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Twenty-six new species of Hoploscopa 1
Twenty-six new species of Hoploscopa (Lepidoptera,
Crambidae) from South-East Asia revealed by
morphology and DNA barcoding
éo Léger1, Christian Kehlmaier2, Charles S. Vairappan3, Matthias Nuss2
1 Museum für Naturkunde – Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Invalidenstr., Berlin,
Germany 2 Senckenberg Museum für Tierkunde Dresden, Königsbrücker Landstr., Dresden, Germany 3Institu-
te for Tropical Biology and Conservation, Universiti Malaysia Sabah, 88400 Kota Kinabalu, Sabah, Malaysia
Corresponding author: éo Léger (theo.leger@mfn.berlin)
Academic editor: Colin Plant|Received 27 May 2019|Accepted 25 August 2019|Published 29 January 2020
http://zoobank.org/DBF339E5-EBBC-4619-9438-8359C769473F
Citation: Léger T, Kehlmaier C, Vairappan CS, Nuss M (2020) Twenty-six new species of Hoploscopa (Lepidoptera,
Crambidae) from South-East Asia revealed by morphology and DNA barcoding. ZooKeys 907: 1–99. https://doi.
org/10.3897/zookeys.907.36563
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 scientic collections. An iterative approach including
morphological and molecular characters was used in order to explore the diversity of Hoploscopa. e hith-
erto described species are revised, and descriptions authored by T. Léger and M. Nuss are provided for an
additional 26 new species: H. agtuuganonensis sp. nov., H. albipuncta sp. nov., H. albomaculata sp. nov.,
H. anacantha sp. nov., H. boleta sp. nov., H. cynodonta sp. nov., H. danaoensis sp. nov., H. gombongi sp.
no v., H. gracilis sp. nov., H. ignitamaculae sp. nov., H. isarogensis sp. nov., H. jubata sp. nov., H. kelama
sp. nov., H. kinabaluensis sp. nov., H. mallyi sp. nov., H. marijoweissae sp. nov., H. matheae sp. nov., H.
niveofascia sp. nov., H. pangrangoensis sp. nov., H. parvimacula sp. nov., H. pseudometacrossa sp. nov.,
H. sepanggi sp. nov., H. sumatrensis sp. nov., H. titika sp. nov., H. tonsepi sp. nov., H. ypsilon sp. nov.
Using a protocol specic for the amplication 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 reecting allopatric divergence, and
two further cases found in sympatry.
Keywords
Indo-malayan, integrative taxonomy, historic DNA, Melanesia, Pyraloidea, taxonomy
ZooKeys 907: 1–99 (2020)
doi: 10.3897/zookeys.907.36563
http://zookeys.pensoft.net
Copyright Théo Léger et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0),
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éo Léger et al. / ZooKeys 907: 1–99 (2020)
2
Table of Contents
Introduction ............................................................................................................. 3
Materials and methods ............................................................................................. 4
Material acquisition ........................................................................................... 4
Iterative approach ............................................................................................... 4
Molecular work .................................................................................................. 5
Data analyses ...................................................................................................... 5
Species delimitation ........................................................................................... 6
Systematic treatment .......................................................................................... 7
Results ...................................................................................................................... 7
Molecular dataset ............................................................................................... 7
Species delimitation ........................................................................................... 8
Systematics ......................................................................................................... 8
Hoploscopa Meyrick, 1886 ............................................................................ 8
Hoploscopa albipuncta Léger & Nuss, sp. nov. ....................................... 12
Hoploscopa matheae Léger & Nuss, sp. nov. .......................................... 15
Hoploscopa sepanggi Léger & Nuss, sp. nov. .......................................... 17
Hoploscopa cynodonta Léger & Nuss, sp. nov. ....................................... 18
Hoploscopa parvimacula Léger & Nuss, sp. nov. .................................... 20
Hoploscopa kinabaluensis Léger & Nuss, sp. nov. .................................. 22
Hoploscopa luteomacula Nuss, 1998 ...................................................... 24
Hoploscopa obliqua (Rothschild, 1915) ................................................. 25
Hoploscopa niveofascia Léger & Nuss, sp. nov. ...................................... 26
Hoploscopa gombongi Léger & Nuss, sp. nov. ........................................ 27
Hoploscopa tonsepi Léger & Nuss, sp. nov. ............................................ 28
Hoploscopa marijoweissae Léger & Nuss, sp. nov. .................................. 30
Hoploscopa titika Léger & Nuss, sp. nov. .............................................. 31
Hoploscopa pangrangoensis Léger & Nuss, sp. nov. ................................ 32
Hoploscopa isarogensis Léger & Nuss, sp. nov. ....................................... 33
Hoploscopa ypsilon Léger & Nuss, sp. nov. ............................................ 35
Hoploscopa danaoensis Léger & Nuss, sp. nov. ....................................... 36
Hoploscopa aurantiacalis (Snellen, 1895) ............................................... 39
Hoploscopa brunnealis (Snellen, 1895) .................................................. 40
Hoploscopa ocellata (Hampson, 1919) .................................................. 41
Hoploscopa quadripuncta (Rotschild, 1915) .......................................... 42
Hoploscopa semifascia (Hampson, 1919) ............................................... 43
Hoploscopa subvariegata (Rotschild, 1915) ............................................ 44
Hoploscopa persimilis (Rotschild, 1915) ................................................ 45
Hoploscopa diusa (Hampson, 1919) .................................................... 46
Hoploscopa triangulifera (Hampson, 1919) ........................................... 47
Hoploscopa astrapias Meyrick, 1886 ...................................................... 48
Hoploscopa anamesa Tams, 1935 ........................................................... 50
Hoploscopa nauticorum Tams, 1935 ...................................................... 51
Twenty-six new species of Hoploscopa 3
Hoploscopa anacantha Léger & Nuss, sp. nov. ....................................... 52
Hoploscopa kelama Léger & Nuss, sp. nov. ............................................ 53
Hoploscopa ignitamaculae Léger & Nuss, sp. nov. ................................. 55
Hoploscopa albomaculata Léger & Nuss, sp. nov. .................................. 63
Hoploscopa sumatrensis Léger & Nuss, sp. nov. ...................................... 64
Hoploscopa mallyi Léger & Nuss, sp. nov. ............................................. 65
Hoploscopa gracilis Léger & Nuss, sp. nov. ............................................ 68
Hoploscopa agtuuganonensis Léger & Nuss, sp. nov. .............................. 69
Hoploscopa boleta Léger & Nuss, sp. nov. .............................................. 70
Hoploscopa pseudometacrossa Léger & Nuss, sp. nov. ............................. 72
Hoploscopa metacrossa (Hampson, 1917) .............................................. 73
Hoploscopa jubata Léger & Nuss, sp. nov. ............................................. 75
Misplaced species ....................................................................................... 76
Discussion .............................................................................................................. 88
Acknowledgements ................................................................................................. 89
References .............................................................................................................. 90
Appendix ................................................................................................................ 95
Supplementary material 1 ....................................................................................... 98
Supplementary material 2 ....................................................................................... 98
Supplementary material 3 ....................................................................................... 98
Supplementary material 4 ....................................................................................... 99
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. e genus is distributed across the
Oriental and Australasian regions, ranging from the North of ailand eastwards to the
Samoa Islands, but is virtually absent from the tropical rainforests of Northern Queens-
land, 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. e 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 Perimec-
eta 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). e systematic
éo Léger et al. / ZooKeys 907: 1–99 (2020)
4
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 He-
liothelinae an ingroup of Scopariinae (Leraut 1980, Minet 1982, Munroe and Solis 1998).
e rst molecular-based phylogeny including representatives of Hoploscopa suggest it to
represent a separate lineage outside of Scopariinae + Crambinae (Léger et al. 2019).
is paper aims at establishing an iterative approach following Yeates et al. (2011)
for species discovery and description in Hoploscopa, serving as a rst step of a compre-
hensive revision of the genus. With an iterative approach using morphology and COI
barcode, we provide redescriptions for fteen of the 16 described species and describe
26 new species. Incongruence between morphology and molecular datasets, as well as
factors that possibly inuenced the evolution of the group are discussed.
Materials and methods
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 Mu-
seum 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, Mu-
nich (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. Speci-
mens 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 fol-
lowed the best practice outlined by Schlick-Steiner et al. (2010), here referred to as
“iterative taxonomy” (Yeates et al. 2011). is workow 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 mor-
phospecies, specimens with slightly diering wing pattern and/or of each collecting
locality were selected for amplication 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 diver-
gence 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.
Twenty-six new species of Hoploscopa 5
Molecular work
One hundred and fty-seven dried museum specimens were considered for DNA extrac-
tion. 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. e 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 meas-
ured with a NanoDropOne spectrophotometer. We followed a workow previously used
by other authors (Hebert et al. 2013, Mitchell 2015, Hundsdoerfer et al. 2017) by am-
plifying up to ve 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 rst step (recently collected samples only), amplication
of the whole COI barcode sequence (658 bp) was attempted with the primer pair Hy-
bLCO/HybNancy, each anked with universal primer tails (“Hyb-”) facilitating sequenc-
ing (Wahlberg and Wheat 2008). If amplication was not successful, we proceeded in the
second step with the amplication of fragments 1a (LepF1/K699) and 1b (f220/LepR1
or HybNancy). e latter reverse primer showed a slightly better performance with re-
spect to the amplication of fragment 1b and was preferentially used. If amplication of
fragments 1a or 1b in step two failed, we proceeded in step three with amplication of
fragments A, B, C (covering fragment 1a) and C, D, E (covering fragment 1b). PCR-mix
included 2 µl of BIORON complete buer 10× incl. Mg2+ (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 (Bi-
oron 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 lled up with water to nal
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 lled up with water
to nal 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 nal elongation at 72 °C for 10 min. Amplication 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 origi-
nal 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 amplication of the correct fragment, sequences were
éo Léger et al. / ZooKeys 907: 1–99 (2020)
6
blasted with the blastn program against the nucleotide collection (nt), as provided by
the National Center for Biotechnology Information (NCBI). In order to identify po-
tential 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 se-
quences 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 subse-
quently 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 dierent methods were used to investigate species delimitation in our molecular
dataset: Automatic Barcode Gap Discovery (ABGD) (Puillandre et al. 2012) and Gen-
eral Mixed Yule-coalescent method (GMYC) (Pons et al. 2006, Fujisawa and Barra-
clough 2013). ese 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 eect of non-compliant bar-
code sequences (< 487 bp) on the estimation of species in densely sampled clades (n >
2), where these sequences were agged 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 xed 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.
e ultrametric tree required for the GMYC analysis was generated using BEAST
1.10.4 (Suchard et al. 2018). e 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 1000th generation. Convergence was
checked on Tracer (Rambaut et al. 2018) and the rst 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 and
Barraclough 2013). e single-threshold option was used as suggested in Fujisawa
and Barraclough (2013).
Twenty-six new species of Hoploscopa 7
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 paren-
theses. e specimen depositories are reported with the use of the corresponding acronyms.
GPS coordinates for localities were retrieved from the online platform geographic.org (htt-
ps://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 conspecicity was uncertain were listed under “Other specimens exam-
ined”. Nomenclature follows that of Landry (1995), except for the use of the term phallus
(Kristensen 2003). e 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.
Results
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 nal
dataset for species with no or one barcode sequence. Fifteen further non-compliant
barcode sequences were discarded from the nal dataset (see Suppl. material 2: Tab.
S1). We recovered the COI barcode for all but one (H. marijoweissae sp. nov.) spe-
cies described in this paper and for seven of the sixteen previously described species.
Twenty-ve Hoploscopa sequences, as well as 13 sequences from the hypothesised sister-
group 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. e oldest sample from which we
could successfully amplify COI sequence data were collected in 1955 (63 years old).
éo Léger et al. / ZooKeys 907: 1–99 (2020)
8
e 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 reect allopatric diver-
gence. 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 dierentiation is also observed among Philippines
islands: specimens of H. danaoensis sp. nov. from Danao, Negros, and Mindanao rep-
resent three dierent 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.). e two H. sumatrensis MOTUs show a divergence of 4.1–6%. Sec-
ond MOTU, represented by samples MTD8258 and BC_MTD_LEP01422, include
only females and need examination of male specimens.
Systematics
Hoploscopa Meyrick, 1886
Haploscopa Hampson, 1897: 223.
Syncrotaula Meyrick, 1933: 378. Type species Eudorina aurantiacalis Snellen, 1895, by
subsequent designation (for Eudorina Snellen, 1895) by Joannis, 1930.
Type species. Hoploscopa astrapias Meyrick, 1886, by monotypy.
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
Twenty-six new species of Hoploscopa 9
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. e forewings of Perimeceta are slightly larger, display a yellow to brown
ground colour, with one basal elliptic and one postmedian lunule-shaped white spot.
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 geni-
talia 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, rst 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.
orax (Figs 1–45). 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. e pattern elements can
be described as follows (Fig. 3):
cf costal eld stretched along costa up to postmedian area;
bs basal streak small, weakly or not marked;
bp basal patch often quadrangular, ill-dened;
pds and dds proximal and distal discoidal stigmata along costal eld;
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 bare-
ly marked, mostly visible close to costa;
pma postmedian area often suused 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 eld more or less suused with brown, red or yellow.
éo Léger et al. / ZooKeys 907: 1–99 (2020)
10
Figures 1–3. Habitus, venation and forewing pattern in Hoploscopa. 1 Habitus of Hoploscopa mathe-
ae sp. nov., holotype, ♀, Malaysia, Sabah, Kundasang, Kinabalu Mt. Lodge veranda, 6°0'42.15"N,
116°32'3.63"E, 1570 m (T. Léger & R. Mally) (genitalia on slide TL315 ♀) 2 wing venation of Hoplosco-
pa species: H. jubata sp. nov., paratype, ♂, Papua New Guinea, Morobe Prov., nr Bulolo, Mt Susu Nat.
Res., Araucaria For., 975 m, 27–28 Aug 1983 (S. Miller) (wing preparation TL706) 3 nomenclature used
for description of forewing pattern of Hoploscopa spp. Abbreviations: bp (basal patch), bs (basal streak), cf
(costal eld), dds (distal discoidal stigma), mcp (median cubital patch), mds (median discoidal stigma),
mdp (median dorsal patch), pds (proximal discoidal stigma), pma (postmedian area), pml (postmedian
line), pmp (postmedian patch), sbf (subterminal eld), sbl (subterminal line).
Twenty-six new species of Hoploscopa 11
e 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 an-
droconial 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 grey-
ish 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.
Tympanal organs (Figs 83–84).
tb Tympanic bridge deeply divided in the centre;
td Tympanic drum (Fig. 50) ovoid, antero-ventrally semi-closed, posteriorly
not reaching transverse ridge;
tdp Tympanic depressions broad, opened ventrally;
tm Tympanum subtriangular;
tr Transverse ridge (“”) not sclerotised.
Abdomen. Pale brown to brown. In males, sternum A8 more or less broadly in-
dented, in some species with short, rounded lateral projections (Figs 85, 86).
Male genitalia (Figs 46–82).
c cornutus on phallus elongated, at, spatula-shaped, except otherwise men-
tioned;
de ductus ejaculatorius;
g Gnathos arms (Fig. 54) originating from the dorsal part of tegumen, form-
ing ring, with or without dorso-median extension;
jx Juxta basally wide with anterior margin rounded to quadrangular, narrow-
ing at basal 1/4;
s saccus triangular to quadrangular, pointing dorsally;
ta Tegumen arms anteriorly sclerotised, with edge marked, posteriorly mem-
branous; dorsally fused into a bridge of various width, articulated or fused
to uncus base;
uc Uncus well-developed, densely haired apically;
v Valva elongated, hairy, sclerotised, with strongly sclerotised dorsal edge;
vc Vinculum U-shaped in posterior view, dorso-laterally on each side with an-
terior projection bearing brush-shaped coremata.
Female genitalia (Figs 87–122).
aa Anterior apophyses (Fig. 88) bluntly angled at posterior 1/3, bent ventrad;
at Antrum sclerotised or membranous;
éo Léger et al. / ZooKeys 907: 1–99 (2020)
12
cb Corpus bursae globular or ovoid, its membrane with a reticulate
structure;
cs Broad sclerotisation of the corpus bursae;
db Ductus bursae membranous with longitudinal wrinkles, variable in
length and shape;
pa Posterior apophyses slender, straight, directed posterad except other-
wise mentioned;
pp Papillae anales thin in lateral view, dorsally and ventrally connected,
setose;
segm. VIII Segment VIII faintly sclerotised, ventrally membranous; setae scat-
tered across segment, more densely concentrated on posterior margin;
t thorn on corpus bursae large, sclerotised, inwardly projecting, its base
slighly extending outwards into a small sclerotised protuberance.
Distribution. Hoploscopa is found from the Malay Peninsula and Sumatra in the
West to the Samoa Islands in the East, as well as from the North of ailand to Va-
nuatu and Fiji in the South. It is absent from Queensland (Australia) and New Cal-
edonia. It is predominantly found in tropical mountain forests, with only few species
encountered in the lowlands.
Biology. Host plant data is available for ve Hoploscopa species, all feeding on
ferns. e larvae of H. gombongi sp. nov., H. obliqua and H. tonsepi sp. nov. from
Papua New Guinea are reported from Diplazium esculentum (Retzius in Retzius &
König, 1791) Swartz, 1803 (Athyriaceae) (Miller et al. 2015, Mally et al. 2017). An-
other undescribed Hoploscopa species (sample USNM_ENT_00739239) from Papua
New Guinea is reported from Sphaerostephanos unitus (Linnaeus, 1759) Holttum,
1794 (elypteridaceae) (ibid). Lastly, one undescribed Hoploscopa species from Bor-
neo is reported from the fern Dicranopteris linearis (Burman, 1768) Underwood, 1907
(Gleicheniaceae) (Mally et al. 2017).
Hoploscopa albipuncta Léger & Nuss, sp. nov.
http://zoobank.org/26FE78DF-5D7E-4034-A730-FA5C7177466E
Figs 4, 46, 87
Material examined. Holotype: ♂, with labels: “MALAYSIA: Sabah, Kinabalu Park
H[ead]Q[uarter], | Timpohon Gate, 700m from Liwagu | Trail starting point, near
Liwagu River, | 6°1'40"N, 116°32'59"E, 1760m, UV light, | 18.vi.2015, leg. T. Léger
& R. Mally”; “TL335 | ♂”. Deposited in BORN.
