g., on the Philippines (H. danaoensis sp. nov., H. isarogensis sp. nov.) and the Melanesian islands (H. astrapias, H. anamesa, H. nauticorum). Mountain of South-East Asia are also home to other species-rich crambid groups, e.g., Glaucocharis, Micraglossa, and Scoparia (Gaskin 1985, Nuss 1998. Based on our data, we hypothesise isolation through elevation, higher specialisation, and seasonality as three potential factors driving the diversification of Hoploscopa. Isolation through elevation promotes species diversity and endemism (Steinbauer et al. 2016). The study of Merckx et al. (2015) on a broad range of taxonomic groups from Mount Kinabalu suggested endemic species on tropical mountains to originate either from long-distance dispersal (eccentric species) or from colonisation from local lowlands (centric species). All seven new species from Mount Kinabalu described in this paper do not share morphological similarities and no close relationships was recovered among them. On the contrary, H. mallyi sp. nov. from Mt Kinabalu forms a well-supported clade with H. agtuuganonensis sp. nov. from Mindanao and H. gracilis sp. nov. from Sumatra and represents a possible case of long-distance dispersal across islands. Colonisation from local lowland is also conceivable in Hoploscopa: H. cynodonta sp. nov. is reported at an altitude of 300 m in Brunei, and other Hoploscopa specimens have been collected at altitudes of 500 m at the foot of Mount Kinabalu (Schulze 2000). Secondly, on-site specialisation on host plant is another possible driver of speciation. Moths were shown to be high specialists in tropical mountains (Rodríguez-Castañeda et al. 2010). Larvae of Hoploscopa feed on ferns, which show a high species diversity in montane wet forests of South-East Asia (Ebihara and Kuo 2012). Current host plant records report five Hoploscopa species to feed on ferns belonging to three distinct genera (Miller et al. 2015;Mally et al. 2017). Ferns are scarcely used by Lepidoptera (Weintraub et al. 1995) and thus represent a large palette of potential larval food. Finally, there is a possible seasonality occurring in Hoploscopa species. The Hoploscopa larva described by Mally et al. (2017) and collected during a field trip on Mount Kinabalu did not match any of the eight species we collected there, although we cannot rule out the possibility that we simply did not collect it. Seasonality is observed in Pyraloidea from lowlands surrounding Mount Kinabalu (Schulze and Fiedler 2003) but no studies are available on the seasonality of moths at higher elevations in South-East Asia. Further systematic work on Hoploscopa, phylogeographic investigations, as well as accumulation of host plant records will assist in completing a more comprehensive picture of its diversity.
Appendix 1 Table A1. Collecting localities of the Hoploscopa specimens examined. GPS coordinates for historical localities were approximated with help of the online gazetteer https://geographic.org/geographic_names/index.html.