Paratypes: 13 ♂, 1 ♀. MALAYSIA: 8 ♂ (1 with genitalia on slide TL336 ♂),
same data as holotype; 3 ♂ (1 with genitalia on slide TL536 ♂), Sabah, Kinabalu
National Park, Timpohon Gate, 300 m from Ligawu trail starting point, 6°1'41"N,
116°32'54"E, 1820 m, UV light, 18.vi.2015, leg. T. Léger & R. Mally (MTD); 1 ♂,
Twenty-six new species of Hoploscopa 13
Sabah, Mesilau, Kopogon, 24.ii.2006, light trap, leg. W. & M. Mey; 1 ♂ (genitalia on
slide TL635 ♂), Sabah, Mesilau River, 24–25.ii.2006, light trap, leg. W. & M. Mey; 1
♀ (DNA voucher MTD7434, genitalia on slide TL609 ♀), Sabah, Mesilau, 2000 m,
14–17.xi.2006, light trap, leg. W. Mey & K. Ebert (MFNB).
Other specimens examined. 1 ♂. MALAYSIA: 1 ♂ (DNA voucher ITBC09,
genitalia on slide TL309 ♂), same data as holotype.
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.
Similar species. No similar species known.
Description. Head. Antennae dorsally with brown scales. Proboscis pale yellow.
Maxillary palpi brown, basally light brown. Labial palpi brown, ventrally pale yellow.
orax (Fig. 4). Collar white. Forewing length: 9–10 mm (♂ & ♀); forewing
ground colour brown; rounded snow white basal cubital spot; median discoidal stigma
rhomboid to ellipsoid, dark red, edged with pale yellow; postmedian spot trapezoid,
dark red, proximally pale yellow; subterminal line snow white, parallel to termen,
slightly angled at M1, broader at costa; fringes brown, with white dots. Hindwing
dirty pale yellow. Forelegs bronze. Midlegs brown, with tibia and tarsi segments distally
white. Hindlegs brown, with tibia and tarsi segments distally white.
Abdomen. Male sternum A8 posterior margin straight.
Male genitalia (N = 5) (Fig. 46). Uncus long and slender, gently narrow-
ing on basal half, apex tongue-shaped. Gnathos projection broad, triangular, with
rounded apex. Valva ventral margin straight, dorsal margin strongly protruded
dorsad, apex rounded. Juxta with base rounded, medially narrowed, apex split
into two conspicuous tips. Saccus small, pointing dorsad. Phallus with sclerotised,
anvil-shaped cornutus.
Female genitalia (N = 1) (Fig. 87). Anterior apophyses with dorsal bump at pos-
terior 1/3. Antrum forming a membranous rounded pouch. Ductus bursae of medium
length, slender, straight. Corpus bursae small, globular, reticulate, with barely marked
sclerotisation at thorn base. orn curved, with small dents pointing toward thorn
apex on its inner side, glabrous on the outer side.
Distribution. Known from the slopes of the Mount Kinabalu (4,095 m) on Bor-
neo, 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. e species name albipuncta refers to the Latin albus, white, and punc-
tus, forming a point.
Remarks. Two female specimens from the Philippines with similar habitus and
genitalia but a COI barcode divergence of 4.5–4.9% are recovered as dierent MOTU
in the species delimitation analyses. Unfortunately, no male was available for this study,
thus we refrained from describing a new species based on these two specimens.
éo Léger et al. / ZooKeys 907: 1–99 (2020)
14
Figures 4–18. Forewing of Hoploscopa species. 4 Hoploscopa albipuncta sp. nov., holotype, ♂, Malaysia,
Sabah, Kinabalu Park HQ, Timpohon Gate, 700 m from Liwagu Trail starting point, near Liwagu
River, 6°1'40"N, 116°32'59"E, 1760 m, 18.vi.2015 (T. Léger & R. Mally) (genitalia on slide TL335
♂) 5 Hoploscopa sepanggi sp. nov., holotype, ♂, Malaysia, Sabah, Mesilau Nature Resort, 6°2'43.71"N,
116°35'48.03'E, 1925 m, at lamps, 30.v.2015 (T. Léger & R. Mally) 6 Hoploscopa cynodonta sp.
nov., holotype, ♂, Malaysia, Sabah, Kinabalu Park HQ, junction Kiau View- and Pandanus Trail,
6°0'32.84"N, 116°32'14.94"E, 1690m, UV light, 07.vi.2015 (T. Léger & R. Mally) (genitalia on slide
TL327 ♂) 7 Hoploscopa parvimacula sp. nov., paratype, ♀, Sabah, Mesilau Nature Resort, 6°2'43.71"N,
116°35'48.03"E, 1925 m, at lamps, 30.v.2015 (T. Léger & R. Mally) (genitalia on slide TL312 ♀.
Twenty-six new species of Hoploscopa 15
Hoploscopa matheae Léger & Nuss, sp. nov.
http://zoobank.org/D792135D-FFA9-49ED-8B76-C9802548E8E5
Figs 1, 47, 88
Material examined. Holotype: ♀, with labels: “MALAYSIA: Sabah, Kundasang,
| Kinabalu Mt. Lodge veranda, 6°0'42.15"N, 116°32'3.63"E, 1570 m, at day,
15.vi.2015, | leg. T. Léger & R. Mally”; “DNA barcoding | BC MTD Lep 3004”;
“ITBC | 15”; “TL315 | ♀”. Deposited in BORN.
Paratypes: 1 ♂, 2 ♀. BRUNEI: 1 ♂ (NHMUK010923334, DNA voucher
MTD8245, genitalia on slide TL730 ♂), 1 ♀ (NHMUK010923427), Bukit Retak,
LP 238, GR 873804, 1365m, moss forest, 1–4.v.1989, leg. M. G. Allen & K. R. Tuck
(NHMUK). MALAYSIA: 1 ♀ (DNA voucher MTD7426, genitalia on slide TL599
♀), Sabah, Tawau Hills, Gelas River, 3.iii.2006, leg. W. + M. Mey (MFNB).
Other specimens examined. 2 ♀. InDONESIA: 1 ♀ (NHMUK010923426),
Java, Singolangoe, Tengger, 5000 feet, 05.1934 (F. P. A. Kalis) [abdomen missing]
(NHMUK). MALAYSIA: 1 ♀ (NHMUK010923425, DNA voucher MTD8244
& genitalia on slide TL729 ♀), Cameron Highlands, Gunung Brinchang, 15–23.
viii.1986 (G. S. Robinson) (NHMUK).
Diagnosis. Hoploscopa matheae sp. nov. is unique in the genus by its broad pale
yellow forewing fascia crossed by postmedian brown line as well as the basal and distal
discoidal spots circled with pale yellow.
Similar species. No similar species known.
Figures 4–18. Continued. 8Hoploscopa kinabaluensis sp. nov., paratype, ♀, Sabah, Kinabalu Park HQ,
Timpohon Gate, 700 m from Liwagu trail starting point, near Liwagu River, 6°1'40"N, 116°32'59"E,
1760 m, UV light, 18.vi.2015 (T. Léger & R. Mally) (genitalia on slide TL326 ♀) 9Hoploscopa
luteomacula Nuss, paratype, ♀, Indonesia, Sumatra, Barat, N-Padangpanjang, Mt Singgalang, 2100 m,
10–11.ii.1996 (A. Kallies) (genitalia on slide GU prep. Nuss 743) 10 Hoploscopa obliqua Rotschild,
abdomen missing, Papua New Guinea, Madang Province, Wanang village, 05°15'S, 145°17'E, reared
from Diplazium esculentum, 12.vi.2007 11 Hoploscopa niveofascia sp. nov., holotype, ♂, Papua New
Guinea, Morobe Prov., nr. Bulolo, Mt. Susu Nat. Res., 975 m, 27–28.viii. 1983, Araucaria For. (S. Miller)
(genitalia on slide TL442 ♂) 12 Hoploscopa gombongi sp. nov., paratype, ♂, Papua New Guinea, Yawan
village, 06°10'S, 146°5'E, NG Binatang Res. Ctr. (B Gewa, J. Kua, S Sau, A. Kinibel) 13 Hoploscopa
tonsepi sp. nov., holotype, ♂, Papua New Guinea, Yawan village, 06°10'S, 146°5'E, NG Binatang
Res. Ctr. (J. Valeba, J. Auga, M. Dilu, F. Philip, R. Lilip) (genitalia on slide TL655 ♂) 14Hoploscopa
marijoweissae sp. nov., ♂, Papua New Guinea: Morobe Province, Mount Kaindi, 2350m, 11.xii.1976
(G. F. Hevel & R. E. Dietz) (genitalia on slide TL437 ♂) 15 Hoploscopa titika sp. nov., holotype, ♂,
Sumatra-Holzweg, 25 km SSW-Pematangsiantar, Straße nach Prapat, 13.ii.1996 (A. Kallies) (genitalia
on slide TL505 ♂) 16 Hoploscopa pangrangoensis sp. nov., paratype, ♀, Indonesia, Java, Mt. Pangrange,
SE Bogor, 6.30S 107.10E, 1625 m, primary forest, 16–20.ii.1996 (Siniaev & Afonin) (genitalia on
slide TL627 ♀) 17Hoploscopa isarogensis sp. nov., paratype, ♀, Philippines, South Luzon, Mt Isarog,
13°40'N, 123°20'E, 530 m, submontane forest, at light, 22.iii.2000 (M. Nuss) (genitalia on slide TL523
♀) 18 Hoploscopa ypsilon sp. nov., paratype, ♀, Philippines, Luzon, Mountain Province, Chatol, 2100
m, 16–18.xi.1997 (Mey, Ebert & Nuss).
éo Léger et al. / ZooKeys 907: 1–99 (2020)
16
Description. Head. Antennae dorsally with pale yellow scales. Proboscis dark
brown. Maxillary palpi dark brown, basally and inwardly pale yellow. Labial palpi dark
brown, ventro-basally pale yellow.
orax (Fig. 1). orax dark brown, with two pale yellow stripes laterally and
one dorsally. Collar pale yellow. Forewing length: 10 mm (♂ & ♀); forewing ground
colour dark brown, with darker scales at its base; basally with a roughly dened Z-
line pale yellow; basal and distal discoidal stigma of a darker brown, circled with
pale yellow, median discoidal stigma therebetween dark red in some specimens; broad
median triangular pale yellow patch near dorsum, connected distally by a line to post-
median triangular pale yellow patch; subterminal pale yellow line following median
line shape, originating at distal 1/4 of dorsum, inwardly incurved between A1+2 and
Cu2, costally diverging toward apex; subterminal eld broadly marked with scales of
a lighter brown; fringes dark brown, with pale yellow dots. Hindwing pale brown.
Forelegs dark brown. Midlegs brown; femur and tibia distally pale yellow; tarsi pale
yellow. Hindlegs dark brown, femur-tibia articulation pale yellow; tibia distally pale
yellow; tarsi bronze.
Abdomen. Male sternum A8 posterior margin straight.
Male genitalia. (N = 1) (Fig. 47). Uncus broad, narrowing on distal half, with
apex truncate. Gnathos without posterior projection. Valva ventral margin slightly
concave, bent dorsad on apical 1/5; dorsal margin conspicuously convex; apex blunt.
Juxta with base slightly quadrangular, medially narrow, apex broadly incurved. Saccus
not pronounced.
Female genitalia (N = 2) (Fig. 88). Anterior apophyses with dorsal tip pointed
dorsad at posterior 1/3. Antrum sclerotisation twice as long as broad. Ductus bur-
sae of medium length, roughly straight. Corpus bursae pear-shaped, reticulate on
posterior half, sclerotised between thorn and corpus opening. orn straight, with
small dents pointing toward thorn apex, basally with small outwardly projected
extension.
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 K2P-
distance 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).
Etymology. e species is named after Mathéa Léger, sister of the rst author.
Remarks. One specimen from the NHMUK collected on Mount Tengger (In-
donesia, Java) in 1934 shares a similar wing pattern and thus potentially represents
a conspecic specimen. Unfortunately, the abdomen of the specimen was missing.
Specimen MTD8244 from the Malay Peninsula shows minor dierences in wing pat-
tern with those from Borneo: costal eld, median discoidal stigma and subterminal
eld are dark red, and base of thorn on corpus bursae is thinner.
Twenty-six new species of Hoploscopa 17
Hoploscopa sepanggi Léger & Nuss, sp. nov.
http://zoobank.org/81756486-A524-4D2A-912E-F824C234F835
Figs 5, 48, 89
Material examined. Holotype: ♂, with labels: “Malaysia, Sabah, Mesilau Nature Re-
sort | 6°2'43.71"N, 116°35'48.03"E, 1925 m, at | lamps, 30.05.2015, leg. T. Léger &
R. Mally”. Deposited in BORN.
Paratypes: 6 ♂, 3 ♀. MALAYSIA: 1 ♂ (genitalia on slide TL260 ♂) with same data
as holotype except 02.vi.2015; 3 ♂ (genitalia on slide TL641 ♂ & TL338 ♂), 1 ♀ (gen-
italia on slide TL547 ♀), Sabah, Mesilau, logging site 400 m before entrance to Mesilau
Nature Resort, 6°2'22"N, 116°35'54"E, 1930 m, UV light, 02.vi.2015, leg. T. Léger
& R. Mally; 1 ♀ (genitalia TL337 on slide TL337 ♀), same data except 01.vi.2015;
1 ♂ (DNA voucher MTD7894, genitalia on slide TL671, wing preparation on slide
TL237), Sabah, Kinabalu National Park, Timpohon Gate, 300 m from Ligawu trail
starting point, 6°1'41"N, 116°32'54"E, 1820 m, UV light, 18.vi.2015, leg. T. Léger &
R. Mally; 1 ♂ (DNA voucher MTD LEP123, genitalia on slide TL365 ♂), 1 ♀, Sabah,
Mesilau, 2000 m, 14–17.xi.2006, light trap, leg. W. Mey & K. Ebert (MFNB).
Other specimens examined. 1 ♀. MALAYSIA: 1 ♀ (DNA voucher ITBC02,
genitalia on slide TL302 ♀), same data as holotype.
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. e postmedian cubital and subterminal
patch are missing or reduced to traces in H. sepanggi sp. nov., while they are well-
marked, pale yellow in H. luteomacula. e 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.
orax (Fig. 5). orax 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 eld 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 in-
terspaced sulphur yellow dashes, with triangular sulphur costal spot; subterminal eld
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.
Abdomen. Male sternum A8 posterior margin bilobed, with short, rounded lat-
eral projections.
éo Léger et al. / ZooKeys 907: 1–99 (2020)
18
Male genitalia (N = 5) (Fig. 48). Uncus long and slender, gently tapering toward
tongue-shaped apex. Gnathos projection slender, ca. half the uncus length. Valva
ventral margin gently bent dorsad on distal 1/3, dorsal margin conspicuously convex,
apex slightly pointed. Juxta with base roughly quadrangular, medially narrowed, apex
narrow, faintly notched. Saccus broad, triangular, pointing dorsad.
Female genitalia (N = 3) (Fig. 89). Anterior apophyses with small dorsal bump at
1/3. Antrum sclerotisation twice as long as broad. Ductus bursae of medium length,
slender, elbowed at 1/4 and 3/4. Corpus bursae ovoid, reticulated, diuse sclerotisa-
tion between thorn and corpus opening, medially with faintly sclerotised band. orn
straight, small, with tiny dents pointing toward thorn base, basally with small out-
wardly projected extension.
Distribution. Known from the slopes of the Mount Kinabalu on Borneo, at alti-
tudes between 1,800 m and 2,000 m.
Etymology. is species is described in honour of the late Malaysian mountain
guide Robbie Sepanggi who died on Mount Kinabalu during the 2015 Sabah earth-
quake while trying to save hikers.
Hoploscopa cynodonta Léger & Nuss, sp. nov.
http://zoobank.org/55B144AD-1AF7-4023-BC82-E5E2750AFE88
Figs 6, 49, 90
Material examined. Holotype: ♂, with labels: “Malaysia, Sabah, Kinabalu Park
H[ead]Q[quarters], | junction Kiau View- and Pandanus | Trail, 6°0'32.84"N,
116°32'14.94"E, | 1690 m, UV light, 07.vi.2015, | leg. T. Léger & R. Mally”; “DNA
voucher | Lepidoptera | MTD 2015 | [vertically written:] no. 3066”; “TL327 | ♂”.
Deposited in BORN.
Paratypes: 4 ♂, 2 ♀. BRUNEI: 2 ♂ (1 with NHMUK010923398, DNA
voucher MTD8233 & genitalia on slide TL746; 1 with NHMUK010923401), 1
♀ (NHMUK010923400), Ulu Temburong, LP 298, GR 838892, 300 m, 26–30-
iv-1989, leg. M. Allen & K. Tuck (NHMUK). MALAYSIA: 1 ♂ (genitalia on
slide TL531 ♂), Sabah, Kinabalu National Park, Timpohon Gate, 700 m from
Liwagu Trail starting point, near Liwagu River, 6°1'40"N, 116°32'59"E, 1700 m,
18.vi.2015, leg. Léger & R. Mally (MTD); 1 ♀ (DNA voucher MTD7425 & geni-
talia on slide TL625 ♀), Sabah, Tawau Hills National Park, Headquarter, waterfall,
23–27.xi.2006, leg. W. Mey & K. Ebert (MFNB). 1 ♂ (NHMUK010923352),
Sabah, Gunung Monkobo, 5.48N, 116.56E, dipterocarp forest, 945m, 14–23.
viii.1987, leg. K. Tuck (NHMUK).
Diagnosis. is relatively small-sized species (forewing length 8–10 mm) shows
brown forewings with red markings edged with pale yellow. e 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. e postmedian patch consists of a mix of
red and brown scales, at costa forming triangular pale yellow patch. Male genitalia are
Twenty-six new species of Hoploscopa 19
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. Medi-
an 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 geni-
talia, 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 diuse.
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.
orax (Fig. 6). Collar white. Forewing length: 8–10 mm (♂), 8 mm (♀); fore-
wing ground colour brown; basal longitudinal red streak basally surrounded with pale
yellow U-shaped spot; costal eld red; median discoidal stigma oval, red with pale yel-
low edges; median cubital patch forming longitudinal pale yellow rectangle; postme-
dian patch red, with basal edge pale yellow, distally abutted with pale yellow triangular
costal patch; postmedian line costally marked; postmedian suusion of pale yellow
and pale brown scales, distally edged by pale yellow subterminal line; subterminal eld
marked with red; fringe chequered pale yellow and brown. Hindwing pale brown.
Forelegs and midlegs brown. Hindlegs with femur brown; tibia pale yellow, distally
brown; tarsi bronze speckled with pale yellow.
Abdomen. Male sternum A8 posterior margin forming two conspicuous triangular
tips, with short, rounded lateral projections.
Male genitalia (N = 2) (Fig. 49). Uncus medially slightly widened, narrowing
toward apex, apex duck beak-shaped, medio-ventrally with a bump and two ridges.
Gnathos projection roughly triangular, ca. 1/4 of uncus length, with rounded apex.
Valva ventral margin with basal half sclerotised, at 1/4 conspicuously incurved into
a semi-circle, protruded medio-ventrally into a tooth-like tip pointing ventrad; valva
dorsal margin convex, apex truncate. Juxta with base rounded, medially narrowed,
apical half laterally protruded, apex broad, not sclerotised. Saccus quadrangular, con-
spicuously produced anterad.
Female genitalia (N = 1) (Fig. 90). Posterior apophyses bent ventrad. Anterior
apophyses with small dorsal bump at 1/3. Antrum sclerotisation twice as long as broad,
barely sclerotised. Ductus bursae of medium length, broad, with a narrow loop before
corpus bursae. Corpus bursae globular, reticulated, with crest of sclerotised acanthae
éo Léger et al. / ZooKeys 907: 1–99 (2020)
20
between thorn and corpus opening. orn straight, slightly incurved at apex, with
small dents pointing toward thorn base.
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.
DNA barcoding. Specimens MTD LEP3066 from Mount Kinabalu and
MTD7425 from Tawau Hills dier by 0.4%. e nearest neighbour is H. isarogensis
sp. nov. from the Philippines (K2P-dist = 2.3–3%).
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.
Hoploscopa parvimacula Léger & Nuss, sp. nov.
http://zoobank.org/4E445DD3-8C71-40DE-9EDB-870935206EDE
Figs 7, 50, 91
Material examined. Holotype: ♂, with labels: “MALAYSIA, Sabah, Mesilau, logging
| site 400m before entrance to Mesilau | Nature Resort, 6°2'21.97"N, 116°35'54.12"E,
1930m, UV light, | 02.vi.2015, leg. T. Léger & R. Mally”; “DNA voucher | Lepidop-
tera | MTD2016 | [vertically written:] n°. 3196 ”; “TL510 | ♂”. Deposited in BORN.
Paratypes: 7 ♂, 2 ♀. MALAYSIA: 1 ♂ (DNA voucher MTD7888 & genitalia
on slide TL552 ♂) with same data as holotype, 2 ♂ (DNA vouchers MTD7884 &
MTD7885, genitalia on slides TL673 ♂ & TL261 ♂), 1 ♀ (DNA voucher MTD7883
& genitalia on slide TL551 ♀) with same data as holotype except 02.vi.2015; 1 ♂
(DNA voucher MTD7886, genitalia on slide TL537 ♂), Sabah, Mesilau Nature Re-
sort, 6°2'44"N, 116°35'48"E, 1925 m, at lamps, 30.v.2015, leg. T. Léger & R. Mally;
1 ♀ (DNA voucher ITBC12, DNA barcoding voucher BC MTD LEP3003, genitalia
on slide TL312 ♀), same data except 01.vi.2015; 2 ♂ (DNA vouchers MTD7895
& MTD7896, genitalia on slides TL550 ♂ & TL549 ♂), Sabah, Kinabalu Na-
tional Park, Timpohon Gate, 300 m from Liwagu trail starting point, 6°1'41"N,
116°32'54"E, 1820 m, UV light, 18.vi.2015, leg. T. Léger & R. Mally; 1 ♂ (DNA
voucher MTD7890, genitalia on slide TL672 ♂), Sabah, Kundasang, Kinabalu Moun-
tain Lodge veranda, 6°0'42"N, 116°32'4"E, 1570 m, at light, 18.vi.2015, leg. T. Léger
& R. Mally (MFNB).
Diagnosis. Hoploscopa parvimacula sp. nov. displays brown forewings with re-
duced pale yellow markings. e median discoidal stigma is crescent-shaped, and
the median cubital patch forms a streak not connected to the median discoidal
stigma. e 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.
Twenty-six new species of Hoploscopa 21
Similar species. Hoploscopa kinabaluensis sp. nov., H. pangrangoensis sp. nov. e
forewings of H. kinabaluensis sp. nov. display more contrasting postmedian patch and
subterminal line, and a subterminal eld suused with pale yellow. In H. pangrangoen-
sis 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. e nger-like
gnathos projection of H. pangrangoensis sp. nov. is narrower than that of H. parvimacu-
la 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 mem-
branous with longitudinal sclerotised lines, corpus bursae is larger and thorn is smaller.
Description. Head. Antennae dorsally striped with brown and bronze scales. Pro-
boscis pale yellow, speckled with pale brown. Maxillary palpi dark brown, base and
inner side pale yellow. Labial palpi dark brown, ventro-basally pale yellow.
orax (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 yel-
low, speckled with brown; tarsi bronze. Hindlegs with femur brown; tibia pale yellow,
speckled with brown; tarsi pale yellow to pale brown.
Abdomen. Male sternum A8 posterior margin bilobed, with short, rounded lat-
eral projections.
Male genitalia (N = 8) (Fig. 50). Uncus medially broadened, narrowing on distal
half, apex slightly obtuse. Gnathos projection ca. 2/3 of uncus length, spatula-shaped.
Valva ventral margin straight, gently bent dorsad on distal 1/4, dorsal margin convex,
apex rounded or slightly pointed. Juxta slender, base roughly quadrangular, apex trun-
cate. Saccus small, pointing dorsad.
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 diuse sclerotisation between thorn
and corpus opening, medially with faintly marked sclerotised band. orn 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.
éo Léger et al. / ZooKeys 907: 1–99 (2020)
22
Remarks. e specimen with DNA voucher MTD8229 belongs to a series of fe-
males 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.
Hoploscopa kinabaluensis Léger & Nuss, sp. nov.
http://zoobank.org/F75F92C1-7C04-4BE9-A602-0FE822BF66D2
Figs 8, 51, 92
Type material. Holotype: ♀, with labels: “MALAYSIA, Sabah, Kundasang | Kinabalu
Mt. Lodge veranda, | 6°0'42.15"N, 116°32'3.63"E, | 1570m, at light, 16.vi.2015, |
leg. T. Léger & R. Mally”; “DNA voucher | Lepidoptera | MTD 2015 | [vertically writ-
ten:] no. 3064”; “TL325 ♀”. Deposited in BORN.
Paratypes: 3 ♂, 4 ♀. MALAYSIA: 1 ♂ (DNA voucher MTD7889, genitalia on
slide TL667 ♂), same data as holotype; 1 ♂ (DNA voucher MTD7893 & genitalia
on slide TL342 ♂), 1 ♀ (DNA voucher MTD LEP3065 & genitalia on slides TL326
♀), Sabah, Kinabalu National Park, Timpohon Gate, 700 m from Liwagu trail start-
ing point, near Liwagu River, 6°1'40"N, 116°32'59"E, 1760 m, UV light, 18.vi.2015,
leg. T. Léger & R. Mally; 1 ♂ (DNA voucher MTD7897, genitalia on slide TL341
♂), Sabah, Kinabalu National Park, Timpohon Gate, 300 m from Liwagu trail starting
point, 6°1'41"N, 116°32'54"E, 1820 m, UV light, 18.vi.2015, leg. T. Léger & R. Mal-
ly; 1 ♀ (DNA voucher MTD7882, TL666 ♀), Sabah, Kinabalu National Park Head-
quarters, junction Kiau View- and Pandanus Trail, 6°0'32.84"N, 116°32'15"E, 1690
m, UV light, 07.vi.2015, leg. T. Léger & R. Mally; 1 ♀ (DNA voucher MTD7887,
genitalia on slide TL662 ♀), Sabah, Kundasang, road 200 m before Kinabalu Mt.
Lodge, 6°0'37"N, 116°32'4"E, 1535m, UV light, 03.vi.2015, leg. T. Léger & R. Mally
(MTD).
Other specimens examined. 1 ♂, 2 ♀. MALAYSIA: 1 ♂ (DNA voucher
MTD7892, genitalia on slide TL546 ♂), 1 ♀ (DNA voucher MTD7891 & genitalia
on slide TL674 ♀), Sabah, Kinabalu National Park, Timpohon Gate, 700 m from
Liwagu trail starting point, near Liwagu River, 6°1'40"N, 116°32'59"E, 1760 m, UV
light, 18.vi.2015 (T. Léger & R. Mally); 1 ♀ (NHMUK 010923435, DNA voucher
MTD8232 & genitalia on slide TL745 ♀), Sabah, Gunung Monkobo, dipterocarp
forest, 975m, 5.48N, 116.56E (7–13.viii.1987) (NHMUK).
Diagnosis. e 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. ese species share similar forewing markings with
Twenty-six new species of Hoploscopa 23
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.
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.
orax (Fig. 8). orax dorsally pale yellow, laterally brown. Collar pale yellow.
Forewing length: 9.0–10.5 mm (♂), 10–11 mm (♀); forewing ground colour brown;
basal dark brown dash, distally pale yellow; median discoidal stigma and median cu-
bital patch forming together a pale yellow-coloured Y, disrupted with brown at vein,
edged basally and distally with dark brown; postmedian patch roughly triangular,
pale yellow, invaded with brown, edged with dark brown; subterminal line thick,
pale yellow; fringe chequered pale yellow and brown. Hindwing pale brown. Forelegs
brown. Midlegs with femur brown; tibia brown, inwardly pale yellow; tarsi bronze.
Hindlegs with femur brown; tibia pale yellow, speckled with brown; tarsi bronze,
speckled with brown.
Abdomen. Male sternum A8 posterior margin broadly indented, with short,
rounded lateral projections.
Male genitalia (N = 4) (Fig. 51). Uncus broad, gently narrowing on distal half,
apex obtuse. Gnathos projection ca. 2/3 of uncus length, narrow at base, widening
toward apex, apex spatula-shaped. Valva ventral margin straight, gently bent dorsad on
distal 1/3, dorsal margin convex, apex rounded or slightly pointed. Juxta slender, base
roughly quadrangular, apex slightly incurved. Saccus small, pointing dorsad. Phallus
with elongated cornutus, at posterior end with narrow bump pointing dorsad.
Female genitalia (N = 5) (Fig. 92). Anterior apophyses with tip or bump at poste-
rior 1/3. Antrum sclerotisation ca. twice as long as broad. Ductus bursae long, slender,
with one loop, curved before corpus bursae. Corpus bursae conspicuously pear-shaped,
reticulated, with sclerotisation between thorn and corpus opening and faintly marked
sclerotised band medially. orn long, slightly curved, with small dents pointing to-
ward thorn apex.
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. kina-
baluensis sp. nov. ese two lineages are found in sympatry at Mount Kinabalu.
Etymology. e species name kinabaluensis refers to Mount Kinabalu on Borneo,
where the species occurs.
Remarks. e two barcode lineages show no dierences in wing pattern and
female genitalia. Only minor dierences 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.
éo Léger et al. / ZooKeys 907: 1–99 (2020)
24
Hoploscopa luteomacula 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”; “Coll. M. Nuss | Geschenk 2000 | Museum für Tier- | kunde
Dresden”. Deposited in MTD.
Paratype: 2 ♀. INDONESIA: 1 ♀ (genitalia on slide GU 743, DNA barcode BC
MTD 01419), same data as holotype; 1 ♀ (genitalia in capsule under specimen), Su-
matra, Holzweg 2, 25km SSW-Pematangsiantar, Strasse nach Prapat [road to Prapat],
25.x.1989, leg. E. W. Diehl (MTD).
Other material examined. 1 ♂, 1 ♀. BRUNEI: ♀ (NHMUK010923403, DNA
voucher MTD8234 & genitalia on slide TL747 ♀), Ulu Temburong, LP 298, GR
838892, 300m, 26–30.iv.1989 (M. G. Allen & K. R. Tuck). MALAYSIA: 1 ♂ (DNA
voucher MTD LEP3195 & genitalia on slide TL525 ♂), Sabah, Kundasang, road
200m before Kinabalu Mt. Lodge, 6°0'37.38"N, 116°32'0.35"E”, 1535m, UV light,
03.vi.2015, leg. T. Léger & R. Mally (MFNB).
Diagnosis. Hoploscopa luteomacula displays broad pale yellow patches in the fore-
wing. 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.
Similar species. Hoploscopa sepanggi (q.v.).
Description. Head. Antennae dorsally pale yellow. Proboscis pale yellow. Maxil-
lary palpi brown, base and inner side pale yellow. Labial palpi brown, ventral base and
inner side pale yellow.
orax (Fig. 9). orax brown, dorsally pale yellow. Collar pale yellow. Forewing
length: 9–11 mm (♂ & ♀); forewing ground colour brown, with markings pale yel-
low; basal patch large, rhomboid, slightly encroached with brown near dorsum; small
basal discoidal spot; median discoidal stigma trapezoid; costal eld pale yellow, speck-
led with brown; median cubital and dorsal patches rhomboid; postmedian patch quad-
rangular to elliptic, dorsally invaded with brown; postmedian cubital patch rhomboid;
subterminal costal patch triangular; subterminal eld more or less broadly marked
with pale yellow; fringes pale yellow, sometimes with tiny brown dots, apex brown.
Hindwing pale yellow to pale brown. Forelegs femur pale yellow; tibia brown, inner
side pale yellow; tarsi pale yellow speckled with bronze. Midlegs with femur pale yel-
low; tibia and tarsi brown. Hindlegs pale yellow.
Male genitalia (N = 1) (Fig. 52). Uncus large, rectangular, slightly narrowed at
apical 1/4, apex truncate. Gnathos projection tongue-shaped, ca. 1/3 of uncus length.
Valva ventral margin nearly straight, dorsal margin convex, apex blunt. Juxta with
large rounded base, medially conspicuously narrowed, apex duck beak-shaped. Saccus
small, pointing dorsad.
Female genitalia (N = 1) (Fig. 93). Anterior apophyses with dorsal bump at
posterior 1/3. Antrum sclerotisation reduced to sclerotised ring. Ductus bursae short,
Twenty-six new species of Hoploscopa 25
nearly straight. Corpus bursae large, pear-shaped, reticulated on posterior half, ante-
rior half membranous, with faint sclerotisation between thorn and corpus opening.
orn nearly straight, with dents pointing toward thorn apex, basally with small out-
wardly projected extension.
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.
Remarks. Deep barcode divergence between morphologically similar specimens
from Borneo and Sumatra might represent phylogeographic variability or distinct spe-
cies. Additional material from both regions and a broader set of characters is needed to
test these hypotheses.
Hoploscopa obliqua (Rothschild, 1915)
Figs 10, 53, 94
Material examined. Holotype: ♂, with labels: “Holo- | type” [round label, red ringed];
“Utakwa R[iver]., | Dutch N[ew]. Guin[ea]., | 3000 f[ee]t., Jan[uary]. 1913. | A.F.R.
Wollaston.”; “437”; “Eudorina | obliqua | Type Rotsch[ild].” [handwritten]; “♂ | Pyral-
idae | Brit[ish].Mus[eum]. | Slide N°. | 20252”; "NHMUK 010923283” [barcode ap-
pended]. Deposited in NHMUK.
Other specimens examined. Papua New Guinea: 2 specimens [abdomens lost]
(1 with DNA voucher “USNM ENT 00665932”, 1 with DNA voucher “USNM
ENT 00514731”), 1 ♀ (DNA voucher “USNM ENT 00514750”, genitalia on slide
TL656 ♀), Madang Province, Wanang village, 05°15'S, 145°17'E, 1700 m, reared
from Diplazium esculentum, 12.vi.2007, leg. Auga, Molem, Tamtiai, Lilip, Ibalim, Pos-
man, Rimandai, Brus, Novotny, Hrcek (USNM).
Diagnosis. e antennae striped with white and the antemedian oblique white
streak extended distally into white suusion 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.
orax (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 suf-
fusion; postmedian patch reduced to costal white blotch; postmedian line of a darker
brown at costa; barely marked subterminal white line incurved inwards at CuA2, run-
ning up to M2; fringes pale brown. Hindwing pale brown. Fore- and midlegs brown,
tibia and tarsi segments distally white. Hindlegs brown.
éo Léger et al. / ZooKeys 907: 1–99 (2020)
26
Male genitalia (N = 1) (Fig. 53). Uncus medially conspicuously widened, nar-
rowed at apical 1/4, apex spatulate. Gnathos projecting into a broad spatula-shaped
plate with small apical tip on its middle. Valva ventral margin gently bent dorsad on
distal 1/3, dorsal margin convex, apex pointed. Juxta slender, with base rounded, apex
obtuse. Saccus small, pointing dorsad. Phallus with club-shaped cornutus.
Female genitalia (N = 1) (Fig. 94). Anterior apophyses bent ventrad at 1/3. An-
trum membranous. Ductus bursae long, straight. Corpus bursae globular, with one
half reticulated, one half with erect papillae, with sclerotised bump on each side of
thorn. orn straight, broad at base, tapering at mid-length, with small dents pointing
toward thorn apex.
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).
Remarks. Nuss (1998) transferred this species from Eudorina to Hoploscopa.
Hoploscopa niveofascia Léger & Nuss, sp. nov.
http://zoobank.org/4112C427-AF2B-4CAE-BDB3-E94033B8E8C2
Figs 11, 56
Material examined. Holotype: ♂, with labels: “PAPUA NEW GUINEA | Morobe
Prov[ince]., n[ea]r. Bulolo | Mt. Susu Nat[ional]. Res[erve]., 975m | 27–28 Aug.1983,
S. Miller | UV Lite, Araucaria For[est]”; “DNA voucher | Lepidoptera | MTD2016 |
[vertically written:] no. 3162”; “TL442 | ♂”. Deposited in USNM.
Diagnosis. Hoploscopa niveofascia sp. nov. displays snow white postmedian patch,
subterminal line and well-marked marginal spots on the forewing. In male genitalia, the
uncus is broad, with straight lateral margin and a blunt apex. Female genitalia not known.
Similar species. Hoploscopa diusa. Hoploscopa niveofascia sp. nov. shares with H.
diusa the white markings of the forewing. However, postmedian patch of H. diusa
is broadly marked with brown, marginal spots are reduced to small dots, and fringes
exhibit white spots, while fringes are brown in H. niveofascia sp. nov.
Description. Head. Antennae dorsally dark brown. Proboscis white. Maxillary
palpi dark brown, basally pale brown. Labial palpi dark brown, ventro-basally white.
orax (Fig. 11). orax dark brown with mesodorsal transversal white line. Collar
white. Forewing length: 9.5 mm; forewing ground colour dark brown; basal area with-
out marked patches, white scales scattered near dorsum; median discoidal stigma darker
coloured, rectangular, basally and distally thinly edged with white; postmedian patch
diuse, white; postmedian line marked on costal half; subterminal line thick, zigzagging,
disrupted distally at M1; margin with large white spots; fringes bronze. Fore- and midlegs
dark brown, with tibia and tarsi segments distally white. Hindlegs brown to bronze.
Abdomen. Male sternum A8 posterior margin straight.
Male genitalia (N = 1) (Fig. 56). Uncus broad with straight lateral margin, apex
truncate, medially slightly incurved. Gnathos reduced to thin band without poste-
Twenty-six new species of Hoploscopa 27
rior projection. Valva ventral margin bent dorsad on distal half, dorsal margin slightly
convex, apex pointed. Juxta elongated, with base rounded, medially narrowing, apex
blunt. Saccus small, pointing dorsad.
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).
Etymology. From the Latin niveus, snowy, and fascia, band, referring to the snow-
white markings on the forewing.
Hoploscopa gombongi Léger & Nuss, sp. nov.
http://zoobank.org/74AD0BB2-8718-42BA-88A3-0C6236302D18
Figs 12, 54, 96
Material examined. Holotype: ♂, with labels: [exuvia pinned under the specimen]
“Sp. PYRA-106 | YS.2H.3826 | CATY 001 L[ength] 9mm | YC 29975 Leaf M [cir-
cled] Y | Roll [circled] - Tie - Chew - Skel | 10-SEP-2012 [handwritten]”; “Papua New
Guinea | Yawan village | 06°10'S, 146°5'E [1700m] | L[e]g[i]t B Gewa, J. Kua, | S Sau,
A. Kinibel | NG Binatang Res[earch]. C[en]t[e]r.”; “USNM ENT: PNG | Madang
Ecology Project | [barcode] | 00739216” [DNA voucher]; “DNA 2013”; “TL | 653
♂”. Deposited in USNM.
Paratypes: 1 ♂, 2 ♀. PAPUA NEW GUINEA: 1 ♂ (“YC29970”, DNA voucher
“00739200”), 1 ♀ (“YC29974”, DNA voucher “00739238”, genitalia on slide TL654
♀), same data as holotype except “L=19mm”, “Chew” (♂), “Leaf Y”, “L=14mm”,
“Tie” (♀), leg. M. Rimandai, S. Sau, A. Kinibel, M. Jimbudo; 1 ♀ (“YC28238”, DNA
voucher “00739199” & MTD7872, genitalia on slide TL657 ♀), same data as holo-
type except “Chew”, “L=15mm”, 11.ix.2012, leg. M. Rimandai, S. Sau, A. Kinibel,
M. Jimbudo (USNM).
Diagnosis. e well-marked median cubital and dorsal snow-white patches on
the forewing segregate this species from its congeners. Median discoidal stigma is trap-
ezoid, reddish brown, postmedian patch is reddish brown, distally edged by thin white
streak. In male genitalia, the narrow uncus-tegumen connection and the broadly in-
dented 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.
orax (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
éo Léger et al. / ZooKeys 907: 1–99 (2020)
28
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 eld 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.
Abdomen. Male sternum A8 posterior margin bilobed.
Male genitalia (N = 1) (Fig. 54). Uncus slender, entirely sclerotised, forming nar-
row connection to tegumen, narrowed on its middle, apex large, broadly indented.
Gnathos reduced to barely sclerotised band. Tegumen arms dorso-posteriorly not
fused. Valva slender, ventral margin nearly straight, gently bent dorsad on distal 1/4,
dorsal margin convex, valva apex pointed. Juxta with base rounded, medially slightly
narrowed with weakly sclerotised edges, apex broadly rounded, anked on each side
with sclerotised bump covered with setae. Saccus not pronounced. Phallus apically
with sclerotised spine.
Female genitalia (N = 2) (Fig. 96). Papillae anales thick, dorsally and ventrally not
connected. Posterior apophyses bent dorsad. Anterior apophyses widened at posterior
1/3, with tip pointed dorsad. Antrum sclerotisation as long as broad. Ductus bursae
long, more or less straight. Corpus bursae small, globular, reticulated, with sclerotisa-
tion between thorn and corpus opening. orn straight, with small dents pointing
toward thorn base, basally with small outwardly projected extension.
Distribution. Known from Yawan village (1,700 m) in the Eastern Highlands
Province (Papua New Guinea).
Biology. e moths were reared from the fern Diplazium esculentum (Retzius in
Retzius & König, 1791) Swartz, 1803 (Athyriaceae) (S. Miller, C. Redmond & T.
Whiteld, pers. comm.).
Etymology. e species epithet gombongi comes from “gombong”, the name for
fern in the Yau language (https://www.ethnologue.com/language/yuw), referring to the
larval host plant. is name was suggested by Vojtěch Novotný and Gibson Mayiah.
Hoploscopa tonsepi Léger & Nuss, sp. nov.
http://zoobank.org/B0664603-AEC3-4D7B-8A99-146A6359A88C
Figs 13, 55, 101
Material examined. Holotype: ♂, with labels: [exuvia in a capsule pinned under the
specimen] “Sp. PYRA-106 | YP.4B.4041 | CATY 002 L 11 mm | YC 30471 Leaf M
[circled] Y | Roll - Tie - Chew [circled] - Skel | 13 OCT 2012 [handwritten]”; “Papua
New Guinea | Yawan village | 06°10'S, 146°5'E [1700m] | L[e]g[i]t J. Valeba, J. Auga
| M. Dilu, F. Philip, R. Lilip | NG Binatang Res[earch]. C[en]t[e]r.”; “USNM ENT:
PNG | Madang Ecology Project | [barcode] | 00739227” [DNA voucher]; “DNA
2013”; “TL | 655 ♂”. Deposited in USNM.
Twenty-six new species of Hoploscopa 29
Paratypes: 2 ♂, 1 ♀. PAPUA NEW GUINEA: 1 ♂ (“YC30467”, DNA voucher
MTD7874 & “00739207”, genitalia on slide TL661 ♂), same data as holotype except
“CATY 001, L = 15 mm”; 1 ♂ (“YC30472”, DNA voucher “00739208”), same data
as holotype except “CATY 001, L = 15 mm”, leg. B. Gewa, J. Kua, S. Sau, A. Kinibel;
1 ♀ (“YC30473”, DNA voucher “00739234” & MTD7873, genitalia on slide TL658
♀), same data as holotype except “L = 10 mm” “Roll”, leg. B. Gewa, J. Kua, S. Sau, A.
Kinibel (USNM).
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. Pro-
boscis 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.
orax (Fig. 13). Collar pale yellow. Forewing length: 9–10 mm (♂ & ♀); fore-
wing ground colour brown, broadly suused 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, to-
gether with narrow white cubital streak forming a Y; postmedian patch roughly tri-
angular, reddish brown, crossed with brown lines, abutted with pale yellow blotch
at costa; postmedian line broad, marked on costal half; postmedian suusion faintly
marked, pale yellow; subterminal line pale yellow, diuse, running more or less straight
from dorsum distal 1/4 to apex; subterminal eld 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.
Abdomen. Male sternum A8 posterior margin bilobed.
Male genitalia (N = 2) (Fig. 55). Uncus broad, entirely sclerotised, on distal half
extending into two arms with rounded tip, with small dent on inner side, margin be-
tween arms with two small tips on its middle. Gnathos projection plump, ca. 2/3 of un-
cus length. Valva slender, ventral margin nearly straight, dorsal margin slightly convex,
apex pointed. Juxta with base roughly rounded, medially narrowed, apex blunt, slightly
incurved. Saccus slightly quadrangular. Coecum penis reduced to a short protrusion.
Female genitalia (N = 1) (Fig. 101). Papillae anales thick, dorsally and ventrally not
connected. Posterior apophyses absent. Anterior apophyses without bump at posterior
1/3. Antrum sclerotisation short. Ductus bursae short, slender, straight. Corpus bursae
very small, ovoid, covered with tiny papillae. Corpus sclerotisation and thorn absent.
Distribution. Known from the Yawan village (1,700 m) in the Eastern Highlands
Province (Papua New Guinea).
éo Léger et al. / ZooKeys 907: 1–99 (2020)
30
Biology. e moths were reared from Diplazium esculentum (Retzius in Retzius
& König, 1791) Swartz, 1803 (Athyriaceae) (S. Miller, C. Redmond, & T. Whit-
eld, pers. comm.).
Etymology. Named after Tonsep, the village leader of Yawan in Papua New Guinea,
who locally conducted the project on which the larvae were collected and reared to adults.
Hoploscopa marijoweissae Léger & Nuss, sp. nov.
http://zoobank.org/D8FBB3AC-2B4B-436E-B939-70FB9B5E9336
Figs 14, 57, 95
Material examined. Holotype: ♂, with labels: “M[oun]t. Goliath [Mount Yamin], |
5000 f[ee]t., Centr[al]. Dutch N[ew]. Guinea, | ca. 139° long, | February 1911, | (A.
S. Meek).”; "NHMUK 010923397” [barcode appended]; “TL | 710 ♂”. Deposited
in NHMUK.
Paratypes: 1 ♂, 1 ♀. INDONESIA: 1 ♂ (NHMUK010923399), 1 ♀
(NHMUK010923351, genitalia on slide TL 709 ♀), same data as holotype (NHMUK).
Other specimens examined. 1 ♂. PAPUA NEW GUINea: 1 ♂, (DNA voucher
MTD LEP3157 & genitalia on slide TL437 ♂) Morobe Province, Mount Kaindi,
2350 m, 11.xii.1976, leg. G. F. Hevel & R. E. Dietz (USNM).
Diagnosis. is species is recognisable by the brown forewings broadly suused
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 fe-
male 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.
orax (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 eld copper;
median discoidal stigma yellow with copper edges, costally lled 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 eld 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.
Twenty-six new species of Hoploscopa 31
Abdomen. Male sternum A8 posterior margin bilobed.
Male genitalia (N = 1) (Fig. 57). Uncus long and slender, with straight lateral
margin, apex blunt. Gnathos projection ca. half the length of uncus, with blunt apex.
Valva ventral margin nearly straight, in basal half slightly concave, dorsal margin con-
spicuously rounded, apex pointed. Juxta with base rounded, notched on its middle,
medially narrowed, apex rounded, faintly notched on its middle. Saccus triangular.
Phallus with truncate cornutus.
Female genitalia (N = 1) (Fig. 95). Anterior apophyses conspicuously widened
at posterior 1/3, with tip pointing dorsad. Antrum sclerotisation weak, ca. as long
as broad. Ductus bursae of medium length, straight, slender. Corpus bursae large,
globular, posterior half reticulated, anterior half membranous, with weak sclerotisa-
tion between corpus opening and thorn. orn long, slightly curved, with small dents
pointing toward thorn base, basally with small outwardly projected extension.
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 éo Léger’s family.
Remarks. e 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.
Hoploscopa titika Léger & Nuss, sp. nov.
http://zoobank.org/8B31F2F7-6FD3-4C8D-85E2-43D87F3EEF3E
Figs 15, 58
Material examined. Holotype: ♂, with labels: “SUMATRA-Holzweg | 25 km SSW-
Pematangsiantar | straße nach Prapat [road to Prapat], L[icht]F[ang] [light trap] |
13.ii.1996, leg. A. Kallies”; “coll[ection]. M. Nuss | Geschenk 2000 | Museum für
Tier- | kunde Dresden”; “DNA voucher | Lepidoptera | MTD 2016 | [vertically writ-
ten:] no. 3206”; “TL505 | ♂”. Deposited in MTD.
Diagnosis. Hoploscopa titika sp. nov. is recognisable by its well-marked white
quadrangular median cubital spot on the forewing. In male genitalia, the gnathos pro-
jection is reduced to a small ridge, and the dorsal margin of the valva is rounded,
broadly sclerotised. Female genitalia not known.
Similar species. No similar species known.
Description. Head. Antennae dorsally with pale yellow to bronze scales. Probos-
cis white. Maxillary palpi brown, inner side white to pale brown. Labial palpi brown,
ventro-basally white.
orax (Fig. 15). Collar white. Forewing length: 9 mm; forewing ground colour
brown; basal patch extending from dorsum to subcosta, red, with small basal pale yel-
low spot; median discoidal stigma triangular, pale yellow, lled with red scales, with
white trapezoid cubital patch abutting dorsally; postmedian patch roughly quadrangu-
lar, pale yellow, lled with red scales; subterminal line diuse, white; subterminal eld
éo Léger et al. / ZooKeys 907: 1–99 (2020)
32
marked with red and pale yellow; fringe missing on specimen. Forelegs lost. Mid- and
hindlegs white to pale yellow.
Abdomen. Male sternum A8 posterior margin bilobed, with short, rounded lat-
eral projections.
Male genitalia (N = 1) (Fig. 58). Uncus long and slender, gently tapering toward
apex, apex tongue-shaped. Gnathos projection limited to small ridge. Valva broad,
ventral margin bent dorsad on distal 1/3, dorsal margin conspicuously convex, apex
pointed. Juxta with base rounded, medially straight, apex blunt. Saccus small, pointing
dorsad. Phallus apically with sclerotised spine.
Female genitalia. not known.
Distribution. Known from North Sumatra (Indonesia).
Etymology. e species name titika comes from the Indonesian word “titik” mean-
ing spot or dot, referring to the white median cubital spot of the forewing.
Hoploscopa pangrangoensis Léger & Nuss, sp. nov.
http://zoobank.org/B24AA14E-0E31-4CF0-BCDC-85DB29385473
Figs 16, 59, 97
Material examined. Holotype: ♀, with labels: “Indonesia, Java, M[oun]t. Pangrange
| 30 km S[outh]E[ast] Bogor, 1625 m | primary forest 16–20.ii.1996 | 6.30S 107.10E
leg. Siniaev & Afonin”; “Lepidoptera | date: i.2018 | MTD 7433 | [vertically written:]
DNA-voucher”; “TL | 636 ♀”. Deposited in MFNB.
Paratypes: 4 ♂, 1 ♀. INDONESIA: 2 ♂ (1 with genitalia on slide TL659 ♂), 1 ♀
(DNA voucher MTD7431, genitalia on slide TL627 ♀), same data as holotype; 2 ♂ (1
with genitalia on slide TL660 ♂), same data as holotype except 21–26.ii.1996 (MFNB).
Diagnosis. e 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 nger-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 crescent-
shaped pale yellow median discoidal stigma on the forewing and the small median
cubital patch. However, median discoidal stigma and postmedian patch are lled 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.
Twenty-six new species of Hoploscopa 33
orax (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 cres-
cent-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, diuse; 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.
Abdomen. Male sternum A8 posterior margin notched, with short, rounded lat-
eral projections.
Male genitalia (N = 2) (Fig. 59). Uncus medially widened, narrowed on apical
1/4, apex blunt. Gnathos projection nger-shaped, ca. half the length of uncus. Valva
ventral margin straight, gently bent dorsad on distal 1/3, dorsal margin conspicuously
convex; apex slightly pointed. Juxta with base quadrangular, medially narrowing, apex
roughly rounded, slightly notched. Saccus triangular, conspicuously pointing dorsad.
Female genitalia (N = 2) (Fig. 97). Anterior apophyses with dorsal bump at pos-
terior 1/3. Antrum elongated, membranous, with thin longitudinal sclerotised lines,
anteriorly with minute sclerotised markings. Ductus bursae short, bent before cor-
pus opening. Corpus bursae pear-shaped, reticulated, with sclerotisation between
thorn and corpus opening and faintly marked sclerotised band medially. orn small,
straight, with small dents pointing toward thorn apex.
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.
Hoploscopa isarogensis Léger & Nuss, sp. nov.
http://zoobank.org/C77DFE2E-3B02-4738-8F3E-E8139D571FEB
Figs 17, 60, 100
Material examined. Holotype: ♀, with labels: “Philippines, South Luzon | M[oun]
t Isarog | 13°40'N, 123°20'E, 530 m | submontane forest, at light | 22.iii.2000, leg.
M. Nuss”; “DNA Barcode | BC MTD 01431”; “DNA voucher | Lepidoptera | MTD
2016 | [vertically written:] no. 3198”; “TL514 | ♀”. Deposited in MTD.
Paratypes: 2 ♂, 7 ♀. PHILIPPINES: 1 ♂ (DNA vouchers MTD LEP14 & MTD
LEP3209, genitalia on slide TL507 ♂), 5 ♀ (1 with DNA voucher MTD LEP43 &
MTD LEP3208, genitalia on slide TL528 ♀; 1 with DNA barcode BC MTD 01430;
1 with DNA voucher MTD LEP3199 & genitalia on slide TL520 ♀, 1 with DNA
voucher MTD LEP3200 & genitalia on slide TL523 ♀), same data as holotype; 1 ♂, 1
♀ (♂ with genitalia on slide TL762 ♂), same date and locality as holotype, leg. Mey &
éo Léger et al. / ZooKeys 907: 1–99 (2020)
34
Ebert; 1 ♀ (DNA voucher MTD7422 & genitalia on slide TL621 ♀), Laguna, Pangil,
50 m, 11.iv.1997, leg. Mey & Speidel (MTD).
Other specimens examined. 2 ♂, 7 ♀. PHILIPPINES: 1 ♂ (DNA voucher
MTD7420 & genitalia on slide TL626 ♂), 6 ♀ (4 with DNA voucher MTD7418,
MTD8143, MTD8144, MTD8146, MTD8148 and genitalia on slide TL630♀,
TL704 ♀, TL705 ♀, TL720 ♀, TL719 ♀ respectively, 2 with DNA voucher
MTD8145 & MTD8147 and abdomens in microvials), Leyte, Lake Danao, 650 m,
14.-17.4.1997, leg. W. Mey & W. Speidel (MFNB); 1 ♂ (DNA voucher MTD7423
& genitalia on slide TL631 ♂), Mindoro, Mt Baco Pass, 1150 m, 14.i.1998 (Mey &
Samarita); 1 ♀ (DNA voucher MTD7424 & genitalia voucher on slide TL619 ♀),
Mindoro, Mt Halcon, 1300 m, 15–17.i.1998 (Mey & Samarita) (MFNB);
Diagnosis. Hoploscopa isarogensis sp. nov. is a relatively small brown-winged spe-
cies (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 pro-
jection is ca. 1/3 of uncus length, with an indented apex. In female genitalia, the cor-
pus 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. e 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 projec-
tion 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.
Description. Head. Antennae dorsally brown. Proboscis pale yellow. Maxil-
lary palpi brown, base and inner side pale brown. Labial palpi brown, ventro-
basally pale yellow.
orax (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 eld red-
dish brown; median discoidal stigma rhomboid to elliptic, pale yellow, lled with
reddish brown; cubital and dorsal pale yellow patches forming with median discoidal
stigma an oblique band disrupted at veins; postmedian patch roughly triangular, red-
dish brown, more or less speckled with pale yellow, with pale yellow blotch at costa;
subterminal line pale yellow, conspicuously incurved inwards at CuA2; subterminal
eld 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.
Abdomen. Male sternum A8 posterior margin broadly indented, with short,
rounded lateral projections.
Twenty-six new species of Hoploscopa 35
Male genitalia (N = 2) (Fig. 60). Uncus long, medially broadened, narrowed
at apical 1/4, apex duck beak-shaped, medio-ventrally with a small bump projecting
ventrad. Gnathos projection ca. 1/3 of the uncus length, with apex notched. Valva
ventral margin nearly straight, gently bent dorsad on distal 1/4, dorsal margin slightly
convex, apex truncated. Juxta broad, with base rounded, medially slightly narrowed,
apex slightly incurved. Saccus quadrangular.
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 rela-
tively short, nearly straight. Corpus bursae globular and reticulated but in some speci-
mens with a rounded pouch-like extension, with well-marked sclerotisation between
thorn and ductus bursae opening, medially with faintly sclerotised band. orn sabre-
like, curved, with small dents pointing toward thorn base.
Distribution. Known from the Luzon and Mindoro islands (Philippines), be-
tween 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 dierences 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.
Hoploscopa ypsilon Léger & Nuss, sp. nov.
http://zoobank.org/06694A15-8524-4AB3-AAC0-2874511AC4F3
Figs 18, 61, 98
Material examined. Holotype: ♂, with labels: “Philippinen, Luzon | M[oun]t[ai]n.
Prov[ince]., Barlig | 1650 m, 14–15.XI.97 | leg. Mey, Ebert, Nuß”; “DNA voucher |
Lepidoptera | date: ix.2018 | MTD8138 | [vertically written:] DNA-voucher”; “TL699
| ♂”. Deposited in MFNB.
Paratypes: 9 ♂, 15 ♀. PHILIPPINES: 4 ♂, 10 ♀ (1 with DNA voucher
MTD7429 & genitalia on slide TL623 ♀), same data as holotype; 3 ♂ (1 with geni-
talia on slide TL677 ♂), 5 ♀ (1 with DNA voucher MTD7427 & genitalia on slide
TL620 ♀, 1 with DNA voucher MTD7428 & genitalia on slide TL637 ♀), Luzon,
Mountain Province, Chatol, 2100 m, 16–18.xi.1997, leg. Mey, Ebert & Nuss; 1 ♂
(DNA voucher 7435 & genitalia on slide TL624 ♂), Luzon, Ifugao, Mount Polis,
2000 m, 13.xi.1997, leg. Mey, Ebert & Nuss; 1 ♂ (DNA voucher MTD8137), Luzon,
Santa Fe, Bald Mountain, 1150 m, 11–13.xi.1997, leg. Mey, Ebert & Nuss (MFNB).
Diagnosis. Hoploscopa ypsilon sp. nov. displays well-marked white coloured mark-
ings on the forewing. Median discoidal stigma and cubital patch are Y-shaped, post-
median patch is triangular and subterminal line is conspicuously incurved inwards at
CuA2. In male genitalia, the gnathos forms a short, broadly rounded projection.
éo Léger et al. / ZooKeys 907: 1–99 (2020)
36
Similar species. Hoploscopa danaoensis sp. nov. is species displays pale yellow
markings on the forewing, the postmedian area is more or less broadly suused 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 condently separated.
Description. Head. Antennae dorsally with brown scales. Proboscis brown, speck-
led with pale yellow. Maxillary palpi brown, inner side and base pale yellow. Labial
palpi brown, ventral base and tip pale yellow.
orax (Fig. 18). Collar pale yellow. Forewing length: 9–10 mm (♂), 9.5–11 mm
(♀); forewing ground colour brown; basal dark brown blotch with oblique white streak
abutting at costa; median discoidal stigma V-shaped, white, edged with darker brown,
together with white cubital streak forming a Y; postmedian patch triangular, white, in-
vaded with light brown toward costa, edged with darker brown; subterminal line white,
conspicuously incurved inwards at CuA2; fringes brown, with pale yellow dots. Hind-
wing pale brown, darker at apex. Forelegs brown; tarsi brown to pale brown. Midlegs with
femur brown; tibia brown, speckled with pale yellow; tarsi pale yellow. Hindlegs with
femur brown; tibia brown speckled with pale yellow, distally pale yellow; tarsi pale yellow.
Abdomen. Male sternum A8 posterior margin bilobed.
Male genitalia (N = 3) (Fig. 61). Uncus long, slightly narrowed on apical 1/4,
apex duck beak-shaped. Gnathos projection short, with broadly rounded apex. Valva
ventral margin gently bent dorsad on distal 1/3, dorsal margin conspicuously convex,
apex slightly pointed. Juxta with base rounded, medially slightly narrowed, apex in-
curved. Saccus small, quadrangular.
Female genitalia (N = 3) (Fig. 98). Anterior apophyses with dorsal bump at 1/3.
Antrum sclerotisation short, ca. as wide as long. Ductus bursae short, bent before
corpus opening. Corpus bursae pear-shaped, reticulated, with sclerotisation between
thorn and corpus opening and faintly marked sclerotised band medially. orn small,
straight, with small dents pointing toward thorn apex.
Distribution. Known from the Ifugao, the Mountain and the Nueva Vizcaya
Provinces on Luzon (Philippines), at altitudes between 1,150–2,100 m.
DNA barcoding. e highest intraspecic divergence is 0.7%. e nearest neigh-
bor is H. danaoensis sp. nov. (2.9–3.1%).
Etymology. e species name ypsilon, for the Greek letter Y, refers to the shape of
the median markings on the forewing.
Hoploscopa danaoensis Léger & Nuss, sp. nov.
http://zoobank.org/CE945458-F671-4EE7-8E5B-FA18C6CC58AC
Figs 19, 62, 99
Material examined. Holotype: ♂, with labels: “Philipinen [sic], Leyte | Lake Danao, 650
m | 14.-17.4.1997 | leg. Mey & Speidel”; “DNA voucher | Lepidoptera | date: i.2018 |
MTD7419 | [vertically written:] DNA-voucher”; “TL | 632 ♂”. Deposited in MFNB.
Twenty-six new species of Hoploscopa 37
Paratypes: 6 ♂, 11 ♀. Philippines: 6 ♂ (2 with genitalia on slide TL721 ♂ &
TL722 ♂), 11 ♀ (1 with DNA voucher MTD7421 & genitalia on slide TL618 ♀,
1 with DNA voucher MTD8142 & genitalia on slide TL703, 1 with DNA voucher
MTD8141 & genitalia on slide TL702 ♀), same data as holotype (MFNB).
Other specimens examined. 2 ♂, 9 ♀. Philippines: 1 ♂ (genitalia on slide TL681
♂), 1 ♀ (DNA voucher MTD7417 & genitalia on slide TL615 ♀), Mindanao, 1050
m, Mt Agtuuganon, 28.v.-7.vi.1996, leg. W. Mey; 1 ♂ (DNA voucher MTD8140
& genitalia on slide TL701 ♂), 8 ♀ (DNA voucher MTD8139 & genitalia on slide
TL700 ♀), Negros, Patag, Lake Danao, 1400m, 21.5.96, leg. W. Mey (MFNB).
Diagnosis. e forewings of H. danaoensis sp. nov. displays pale yellow median mark-
ings forming a Y. In male genitalia, the uncus displays a straight lateral margin and an ob-
tuse apex, and the tongue-shaped gnathos projection reaches ca. half of the uncus length.
e juxta is progressively narrowing toward the notched apex. Female genitalia are very
similar to several other Hoploscopa species, e.g., H. ypsilon sp. nov.
Similar species. Hoploscopa brunnealis (q.v.), H. ypsilon sp. nov. (q.v.), H.
metacrossa (q.v.).
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.
orax (Fig. 19). Collar pale yellow. Forewing length: 8 mm (♂), 8–9 mm (♀); fore-
wing ground colour brown; basal dark brown blotch distally pale yellow; median discoidal
stigma V-shaped, white to pale yellow, with dark brown basal and distal edges; cubital
patch elongated, white to pale yellow, together with median discoidal stigma forming a Y;
postmedian patch roughly triangular, white to pale yellow, broadly invaded with brown,
edged with darker brown; postmedian line marked on costal half; subterminal line thick,
white to pale yellow, incurved inwards at CuA2, angled at M1; fringes brown, with diuse
pale yellow dots. Hindwing pale brown. Forelegs brown, tarsi pale brown. Midlegs with
femur brown; tibia brown, speckled with pale yellow, distally pale yellow; tarsi pale yellow.
Hindlegs with femur brown; tibia pale yellow, speckled with pale brown; tarsi pale brown.
Abdomen. Male sternum A8 posterior margin straight.
Male genitalia (N = 1) (Fig. 62). Uncus long with straight lateral margin, apex
obtuse. Gnathos projection tongue-shaped, ca. half the uncus length. Valva ventral
margin gently bent dorsad on distal 1/3, dorsal margin conspicuously convex, apex
slightly pointed. Juxta with base quadrangular, tapering toward apex, apex notched.
Saccus small, triangular, pointing dorsad.
Female genitalia (N = 1) (Fig. 99). Anterior apophyses with dorsal bump at pos-
terior 1/3. Antrum short, as long as broad. Ductus bursae short, straight, bent before
corpus bursae. Corpus bursae broad, pear-shaped, reticulated, with elongated sclero-
tisation between thorn and corpus opening, medially with sclerotised band. orn
straight, with small dents pointing toward thorn apex.
Distribution. Known from Lake Danao (650 m) on Leyte island (Philippines).
DNA barcoding. Morphologically similar specimens from Mindanao and Negros
are recovered in two separate MOTUs and show a divergence of 3.9–4.2% with H.
danaoensis sp. nov.
éo Léger et al. / ZooKeys 907: 1–99 (2020)
38
Figures 19–33. Forewing of Hoploscopa species 19 Hoploscopa danaoensis sp. nov., paratype, ♂, Philip-
pines, Leyte, Lake Danao, 650 m, 14–17.iv.1997 (Mey & Speidel) 20 Hoploscopa aurantiacalis Snellen lec-
totype, ♀, NHMUK010923286, Indonesia, Java occ., Pengaleng, 4000 ft, 1893 (genitalia on slide Pyrali-
dae Brit. Mus. Slide N° 20246) 21 Hoploscopa brunnealis Snellen, lectotype, ♀, NHMUK010923292,
Indonesia, Java occ., Pengaleng, 4000 ft, 1893 (genitalia on slide Pyralidae Brit. Mus. Slide N° 20247)
22 Hoploscopa ocellata Hampson, holotype, ♀, NHMUK010923358, Indonesia, Moluccas, Batchian,
iii.1892 (W. Doherty) (genitalia on slide Pyralidae Brit. Mus. Slide N° 20258) 23 Hoploscopa quadripuncta
Rotschild, holotype, ♂, NHMUK010923357, Indonesia, Papua, Utakwa River, 3000 ft, i.1913 (A.F.R.
Wollaston) (genitalia on slide Pyralidae Brit. Mus. Slide N° 20257)
Twenty-six new species of Hoploscopa 39
Etymology. Named after Lake Danao on Leyte where the species is found.
Remarks. Minor dierences were observed in male genitalia of the specimens from
Mindanao and Negros. However, we only had one male on hand for each series. In
absence of further evidence, we refrained from describing further species here.
Hoploscopa aurantiacalis (Snellen, 1895)
Figs 20, 102
Material examined. Lectotype: ♀, with labels: “Lecto- | type” [purple ringed]; “SYN-
| TYPE” [blue ringed]; “F1893 | Java occ. | Pengaleng | 4000’ F[ee]t” [handwrit-
ten, label squared with silver]; “Syncrotaula | aurantiacalis | Snellen | det. M. Shaer,
1967” [handwritten]; “Lectotype | Eudorina | aurantiacalis | Snellen | det. M. Nuß”
[handwritten]; “99.113.”; “♀ | Pyralidae | Brit[ish].Mus[eum]. | Slide N°. | 20246”;
“GU Nr 659 | prep. M. Nuß”; "NHMUK 010923286” [barcode appended]. Depos-
ited in NHMUK.
Diagnosis. e median discoidal stigma, cubital and dorsal patches of the fore-
wings form an oblique thin white streak. In female genitalia, the membranous pouch
anterad from the antrum and the leaf-shaped sclerotisation of the corpus bursae allows
to recognise this species. Male genitalia are not known.
Similar species. No similar species known.
Description. Head. Antennae dorsally brown. Proboscis pale yellow. Maxillary
palpi brown, base pale yellow, inner side pale brown. Labial palpi brown, ventral base
and inner side pale yellow.
Figures 19–33. Continued. 24Hoploscopa semifascia Hampson, ♀, NHMUK010923321, Papua
New Guinea, Southern Highlands, Bosavi, 570 m, 6.i.1986 (D. Agassiz) (genitalia on slide TL740 ♀)
25 Hoploscopa subvariegata Rothschild, ♂, Morobe Province, near Wau, Mt. Kaindi, 2360 m, 27–28.
vii.1983 (S. E. & P. M. Miller) (genitalia on slide TL642 ♂) 26 Hoploscopa persimilis Rothschild, lec-
totype, ♂, NHMUK010923328, Indonesia, Papua, Utawka River, 3000 ft., i.1913 (A.F.R. Wollas-
ton) (genitalia on slide Pyralidae Brit. Mus. Slide N° 20255) 27 Hoploscopa astrapias Meyrick, lecto-
type, ♀, NHMUK010923383, Fiji, Vunîdawa, 2.i.1932 (genitalia on slide Pyralidae Brit. Mus. Slide
N° 20241) 28 Hoploscopa anamesa Tams, ♀, NHMUK010923466, Vanuatu, Aneityum Island, Red
Crest, 3 m NE of Anelgauhat, 1200 ft, vi.1955 (E. Cheesman) (genitalia on slide TL718 ♀) 29Hop-
loscopa nauticorum Tams, allotype, ♀, Samoan, Upolu Island, Malololelei, 2000 ft, 21.ii.1925 (P.
A. Buxton & G. H. Hopkins) 30 Hoploscopa diusa Hampson, lectotype, ♂, NHMUK010923338,
Papua New Guinea, Fergusson Island, x-xi.1894 (A. S. Meek) (genitalia on slide Pyralidae Brit. Mus.
Slide N° 1014 ♂) 31 Hoploscopa triangulifera Hampson, ♀, NHMUK010923453, Biagi, Mam-
bare R., 1600 m, iii.1906 (A. S. Meek) (genitalia on slide TL712 ♀) 32 Hoploscopa anacantha sp.
nov., holotype, ♂, NHMUK010923444, Indonesia, North Sumatra, Dumoga-Bone N. P., Gunung
Mogogonipa, summit, 1008 m, 18–20.x.1985 (genitalia on slide TL759 ♂) 33 Hoploscopa kelama
sp. nov., paratype, ♂, NHMUK010923342 , Indonesia, North Sulawesi, Dumoga-Bone National
Park, Clarke Camp, lower montane forest, 1140 m, x.1985 (Royal Entomological Society of London,
Project Wallace) (genitalia on slide TL759 ♂).
éo Léger et al. / ZooKeys 907: 1–99 (2020)
40
orax (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 eld 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 cos-
tal half; postmedian suusion white; subterminal line white; subterminal eld 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.
Male genitalia. Not known.
Female genitalia (N = 1) (Fig. 102). Anterior apophyses bent ventrad at 1/3,
without bump. Antrum sclerotisation short. Ductus bursae of medium length, with
a rounded pouch at base, one loop on its middle. Corpus bursae rounded, extend-
ing lateral into a broad pouch, posterior half reticulated, anterior half with erect
papillae, with leaf-shaped sclerotisation at corpus opening. orn long, attened,
straight, thinly indented.
Distribution. Recorded from West Java (Indonesia), at an altitude of 1,300 m.
Remarks. Nuss (1998) transferred this species from Eudorina to Hoploscopa and
designated the lectotype.
Hoploscopa brunnealis (Snellen, 1895)
Figs 21, 103
Argyria xiphotoma Meyrick, 1938
Material examined. Lectotype: ♀, with labels: “Lecto- | type” [round label, purple
ringed]; “SYN- | TYPE” [round label, ringed blue]; “F1893 | Java occ[idental]. | Pen-
galeng | 4000’ F[ee]t” [handwritten, squared with silver]; “99.113.” [handwritten] |
“Syncrotaula | brunnealis | Snellen [handwritten] | det. M. Shaer, 1967”; “Lecto-
type | Eudorina | brunnealis | Snellen | det. M. Nuß” [handwritten]; “GU Nr. 660 |
prep. M. Nuß”; “♀ | Pyralidae | Brit[ish].Mus[eum]. | Slide N°. | 20247”; "NHMUK
010923292” [barcode appended]. Deposited in NHMUK.
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.
Similar species. Hoploscopa danaoensis sp. nov., H. metacrossa (q.v.), H. parvimacu-
la sp. nov. (q.v.). Female genitalia of these species display a larger antrum sclerotisation.
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.
Twenty-six new species of Hoploscopa 41
orax (Fig. 21). Forewing length: 10 mm; ground colour brown; basal dash dark
brown distally pale yellow; costal eld tawny; median discoidal stigma and median cu-
bital 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 suused with pale yellow; subterminal line
pale yellow; fringes brown, with pale yellow dots.
Male genitalia. Not known.
Female genitalia (N = 1) (Fig. 103). Anterior apophyses without dorsal bump at
posterior 1/3, Antrum sclerotisation short. Ductus bursae of medium length, slender,
more or less straight. Corpus bursae pear-shaped, with posterior half reticulated, an-
terior half membranous, with faintly marked sclerotisation between thorn and corpus
opening, medially with faintly marked sclerotised band. orn small, straight, with
small dents pointing toward thorn base.
Distribution. Recorded from West Java (Indonesia) at an altitude of 1,300 m.
Remarks. Nuss (1998) transferred this species from Eudorina to Hoploscopa and
designated the lectotype. Argyria xiphotoma Meyrick, 1938 is synonymised with Hop-
loscopa brunnealis in the same paper.
Hoploscopa ocellata (Hampson, 1919)
Figs 22, 104
Material examined. Holotype: ♀, with labels: “Holo- | type” [round label, red ringed];
“Batchian [Bacan islands] | Mar[ch]. 1892 | W. Doherty”; “Pyrocrambia | ocellata |
type ♀. H[a]mps[o]n.” [handwritten]; “♀ | Pyralidae | Brit[ish].Mus[eum]. | Slide N°.
| 20258”; "NHMUK 010923358” [barcode appended]. Deposited in NHMUK.
Diagnosis. e large oblique elliptic white patch at base of the forewing, the me-
dian 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.
Similar species. No similar species known.
Description. Head. Antennae dorsally brown. Proboscis pale yellow. Maxillary
palpi brown, basally pale yellow. Labial palpi brown, ventro-basally pale yellow.
orax (Fig. 22). Collar white. Forewing length: 10 mm; forewing ground colour
brown; large elliptic white basal patch with dark brown edges; basal and distal discoidal
patches of a darker brown; median discoidal stigma there between trapezoid- to square-
shaped, tawny, basally thinly edged with white; postmedian patch semi-elliptic, tawny,
distally with white costal streak; subterminal line more or less straight, disrupted at
M1, white, distally dark brown on dorsal 2/3; subterminal eld suused with white;
fringes brown with lighter spots. Hindwing pale brown. Fore- and hindlegs brown,
with tibia and tarsi segments distally white. Midlegs lost.
Male genitalia. Not known.
éo Léger et al. / ZooKeys 907: 1–99 (2020)
42
Female genitalia (N = 1) (Fig. 104). Papillae anales thin, dorsally and ventrally
not connected. Anterior apophyses bent ventrad at 1/3, without bump. Antrum mem-
branous. Ductus bursae long, narrow, wrinkled, multi-coiled. Corpus bursae ovoid,
with one half reticulate, the other half densely covered with long acanthae, medially
with sclerotised band. orn long, slender, straight, without dents.
Distribution. Recorded from Bacan island in the Moluccas (Indonesia).
Remarks. Nuss (1998) transferred this species from Eudorina to Hoploscopa.
Hoploscopa quadripuncta (Rotschild, 1915)
Figs 23, 63, 105
Material examined. Holotype: ♂, with labels: “Holo- | type” [round label, red ringed];
“Utakwa [sic, Oetakwa] R[iver]., | Dutch N[ew]. Guin[ea]., | 3000 f[ee]t., Jan. 1913. |
A.F.R. Wollaston.”; “Eudorina | quadripuncta | Type Rotsch[ild].”; “436”; “♂ | Pyrali-
dae | Brit[ish].Mus[eum]. | Slide N°. | 20257”; "NHMUK 010923357” [barcode ap-
pended]. Deposited in NHMUK.
Other specimens examined. 3 ♂, 1 ♀. INDONESIA: 1 ♂ (NHMUK010923460,
DNA voucher MTD8249 & genitalia on slide TL734 ♂), Moluccas, Seram, Gunung
Kobipoto, north slopes, 570 m (NHMUK010923460), lowland forest, viii-ix.1987, leg.
J. D. Holloway, D. T. Jones et al.; 1 ♂ (NHMUK010923459, DNA voucher MTD8248
& genitalia on slide TL733 ♂), same data except 900 m. PAPUA NEW GUINEA: 1 ♂,
1 ♀, Hydrographers Mountains, 830 m, i.1918, Rothschild bequest (NHMUK).
Diagnosis. e white-coloured basal patch and median discoidal stigma as well
as the oblique postmedian line allows separation of H. quadripuncta from its conge-
neric species. In male genitalia, uncus apex is cuneate and gnathos projection is ogive-
shaped. In female genitalia, the thorn of the corpus bursae is long, slender, curved, and
shows a well-marked rounded sclerotisation at its base.
Similar species. No similar species known.
Description. Head. Antennae dorsally bronze. Proboscis white. Maxillary palpi brown,
base and inner side pale brown. Labial palpi brown, ventral base and inner side white.
orax (Fig. 23). Collar pale yellow. Forewing length: 8.5–10 mm (♂), 10 mm
(♀); forewing ground colour brown; basal quadrangular oblique white patch, basally
and distally edged with dark brown, with tawny to reddish brown fascia along CuA2
spreading distally into postmedian area up to costa; median white rounded patch edged
with dark brown; postmedian line oblique, straight, thick; fringes brown. Hindwing
pale brown. Forelegs brown. Midlegs with femur brown, tibia and tarsi pale yellow.
Hindlegs pale brown.
Abdomen. Male sternum A8 posterior margin notched.
Male genitalia (N = 3) (Fig. 63). Uncus long and slender, narrowed at apical 1/4,
apex cuneate. Gnathos projection ogival, ca. 1/3 of uncus length. Valva slender, ventral
margin straight on basal 2/3, bent dorsad on apical 1/3, dorsal margin convex, apex
pointed. Juxta with base rounded, medially narrowed, apex broad, weakly sclerotised.
Twenty-six new species of Hoploscopa 43
Saccus broad, triangular, pointing dorsad. Phallus with at spatula-shaped cornutus,
apically with narrow bump.
Female genitalia (Fig. 105). Anterior apophyses with tip pointing upward at pos-
terior 1/3. Antrum with a narrow weakly sclerotised ring. Ductus bursae of medium
length, straight. Corpus bursae ovoid, posterior half reticulated, anterior half membra-
nous, sclerotised between thorn and corpus opening, medially with faintly sclerotised
band. orn 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 dierence 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.
Hoploscopa semifascia (Hampson, 1919)
Figs 24, 106
Material examined. Holotype: ♀, with labels: “Holo- | type” [round label, red ringed];
“Fak-Fak | Dutch NewGuinea | Dec’[19]07 | 1700 f[ee]t | (Pratt)”; “1913-216.”; “Eu-
dorina | semifascia | type ♀ H[a]mps[o]n” [handwritten]; “♀ | Pyralidae | Brit[ish].
Mus[eum]. | Slide N°. | 20253”; "NHMUK 010923320” [barcode appended]. Depos-
ited in NHMUK.
Other specimens examined. 2♀; PAPUA NEW GUINEA: 1♀
(NHMUK010923321, DNA voucher MTD8242, genitalia on slide TL740 ♀),
Southern Highlands, Bosavi, 570 m, 6.i.1986, leg. D. Agassiz (NHMUK); 1 ♀ (DNA
voucher MTD LEP3158, genitalia on slide TL438 ♀), Morobe Province, Wau, Wau
Ecology Institute, 24–26.viii.1983, 1360 m, leg. S. E. & P. M. Miller (USNM).
Diagnosis. e forewings of H. semifascia display an antemedian dark brown fas-
cia edged with pale yellow and a marked postmedian pale yellow patch. Median mark-
ings 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.
Similar species. No similar species known.
Description. Head. Antennae dorsally striped with bronze and brown scales. Pro-
boscis pale yellow. Maxillary palpi brown, base and inner side pale yellow. Labial palpi
brown, base pale yellow, inner side pale yellow to pale brown.
orax (Fig. 24). Collar white. Forewing length: 7 mm; forewing ground colour
brown; antemedian broad transversal dark brown fascia, basal edge pale yellow, in-
curved 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-
éo Léger et al. / ZooKeys 907: 1–99 (2020)
44
wing pale brown. Forelegs brown. Midlegs brown; tarsi pale brown. Hindlegs with
femur brown; tibia pale yellow, speckled with brown; tarsi pale brown.
Male genitalia. Not known.
Female genitalia (N = 3) (Fig. 106). Anterior apophyses widened at posterior
1/3, with dorsal tip. Antrum sclerotisation short, as long as broad. Ductus bursae of
medium length, slightly curved twice. Corpus bursae reticulated, with small rounded
sclerotisation at thorn base. orn long and slender, curved, glabrous, basally with
small outwardly projected extension.
Distribution. Recorded from Fak-Fak (Papua, Indonesia), also known from the
Southern Highlands and the Morobe Province (Papua New Guinea) at altitudes be-
tween 550 m and 1,400 m.
DNA barcoding. e 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) transferred this species from Eudorina to Hoploscopa. Other
specimens examined from Papua New Guinea display the same pattern and identical
genitalia as the holotype from Western New Guinea.
Hoploscopa subvariegata (Rotschild, 1915)
Figs 25, 64, 109
Material examined. Holotype: ♀, with labels: “Holo- | type” [round label, red
ringed]; “Angabunga R[iver]., | a[uent]. of St. Joseph | R., Brit[ish].N[ew].Guin-
ea, | 6000 f[ee]t, upwards. | Nov.[19]04.-Febr.[19]05. | (A. S. Meek).”; “Eudo-
rina | subvariegata | Type Rotsch[ild].” [handwritten]; “♀ | Pyralidae | Brit[ish].
Mus[eum]. | Slide N°. | 20254”; "NHMUK 010923326” [barcode appended]. De-
posited in NHMUK.
Other specimens examined. 14 ♂, 4 ♀. PAPUA NEW GUINEA: 10 ♂ (2 with
genitalia on slide TL478 ♂ & TL642 ♂), 1 ♀ (genitalia on slide TL521 ♀), Morobe
Province, near Wau, Mt. Kaindi, 2360 meters, 27–28 July 1983, leg. S. E. & P. M.
Miller; 1 ♂ (genitalia on slide TL534 ♂), same data except 29–30.viii.1983; 1 ♂,
1♀, same locality, 11.xii.1976, leg. G. F. Hevel & R. E. Dietz; 2 ♂ (1 with DNA
voucher 3156, genitalia on slide TL436 ♂), 1 ♀, same locality, 3.x.1992, leg. V. O.
Becker); 1 ♂, 1 ♀ (genitalia on slide TL471 ♀), Morobe Province, Biaro Road, 2000
m, 25.ix.1992, leg. V. O. Becker (USNM).
Diagnosis. Hoploscopa subvariegata displays four roughly triangular yellow spots
lled with reddish brown on forewing costa. In male genitalia, uncus is long and slen-
der 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. e latter species lacks the subterminal tri-
angular yellow blotch at forewing costa. In the male genitalia of H. persimilis, uncus
Twenty-six new species of Hoploscopa 45
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.
Description. Head. Antennae dorsally with bronze scales. Proboscis pale yellow,
basally brown. Maxillary palpi brown. Labial palpi brown, ventro-basally pale yellow.
orax (Fig. 25). Collar white. Forewing length: 10 mm (♂), 9.5–12 mm (♀);
forewing ground colour brown to dark brown; basal area reddish brown, basally with
small yellow dash, at costa with V-shaped yellow patch lled with reddish brown; an-
temedian cubital snow-white dot; costal eld reddish brown; median discoidal stigma
V-shaped, white, yellow toward costa, lled with reddish brown; postmedian patch
reddish brown, with white to yellow basal edge and pale yellow to yellow costal blotch;
subterminal costal spot pale yellow to yellow, encroached with reddish brown; sub-
terminal eld reddish brown; margin brown, with white dots. Hindwing dirty white,
bronze toward margin. Forelegs femur and tibia brown; tarsi pale yellow. Midlegs with
femur brown; tibia pale yellow, dorso-distally brown; tarsi pale yellow. Hindleg femur
brown; tibia pale yellow, dorso-distally brown; tarsi pale yellow to pale brown.
Abdomen. Male sternum A8 posterior margin bilobed.
Male genitalia (N = 4) (Fig. 64). Uncus long and slender, gently tapering to-
ward apex, apex roughly truncate. Gnathos projection tongue-shaped, ca. 1/3 of uncus
length. Valva ventral margin nearly straight, dorsal margin strongly protruded dorsad,
apex truncate. Juxta with broad rounded base, medially wide, apex broadly incurved,
weakly sclerotised. Saccus broad, triangular, pointing dorsad. Phallus with elongated,
at, spatula-shaped cornutus apically with narrow bump.
Female genitalia (N = 2) (Fig. 109). Anterior apophyses with small dorsal tip at
posterior 1/3. Antrum sclerotisation short, as long as broad. Ductus bursae of medium
length, slender, straight. Corpus bursae large, globular, posterior half reticulated, ante-
rior half membranous, with sclerotisation between thorn and corpus opening, medially
with faintly marked sclerotised band. orn slightly curved, with small dents pointing
toward thorn base.
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) transferred this species from Eudorina to Hoploscopa. Other
specimens examined from Papua New Guinea display the same pattern and identical
genitalia as the holotype from Western New Guinea.
Hoploscopa persimilis (Rotschild, 1915)
Figs 26, 65
Material examined. Lectotype: ♂, with labels: “Lecto- | type” [round label, purple
ringed]; “Utakwa [sic, Oetakwa] R[iver]., | Dutch N[ew]. Guin[ea]., | 3000f[ee]t.,
Jan[uary]. 1913. | A.F.R. Wollaston.”; “439”; “Eudorina | persimilis | Type Rotsch[ild].”;
“Lectotype | Eudorina | persimilis | Rothschild | det. Nuß” [handwritten]; “♂ | Pyralidae
| Brit[ish].Mus[eum]. | Slide N°. | 20255”; "NHMUK 010923328” [barcode appended].
éo Léger et al. / ZooKeys 907: 1–99 (2020)
46
Diagnosis. Hoploscopa persimilis displays three nearly triangular yellow spots lled
with reddish brown on the costa of the forewing. In male genitalia, uncus is long, slen-
der, 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.
orax (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 eld 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 eld
tawny; fringes brown with pale yellow spots. Hindwing pale yellow. Legs missing or
badly preserved.
Male genitalia (N = 1) (Fig. 65). Uncus long, slender, slightly narrowed on api-
cal 1/4, apex duck beak-shaped. Gnathos projection triangular. Valva ventral margin
nearly straight, dorsal margin conspicuously convex, apex roughly rounded. Juxta not
clearly visible on the slide. Saccus broad, triangular, pointing dorsad. Phallus apically
with sclerotised spine.
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) transferred this species from Eudorina to Hoploscopa and
designated the lectotype. e worn-out specimens hampered accurate description.
Hoploscopa diusa (Hampson, 1919)
Figs 30, 66
Material examined. Lectotype: ♂, with labels: “Lecto- | type” [round label, purple
ringed]; “Syn- | type” [round label, blue ringed]; “Fergusson I., | X, XI. [18]94 | (A.
S. Meek).”; “97–80”; “Pyrocrambia | diusa. | type ♂. H[a]mpson.” [handwritten];
“Pyralidae | Brit[ish].Mus[eum]. | Slide N°. | 1014 ♂”; “Lectotype | Eudorina | diusa
| Hampson | det. Nuß” [handwritten]; "NHMUK 010923338 [barcode appended]”.
Deposited in NHMUK.
Other specimens examined. 2 ♂. PAPUA NEW GUINEA: 2 ♂ (1 with
NHMUK010923439 & genitalia on slide TL715 ♂, 1 with NHMUK010923440 &
genitalia on slide TL716 ♂), same data as holotype (NHMUK).
Diagnosis. Hoploscopa diusa 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 tongue-
shaped. Female unknown.
Similar species. Hoploscopa niveofascia sp. nov. (q.v.).
Twenty-six new species of Hoploscopa 47
Description. Head. Antennae dorsally with brown scales. Proboscis white. Maxil-
lary palpi brown, pale brown at base. Labial palpi brown, ventro-basally white.
orax (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, lled with brown, with white costal blotch; postmedian line marked on cos-
tal half; postmedian area medially suused 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.
Abdomen. Male sternum A8 posterior margin notched, with short, rounded lat-
eral projections.
Male genitalia (N = 3) (Fig. 66). Uncus long, narrowed on apical 1/4, apex duck
beak-shaped. Gnathos projection triangular, with rounded apex. Valva broad, ventral
margin conspicuously bent dorsad on apical 1/4, dorsal margin strongly protruding
dorsad, apex pointed. Juxta with base rounded, of constant width, apex rounded. Sac-
cus small, pointing dorsad. No cornutus visible on the slide.
Female genitalia. Not known.
Distribution. Recorded from Fergusson Island (Papua New Guinea).
Remarks. Nuss (1998) transferred this species from Eudorina to Hoploscopa and
designated the lectotype.
Hoploscopa triangulifera (Hampson, 1919)
Figs 31, 67, 110
Material examined. Holotype: ♂, with labels: “Holo- | type” [round label, red ringed];
“Fergusson I[sland]. | [word crossed] xii.[18]95, | (A, S, Meek).”; “.97.204.”; “Eudori-
na | triangulifera | type ♂. H[a]mps[o]n”[handwritten]; “Body re-axed | 6.xii.[19]40
R.S.C.” [handwritten]; “♀ | Pyralidae | Brit[ish].Mus[eum]. | Slide N°. | 20256”;
"NHMUK010923353”[barcode appended]. Deposited in NHMUK.
Other specimens examined. 2♂, 2♀. PAPUA NEW GUINEA: 1 ♂
(NHMUK010923452, genitalia on slide TL750 ♂), same locality as holotype, xi-
xii.1894; 1 ♂ (genitalia on slide TL711 ♂, NHMUK010923454), 1 ♀ (genitalia on
slide TL712 ♀, NHMUK010923453), Biagi, Mambare R., 1600 m, iii.1906, leg. A. S.
Meek; 1 ♀ (NHMUK010923455), Dampier Island [Karkar], ii-iii.1914 (NHMUK).
Diagnosis. e 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.
éo Léger et al. / ZooKeys 907: 1–99 (2020)
48
orax (Fig. 31). orax 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 dened 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 yel-
low and brown postmedian fascia on costal half in some specimens, edged distally by
white subterminal line; fringes brown with white spots. Hindwing pale brown. Fore-
legs 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.
Male genitalia (N = 2) (Fig. 67). Uncus broad, narrowing on distal half, apex trun-
cate. Gnathos projection limited to small ridge. Valva ventral margin nearly straight,
dorsal margin conspicuously convex, apex pointed. Juxta with broad rounded base,
medially wide, apex broadly indented, weakly sclerotised. Saccus broad, quadrangular.
Female genitalia (N = 2) (Fig. 110). Anterior apophyses without dorsal bump
at posterior 1/3. Antrum sclerotisation ca. twice as long as broad. Ductus bursae
short, straight, slender. Corpus bursae globular, reticulated, with sclerotisation be-
tween thorn and corpus opening, and faintly marked sclerotised band medially. orn
long, incurved, with small dents pointing toward thorn base on inner side, glabrous
on outer side.
Distribution. Known from the Madang, the Northern and the Milne Bay Prov-
inces (Papua New Guinea) at altitudes of ca. 1,600 m.
Remarks. Nuss (1998) transferred this species from Eudorina to Hoploscopa. e
handwritten label from Hampson stipulates that the type is a male, while the abdomen
dissected shows it is a female. e abdomen was re-axed to the specimen, suggesting
that either the sex wasn’t identied properly by Hampson or that the wrong abdomen
was reaxed to the specimen.
Hoploscopa astrapias Meyrick, 1886
Figs 27, 68, 107
Material examined. Lectotype: ♀, with labels: “Lecto- | type” [round label, purple
ringed]; “Vunîdawa | Fiji | HP. 2. 1. [19]32” [handwritten]; “HOPLOSCOPA Meyr.”
[handwritten]; “astrapias Meyr.” [handwritten]; “Hoploscopa | astrapias | 1/1 Meyrick
[handwritten] | E. Meyrick det. | in Meyrick Coll.”; “Lectotype | Hoploscopa | astra-
pias | Meyrick | det. M. Nuß” [handwritten]; “Meyrick Coll. | B. M. 1938-290.”; “♀ |
Pyralidae | Brit[ish].Mus[eum]. | Slide N°. | 20241”; “GU 654 | Hoploscopa | astrapias |
Matthias Nuß”; "NHMUK 010923383” [barcode appended]. Deposited in NHMUK.
Other specimens examined. 1 ♂, 1 ♀. FIJI: 1 ♂ (DNA voucher MTD8251,
genitalia on slide TL724 ♂), 1 ♀ (DNA voucher MTD8250, genitalia on slide TL723
♀), Viti Levu, Nandarivatu, 820 m, 14.9.1955, leg. H. W. Simmonds (NHMUK).
Twenty-six new species of Hoploscopa 49
Diagnosis. e 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 round-
ed 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 rough-
ly 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 unam-
biguous 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.
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.
orax (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 eld
reddish brown; median discoidal stigma trapezoid, reddish brown, edged basally and
distally with yellow, median cubital patch triangular, snow white, together with me-
dian discoidal stigma forming a canine tooth shape; post-median patch bean-shaped,
reddish brown with yellow edges, with thick snow white streak abutting dorsally; post-
median area suused with reddish brown; subterminal line white, not reaching dor-
sum; subterminal eld 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.
Male genitalia (N = 1) (Fig. 68). Uncus long, slender, with straight lateral mar-
gin, apex narrow, tongue-shaped, dorsally with sclerotised bristles. Gnathos projection
limited to small ridge. Valva slender, ventral margin nearly straight, dorsal margin
slightly convex, apex rounded. Juxta broad, with base rounded, apex weakly sclero-
tised, slightly incurved. Saccus small, pointing dorsad. Phallus with large at spatula-
shaped cornutus.
Female genitalia (N = 1) (Fig. 107). Anterior apophyses with dorsal bump at
posterior 1/3. Antrum lightly sclerotised, twice as long as broad. Ductus bursae of me-
dium length, broadly curved. Corpus bursae globular, with posterior half reticulated,
éo Léger et al. / ZooKeys 907: 1–99 (2020)
50
anterior half membranous, with weak sclerotisation between thorn and corpus open-
ing. orn long and slender, straight, glabrous.
Distribution. Known from the island of Viti Levu (Fiji), at altitudes between
0 and 800 m.
DNA barcoding. Hoploscopa astrapias shows a divergence of 3.9% with H. an-
amesa and 3.3–4.7% with H. nauticorum.
Phylogenetic relationships. Hoploscopa astrapias, H. anamesa, and H. nauticorum
are recovered together in the ML analysis of the COI barcode (BS = 61). is 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 pro-
jection, the phallus bears a large, attened, spatula-shaped cornutus; in female genita-
lia, 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).
Hoploscopa anamesa Tams, 1935
Figs 28, 69, 108
Material examined. Lectotype: ♀, with labels: “LECTO- | TYPE” [round label, pur-
ple ringed]; “SYN- | TYPE” [round label, blue ringed]; "New Hebrides: | Tanna. |
ix.1930. | L.E.Cheesman. | B.M.1931-30.”; “Lectotype | Hoploscopa | astrapias | an-
amesa | Tams | det. Nuß” [handwritten]; "NHMUK 010923391” [barcode append-
ed]. Deposited in NHMUK.
Other specimens examined. 2 ♂, 2 ♀. VANUATU: 1 ♂ (NHMUK010923465),
1 ♀ (NHMUK010923466, genitalia on slide TL718 ♀), Aneityum Island, Red Crest, 3
[k]m NE of Anelgauhat, 1200ft, vi.1955, leg. E. Cheesman; 1 ♂ (NHMUK010923471,
DNA voucher MTD8254, genitalia on slide TL727 ♂), 1 ♀ (NHMUK010923470,
DNA voucher MTD8255, genitalia on slide TL728 ♀), Aneityum Island, Agathis
Camp, 1150 ft, 21.vii.1971, leg. G. S. Robinson (NHMUK).
Diagnosis. Hoploscopa anamesa (Fig. 28) display forewing and male genitalia simi-
lar to those of H. astrapias. Female genitalia dier from those of H. astrapias in follow-
ing 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.
orax (Fig. 28). Collar white. Forewing length: 10–11 mm (♂ & ♀); forewing
ground colour brown; basal yellow streak along 1A+2A, abutted by cubital red fascia
running up to median area, disrupted by median cubital patch; costal eld reddish
brown; median discoidal stigma trapezoid, red, edged basally and distally with yellow;
Twenty-six new species of Hoploscopa 51
median cubital patch thick, streak-like, snow white; post-median patch bean-shaped,
red with yellow edges, dorsally abutted by thick snow white streak; postmedian area
suused with red; subterminal line white, not reaching dorsum; subterminal eld
broadly marked with red; fringes brown. Hindwing pale yellow, bronze toward dis-
tal margin. Forelegs brown. Midlegs brown to bronze; tibia medially white. Hindlegs
brown to bronze, tibia base dorsally pale yellow.
Male genitalia (N = 1) (Fig. 69). Uncus long, slender, with straight lateral margin,
apex narrow, tongue-shaped, dorsally with sclerotised bristles. Gnathos projection lim-
ited to small ridge. Valva slender, ventral margin nearly straight, dorsal margin convex,
apex rounded. Juxta broad, with base rounded, apex weakly sclerotised, blunt. Saccus
small, pointing dorsad. Phallus with broad at spatula-shaped cornutus.
Female genitalia (N = 1) (Fig. 108). Anterior apophyses with dorsal protuberance
at posterior 1/3. Antrum membranous. Ductus bursae long, coiled. Corpus bursae
globular, with posterior half reticulated, anterior half membranous, with weak scleroti-
sation between thorn and corpus opening. orn long and slender, straight, glabrous.
Distribution. Known from the Fiji (see Remarks), the Aneityum and Tanna is-
lands on Vanuatu, at an altitude of ca. 350 m.
Phylogenetic relationships. Hoploscopa astrapias is recovered as sister species
(BS = 78). See H. astrapias for further remarks.
Remarks. Nuss (1998) raised H. anamesa from subspecies of H. astrapias to spe-
cies. He stated that this species occurs in sympatry with H. astrapias on Fiji.
Hoploscopa nauticorum Tams, 1935
Figs 29, 70
Material examined. Holotype: ♂, with labels: “Holo- | type [round label, red ringed]”;
“SAMOAN ISLAND. [typographed]| Malololelei | Upolu | 24.ii.24. [handwritten]|
P.A.Buxton”; “SAMOA: | Brit.-Mus. | 1935-315.”; “Hoploscopa | astrapias | nautico-
rum Tams [typographed] | Holotype ♂ ”; “♂ | Pyralidae | Brit[ish].Mus[eum]. | Slide
N°. | 20242”; “GU Nr. 655 | prep. M. Nuß”; "NHMUK 010923388 [barcode ap-
pended]”. Deposited in NHMUK.
Allotype: ♀, with labels: “Allo- | type [round label, red ringed]”; “SAMOAN IS.
[typographed] | Upolu Is[land] | Malololelei | 21.ii.1925 | 2,000 f[ee]t | P.A. Buxton |
and G. H. Hopkins [handwritten]”; “SAMOA: | Brit. Mus. | 1935-315.”; “Hoplosco-
pa | astrapias | nauticorum Tams [typographed] | Paratype. | Allotype ♀ [handwrit-
ten]”; "NHMUK 010923389 [barcode appended]”. Deposited in NHMUK.
Other specimens examined. 2 ♂. SAMOA: 2 ♂ (DNA vouchers MTD8252 &
MTD8253, genitalia on slide TL725 ♂ & TL726 ♂), West Samoa, Upolu, Tiavi, 600
m, 24.viii.1974, leg. G. S. Robinson (NHMUK).
Diagnosis. e forewings of H. nauticorum display snow white median, cubital
and dorsal markings together forming a streak disrupted with brown at veins, and a
postmedian triangular red patch, distally with snow white streak. In male genitalia, the
éo Léger et al. / ZooKeys 907: 1–99 (2020)
52
small sclerotised bristles on dorsal side of the uncus apex are unique to this species.
Female genitalia were not investigated.
Similar species. Hoploscopa astrapias (q.v.), H. anamesa (see H. astrapias).
Description. Head. Antennae dorsally bronze. Proboscis brown, speckled with
pale yellow. Maxillary palpi dark brown, basally pale yellow, inner side brown. Labial
palpi dark brown, ventral base pale yellow.
orax (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 quad-
rangular, red, dorso-distally with thick snow white streak; subterminal line white, in-
dented inwardly at CuA2; subterminal eld 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.
Abdomen. Male sternum A8 posterior margin broadly indented, with short,
rounded lateral projections.
Male genitalia (N = 3) (Fig. 70). Uncus long and slender, gently tapering toward
apex, apex narrow, tongue-shaped, dorsally with few small sclerotised bristles. Gna-
thos without posterior projection. Valva ventral margin nearly straight, dorsal margin
strongly protruding dorsad on basal half, apex rounded. Saccus broad, medially slightly
incurved. Phallus with large, at, spatula-shaped cornutus.
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.as-
trapias for further remarks.
Remarks. Nuss (1998) raised H. nauticorum from subspecies of H. astrapias to species.
Hoploscopa anacantha Léger & Nuss, sp. nov.
http://zoobank.org/58D67EDA-0687-479C-87A3-854F58BC369C
Figs 32, 71, 111
Material examined. Holotype: ♂, with labels: “G[unung]. Mogogonipa | summit,
1008m. | 18–20.x.1985”; “INDONESIA : | SULAWESI UTARA, | Dumoga-Bone
N[ational]. P[ark]. [Bogani Nani Wartabone National Park] | October 1985.”; “R[oyal].
Ent[omological]. Soc[iety]. Lond[on]. | PROJECT WALLACE | B.M. 1985-10”;
"NHMUK010923444 [barcode appended]”; “TL 759 | ♂”. Deposited in NHMUK.
Paratypes: 3♂, 3♀ INDONESIA: 2♂ (NHMUK010923449 &
NHMUK010923343), Sulawesi Utara, Danau Mooat, site 22, PHPA chalet, open
habitat, 1080 m, 31.viii.1985, leg. J. D. Holloway; ♂ (NHMUK010923446 &
genitalia on slide TL760 ♂), 1 ♀ (NHMUK010923445), Sulawesi Utara, Dumoga-
Twenty-six new species of Hoploscopa 53
Bone National Park [Bogani Nani Wartabone National Park], “Clarke” Camp, lower
montane forest, 1140 m, leg. Royal Entomological Society of London; 2 ♀ (1 with
NHMUK010923447, DNA voucher MTD8247 & genitalia on slide TL732 ♀, 1
with NHMUK010923448), Sulawesi Utara, Danau Mooat, near Kotamobagu, 1200
m, 27–28.ix.1985, leg. Royal Entomological Society of London (NHMUK).
Diagnosis. Hoploscopa anacantha sp. nov. displays dark brown forewings with ba-
sal, 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 fe-
male genitalia, the elongated bulging sclerotisation without thorn on corpus bursae is
unique to this species.
Similar species. No similar species known.
Description. Head. Antennae dorsally dark brown. Proboscis white. Maxillary
palpi dark brown, basally white. Labial palpi dark brown, ventro-basally white.
orax (Fig. 32). Collar white. Forewing length: 9 mm (♂), 9–10 mm (♀); fore-
wing ground colour dark brown; basal cubital streak and median triangular to streak-
like patch white, edged with darker brown and few reddish brown scales; postmedian
streak white, reddish brown toward middle of wing; fringes dark brown, with white
dots. Hindwing pale brown. Legs brown.
Abdomen. Male sternum A8 posterior margin notched, laterally with short,
rounded lateral projections.
Male genitalia (N = 2) (Fig. 71). Uncus long, slender, with straight lateral margin, apex
blunt. Gnathos projection broad, triangular, ca. half the uncus length, with small rounded
apex. Valva ventral margin nearly straight, dorsal margin convex, apex pointed. Juxta with
base quadrangular, medially narrowed, apex wide, truncate. Saccus small, quadrangular.
Female genitalia (N = 1) (Fig. 111). Anterior apophyses without bump at poste-
rior 1/3. Antrum sclerotisation short, as long as broad. Ductus bursae long, broadly
curved before corpus opening. Corpus bursae globular, covered with erect papillae,
reticulated at corpus opening, with well-marked elongated bulging sclerotisation run-
ning from corpus opening to middle of corpus bursae. orn absent.
Distribution. Known from North Sulawesi (Indonesia) at altitudes between
1,000m and 1,200 m.
Phylogenetic relationships. is 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. is name was suggested by
Francesca Vegliante.
Hoploscopa kelama Léger & Nuss, sp. nov.
http://zoobank.org/69D2A893-2F5A-4E90-9E82-8C6FD059A364
Figs 33, 72, 112
Material examined. Holotype: ♀, with labels: “[Indonesia] North-Sulawesi, Danau
Mooat | east of Kotamobagu, 1150m | 30–31.iii.2000, at light | leg. A. Kallies &
éo Léger et al. / ZooKeys 907: 1–99 (2020)
54
S.Naumann”; “DNA voucher | Lepidoptera | MTD 2016 | no. 3214 [vertically writ-
ten]”; “TL | 532 ♀”. Deposited in MTD.
Paratypes: 6 ♂, 2 ♀. 1 ♂ (DNA voucher MTD LEP3215, genitalia on slide
TL511 ♂), same data as holotype; 2 ♀ (1 with DNA voucher MTD7878 & geni-
talia on slide TL665 ♀, 1 with DNA voucher 7879 & genitalia on slide TL474
♀), same collecting locality as holotype, 25–26.iii.2000, leg. A. Kallies & C. Zorn
(MTD); 2 ♂ (NHMUK010923443, NHMUK010923341), North Sulawesi, Danau
Mooat, near Kotamobagu, 1200 m, 27–28.ix.1985, leg. Royal Entomological Soci-
ety of London, Project Wallace; 3 ♂ (NHMUK010923342, NHMUK010923442,
NHMUK010923441), North Sulawesi, Dumoga-Bone National Park [Bogani Nani
Wartabone National Park], Clarke Camp, lower montane forest, 1140 m, x.1985, leg.
Royal Entomological Society of London, Project Wallace (NHMUK).
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 ridg-
es, and female genitalia show a larger thorn located on posterior half of corpus bursae.
Description. Head. Antennae dorsally striped with brown and ochreous scales.
Proboscis brown. Maxillary palpi brown, basally pale yellow, inner side pale brown.
Labial palpi brown, ventro-basally pale yellow.
orax (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, lled with dark brown; distal dis-
coidal patch dark brown, quadrangular, with basal and distal edges pale yellow; dou-
ble 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.
Abdomen. Male sternum A8 posterior margin notched, with short, rounded lat-
eral projections.
Male genitalia (N = 1) (Fig. 72). Uncus long and slender, medially slightly wid-
ened, conspicuously narrowed at apical 1/4, apex duck beak-shaped, with small median
bump. Gnathos projection limited to a small ridge. Valva ventral margin nearly straight,
dorsal margin slightly convex, apex pointed. Juxta slender, with base slightly quadrangu-
lar, medially narrowed, apex broadly incurved, weakly sclerotised. Saccus quadrangular.
Female genitalia (N = 3) (Fig. 112). Anterior apophyses with small dorsal bump
at posterior 1/3. Antrum sclerotisation broad, ca. twice as long as broad. Ductus bursae
long, broad, looped once, bent at corpus opening. Corpus bursae with posterior half
Twenty-six new species of Hoploscopa 55
reticulated, anterior half membranous. orn located on ductus bursae shortly ahead
of corpus opening, very small, with dents pointing toward thorn apex.
Distribution. Known from North Sulawesi (Indonesia) at altitudes between
1,000m and 1,200 m.
Phylogenetic relationships. e 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 signicant support (BS = 30). ese 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, well-
marked 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. e species epithet kelama comes from the Indonesian “kelam”
meaning dark, referring to the dark patterns of the forewing.
Hoploscopa ignitamaculae Léger & Nuss, sp. nov.
http://zoobank.org/DB39215F-2066-454D-BEAE-BE7D0E523CF4
Figs 34, 73, 113
Material examined. Holotype: ♀, with labels: “[Indonesia] North-Sulawesi, Tangkoko- |
Batuangus-Dua-Saudara Reserve | near Batuputih, primary forest | 600m, 21.iii.2000, at
light | leg. A. Kallies & C. Zorn”; “DNA barcode | BC MTD 01427”. Deposited in MTD.
Paratypes: 6 ♂, 13 ♀. INDONESIA: 4 ♂ (3 with genitalia on slide TL475 ♂,
TL480 ♂, TL545 ♂), 7 ♀ (1 with DNA voucher MTD LEP82, genitalia on slide TL364
♀; 1 with DNA barcode BC MTD 01425), same data as holotype; 3 ♀ (1 with DNA
voucher MTD LEP3216, DNA barcoding BC MTD 01426; 1 with DNA voucher
MTD LEP3218, genitalia on slide TL516 ♀), North Sulawesi, Danau Mooat, east of
Kotamobagu, 1000 m, 25–26.iii.2000, at light, leg. A. Kallies & C. Zorn; 1 ♂ (DNA
voucher MTD LEP3217 & genitalia on slide TL509 ♂) with same collecting locality,
30–31.iii.2000, leg. A. Kallies & S. Naumann (MTD); 1 ♂ (NHMUK010923412),
North Sulawesi, Dumoga-Bone National Park [Bogani Nani Wartabone National
Park], Clarke Camp, lower montane forest, 1140 m, ix.1985, leg. Royal Entomologi-
cal Society of London, Project Wallace; 1 ♀ (NHMUK010923411), same data except
Plot B, lowland forest, 300 m; 2 ♀ (NHMUK010923410, NHMUK010923413),
same data except Gunung Mogogonipa, summit, 1008m, 22–23.ix.1985 (NHMUK).
Diagnosis. e forewings of H. ignitamaculae sp. nov. display marked basal, me-
dian 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-dened elongated sclerotisation of the corpus
bursae with the small slightly curved thorn on its middle is unique to this species.
éo Léger et al. / ZooKeys 907: 1–99 (2020)
56
Figures 34–45. Forewing of Hoploscopa species. 34 Hoploscopa ignitamaculae sp. nov., paratype, ♂,
North Sulawesi, Danau Mooat, east of Kotamobagu, 1000 m, 30–31.iii.2000 (A. Kallies & C. Zorn)
(genitalia on slide TL509 ♂) 35 Hoploscopa albomaculata sp. nov., paratype, ♀, Indonesia, North Sulawe-
si, Danau Mooat, east of Kotamobagu, 1000 m, 25–26.iii. 2000, (A. Kallies & C. Zorn) (genitalia on
slide TL540♀) 36 Hoploscopa sumatrensis sp. nov., holotype, ♂, Indonesia, Sumatra-Holzweg, 25 km
SSW-Pematangsiantar, road to Prapat, 6–26.i.1996 (A. Kallies) (genitalia on slide TL755 ♂) 37Hop-
loscopa mallyi sp. nov., paratype, ♂, Malaysia, Sabah, Mt Kinabalu, Mesilau, 2000 m, 14–17.xi.2006
(W. Mey & K. Ebert) 38 Hoploscopa gracilis sp. nov., holotype, ♂, Indonesia, Sumatra-Holzweg, 25 km
SSW-Pematangsiantar, road to Prapat, 13.ii.1996 (A. Kallies) (genitalia on slide TL754 ♂) 39 Hoploscopa
agtuuganonensis sp. nov., holotype, ♀, Philippines, Mindanao, Mt Agtuuganon, 1050 m, 28.v–7.vi.1996
(W. Mey) (genitalia on slide TL616 ♀) 40 Hoploscopa boleta sp. nov., paratype, ♀, Papua New Guinea,
Morobe, Mt Kaindi, 2360 m, 3.x.1992 (V. O. Becker) (genitalia on slide TL448 ♀) 41 Hoploscopa pseu-
dometacrossa sp. nov., holotype, ♀, Papua New Guinea, W. Hhl, Prv, near Mt. Hagen, Kuk Ag, Res.
Sta., 1600 m, 19–20.viii.1983 (Scott E. & Pamela Miller) (genitalia on slide TL689 ♀) 42 Hoploscopa
metacrossa Hampson, ♀, Papua New Guinea, Morobe Province, near Bulolo, Mt Susu National Reserve,
Araucaria forest, 975m, 27–28.viii.1983 (S. Miller).
Twenty-six new species of Hoploscopa 57
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.
orax (Fig. 34). Collar white. Forewing length: 8.0–9.5 mm (♂ & ♀); forewing
ground colour brown; basal patch yellow, with cubital reddish orange line starting
from its middle, running distally up to postmedian area; costal eld reddish orange;
median discoidal stigma trapezoid, reddish orange, edged with yellow; median cubital
and dorsal patches yellow, not connected; postmedian patch reddish orange, basal mar-
gin undulated, yellow, costal blotch yellow; subterminal eld brown to tawny; fringes
brown with pale yellow spots. Hindwing pale brown. Forelegs brown. Midlegs brown;
tibia distally pale yellow. Hindlegs with femur brown; tibia pale yellow speckled with
brown; tarsi brown.
Abdomen. Male sternum A8 posterior margin broadly incurved, with short,
rounded lateral projections.
Male genitalia (N = 4) (Fig. 73). Uncus medially widened, apex duck beak-
shaped, ventrally with a small bump. Gnathos projection triangular, slender, ca. 1/4 of
uncus length, with rounded apex. Valva ventral margin nearly straight, dorsal margin
convex, apex pointed. Juxta with rounded base, narrowing on basal half, apex blunt.
Saccus slightly quadrangular.
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 run-
ning from corpus opening to its middle. orn on middle of sclerotisation, very small,
slightly curved, with dents pointing toward thorn base.
Distribution. Known from North Sulawesi (Indonesia) at altitudes between
300m and 1,150 m.
Phylogenetic relationships. See H. kelama sp. nov.
Etymology. From the Latin ignitus, set on re, and macula, spot or blot, in refer-
ence 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.
Figures 34–45. Continued. 43 Hoploscopa jubata sp. nov., paratype, ♂, Papua New Guinea, Morobe
Province, near Bulolo, Mt Susu National Reserve, Araucaria forest, 975 m, 27–28.viii.1983 (S. Miller)
44 Hoploscopa jubata sp. nov. , paratype, ♀, Morobe Province, near Bulolo, Mt Susu National Reserve,
Araucaria forest, 975 m, 27–28.viii.1983 (S. Miller) 45 Hindwing scent organ in Hoploscopa metacrossa.
Paratype, ♂, Papua New Guinea, Morobe Province, Wau, 1200 m, 8–14.xii.1976 (G. F. Hevel & R. E.
Dietz) (genitalia on slide TL443 ♂).
éo Léger et al. / ZooKeys 907: 1–99 (2020)
58
Figures 46–51. Male genitalia features of Hoploscopa. For each gure left above: genitalia without phal-
lus; left below: phallus; right above: gnathos drawing; right below: juxta drawing. 46 Hoploscopa albi-
puncta sp. nov., paratype, TL336 ♂ (genitalia without phallus), TL635 ♂ (phallus) 47 Hoploscopa matheae
sp. nov., paratype, TL730 ♂ 48 Hoploscopa sepanggi sp. nov., paratype, TL641 ♂ 49 Hoploscopa cynodonta
sp. nov., paratype, TL531 ♂ 50 Hoploscopa parvimacula sp. nov., holotype, TL510 ♂ 51 Hoploscopa kina-
baluensis sp. nov., paratype, TL342 ♂. Abbreviations: c (cornutus), de (ductus ejaculatorius), g (gnathos),
jx (juxta), s (saccus), ta (tegumen arm), v (valva), vc (vinculum), uc (uncus).
Twenty-six new species of Hoploscopa 59
Figures 52–57. Male genitalia features of Hoploscopa. 52 Hoploscopa luteomacula Nuss, holotype, GU
prep. Nuss 744 ♂ 53 Hoploscopa obliqua, holotype, Pyralidae Brit. Mus. Slide N° BMNH20252 ♂ (phal-
lus with juxta attached) 54 Hoploscopa gombongi sp. nov., holotype, TL653 ♂ 55 Hoploscopa tonsepi sp.
nov., paratype, TL661 ♂ 56 Hoploscopa niveofascia sp. nov., holotype, TL442 ♂ 57 Hoploscopa marijo-
weissae sp. nov., holotype, TL710 ♂.
éo Léger et al. / ZooKeys 907: 1–99 (2020)
60
Figures 58–63. Male genitalia features of Hoploscopa. 58 Hoploscopa titika sp. nov., holotype, TL505
♂ (apical part of juxta attached to phallus) 59 Hoploscopa pangrangoensis sp. nov., paratype, TL659 ♂
60Hoploscopa isarogensis sp. nov., paratype, TL762 ♂ 61 Hoploscopa ypsilon sp. nov., paratype, TL624 ♂
62 Hoploscopa danaoensis sp. nov., holotype, TL632 ♂ 63 Hoploscopa quadripuncta, holotype, Pyralidae
Brit. Mus. Slide N° BMNH20257 ♂.
Twenty-six new species of Hoploscopa 61
Figures 64–69. male genitalia features of Hoploscopa. 64 Hoploscopa subvariegata, TL534 ♂ 65 Hop-
loscopa persimilis, lectotype, Pyralidae Brit. Mus. Slide N° BMNH20255 ♂ (no juxta drawing) 66 Hop-
loscopa diusa, holotype, Pyralidae Brit. Mus. Slide N° BMNH1014 ♂ 67 Hoploscopa triangulifera, TL711
♂ 68Hoploscopa astrapias, TL724 ♂ 69 Hoploscopa anamesa, TL727 ♂.
éo Léger et al. / ZooKeys 907: 1–99 (2020)
62
Figures 70–75. Male genitalia features of Hoploscopa. 70 Hoploscopa nauticorum, TL725 ♂ 71 Hoplosco-
pa anacantha sp. nov., holotype, TL759 ♂ 72 Hoploscopa kelama sp. nov., paratype, TL511 ♂ 73Hop-
loscopa ignitamaculae sp. nov., paratype, TL545 ♂ 74 Hoploscopa albomaculata sp. nov., paratype, TL363
♂ 75 Hoploscopa sumatrensis sp. nov., paratype, TL538 ♂.
Twenty-six new species of Hoploscopa 63
Hoploscopa albomaculata Léger & Nuss, sp. nov.
http://zoobank.org/E458DE68-F0CC-48B1-91D2-8F94B21D51DB
Figs 35, 74, 114
Material examined. Holotype: ♀, with labels: “[Indonesia] North Sulawesi, Danau
Mooat | east of Kotamobagu, 1000m | 25.–26. iii. 2000, at light | leg. A. Kallies & C.
Zorn”; “DNA Barcode | BC MTD 01429”; “TL544 | ♀”. Deposited in MTD.
Paratypes: 9 ♂, 5 ♀. INDONESIA: 5 ♂ (4 with genitalia on slides TL473 ♂, TL479
♂, TL543 ♂, TL646 ♂), 3 ♀ (1 with DNA voucher MTD LEP57 & MTD LEP3213,
genitalia on slide TL529 ♀), same data as holotype; 2 ♂ (1 with DNA voucher MTD
LEP81, genitalia on slide TL363 ♂), 1 ♀ (DNA barcoding BC MTD 01428, genitalia on
slide TL540 ♀), same locality as holotype, 30–31.iii.2000, leg. A. Kallies & S. Naumann
(MTD); 1 ♂ (NHMUK010923450), 1 ♀ (NHMUK010923355), North Sulawesi, Da-
nau Mooat, 1200 m, near Kotamobagu, 27–28.ix.1985 (♂), 9.xi.1985 (♀), leg. Royal En-
tomological Society of London, Project Wallace; 1 ♂ (NHMUK010923451), same data,
Site 22, 1080 m, PHPA chalet, open habitat, 31.viii.1985, leg. J. D. Holloway (NHMUK).
Diagnosis. Hoploscopa albomaculata sp. nov. is unique by virtue of its median trap-
ezoid 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, slen-
der, 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.
orax (Fig. 35). Collar pale yellow. Forewing length: 9 mm (♂), 9–10 mm (♀);
forewing ground colour brown; small basal white dash edged with dark brown; median
discoidal stigma trapezoid, white, with basal and distal edges dark brown; postmedian
patch streak-like, white, pale yellow at costa, edges dark brown; subterminal line dark
brown, basally faintly marked with pale yellow; fringes basally pale yellow, distally
brown. Hindwing pale brown. Forelegs bronze. Mid- and hindlegs with femur brown;
tibia pale yellow speckled with brown; tarsi bronze.
Abdomen. Male sternum A8 posterior margin bilobed, laterally with short, round-
ed lateral projections.
Male genitalia (N = 5) (Fig. 74). Uncus medially slightly widened, narrowed at
apical 1/4, apex roughly truncate, ventrally with a triangular tip pointing posterad.
Gnathos projection short, wide at base, triangular. Valva ventral margin nearly straight,
dorsal margin conspicuously convex, apex pointed. Juxta slender, with base rounded,
narrowing toward apex, apex deeply indented. Saccus small, pointing dorsad.
Female genitalia (N = 3) (Fig. 114). Anterior apophyses without dorsal bump at
posterior 1/3. Ductus bursae long, slender, gently curved twice. Antrum sclerotisation
short, as long as wide. Corpus bursae small, globular, reticulated, with marked elon-
éo Léger et al. / ZooKeys 907: 1–99 (2020)
64
gated sclerotisation from corpus opening to thorn. orn straight, with small dents
pointing toward thorn apex.
Distribution. Known from North Sulawesi (Indonesia) at altitudes between
1,000m and 1,200 m.
Phylogenetic relationships. See H. kelama sp. nov.
Etymology. From the Latin albus, white, and maculatus, covered with spots.
Hoploscopa sumatrensis Léger & Nuss, sp. nov.
http://zoobank.org/22F33777-64CB-4E37-9AB2-41A52020ABEB
Figs 36, 75, 83, 115
Material examined. Holotype: ♂, with labels: “SUMATRA-Holzweg | 25km SSW-
Pematangsiantar | Straße nach Prapat [road to Prapat] | 6.-26.i.1995, leg. A. Kallies”;
“coll[ection]. M. Nuss | Geschenk 2000 | Museum für Tier- | kunde Dresden”; “DNA
voucher | Lepidoptera | date: xi.2018 | MTD8260 | [vertically written:] DNA | -vouch-
er”; “TL755 | ♂”. Deposited in MTD.
Paratypes: 3 ♂, 4 ♀. INDONESIA. 3 ♂ (DNA vouchers MTD9119, MTD9129
& MTD9121 and genitalia on slides TL761 ♂, TL763 ♂, TL538 ♂), 3 ♀ (DNA
vouchers MTD LEP3205, MTD8257, MTD8262 & genitalia on slides TL524 ♀,
TL752 ♀, TL757 ♀), same data as holotype; 1 ♀ (DNA voucher MTD7880 & genita-
lia on slide TL670 ♀), Sumatra, Asahan, Huta Padang, 1990, leg. E. W. Diehl (MTD).
Other specimens examined. 4 ♀, 2 ♀ (DNA vouchers MTD LEP3202, MTD
8258 & genitalia on slides TL530 ♀, TL753 ♀), same data as holotype; 1 ♀ (DNA
voucher MTD LEP3210 & genitalia on slide TL522 ♀), same data as holotype except
23.viii.1989, leg. E. W. Diehl; 1 ♀ (DNA voucher MTD8261 & genitalia on slide
TL756 ♀), same data as holotype except 13.ii.1996 (MTD).
Diagnosis. e forewings of H. sumatrensis sp. nov. display a pale yellow crescent-
shaped median discoidal stigma lled with reddish brown, as well as a postmedian area
broadly suused 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 be-
fore 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.
orax (Fig. 36). Collar pale yellow. Forewing length: 9–10 mm (♂ & ♀); fore-
wing ground colour brown; small basal dark brown dash distally pale yellow; base of
dorsum with patch of pale yellow scales; cubital reddish brown fascia running up to
median area; median discoidal stigma reddish brown, edged with crescent-shaped pale
yellow patch; postmedian patch reddish brown, with pale yellow blotch at costa; sub-
Twenty-six new species of Hoploscopa 65
terminal line broad, pale yellow; subterminal eld tawny to brown; fringes brown, with
pale yellow dots. Hindwing pale yellow. Forelegs bronze. Midlegs with femur brown;
tibia pale yellow to brown; tarsi bronze. Hindlegs brown; tibia pale yellow speckled
with brown; tarsi bronze.
Abdomen (Fig. 83). Male sternum A8 posterior margin bilobed.
Male genitalia (N = 4) (Fig. 75). Uncus medially broad, narrowed at apical
1/4, apex tongue-shaped. Gnathos projection slender, ca. 4/5 of uncus length, apex
tongue-shaped. Valva ventral margin straight, bent dorsad on distal 1/4, dorsal margin
convex, apex pointed. Juxta base rounded, distal half narrowed, apex notched. Saccus
triangular, pointed.
Female genitalia (N = 4) (Fig. 115). Anterior apophyses with dorsal tip at poste-
rior 1/3. Antrum sclerotisation short. Ductus bursae short, bent before corpus open-
ing. Corpus bursae large, pear-shaped, with sclerotisation between thorn and corpus
opening and a median sclerotised band. orn small, straight, with small dents.
Distribution. Known from North Sumatra (Indonesia).
DNA barcoding. Two MOTUs are found in morphologically identical specimens
from the same locality. e K2P-distance between the two MOTUs is 4.1–6%. e
rst MOTU, which is the one of the type material, shows an intraspecic variation of
0.7%. e second MOTU is found in three females (samples BC MTD LEP01422,
MTD8258 and MTD8261) and shows an intraspecic 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.
Hoploscopa mallyi Léger & Nuss, sp. nov.
http://zoobank.org/13552C8E-ED5E-447B-859A-8402DD48BA72
Figs 37, 76, 116
Material examined. Holotype: ♀. “Borneo, Mt Kinabalu | Headquarter, 1600m |
10–13.XI.2006, L[icht]F[ang] | leg. W. Mey & K. Ebert”; “Liwagu River | 1500m,
Turm”; “DNA voucher | Lepidoptera | M. Nuss 2007 | [vertically written:] no. 124”;
“TL366 | ♀”. Deposited in BORN.
Paratypes: 4 ♂, 2 ♀. MALAYSIA: 1 ♂, 1 ♀ (genitalia on slides TL533 ♂ and
TL340 ♀ respectively), Sabah, Kinabalu National Park Headquarters, 250 m from
Pandanus Trail starting point, 6°0'34"N, 116°32'20"E 1640 m, UV light, 07.vi.2015,
leg. T. Léger & R. Mally; 2 ♂ (1 with genitalia on slide TL339 ♂), Sabah, Mesilau log-
ging site, 400 m before entrance to Mesilau Nature Reserve, 6°2'22"N, 116°35'54"E,
1930 m, UV light, 02.vi.2015, T. Léger & R. Mally; 1 ♀ (DNA voucher 3197, geni-
talia on slide TL513 ♀), Sabah, Kinabalu Park Headquarters, ca. 300 m from starting
point of Kiau View Trail, 6°0'25"N, 116°32'21"E, 1660 m, UV light, 06.vi.2015,
T. Léger & R. Mally (MTD); 1 ♂, Borneo, Mt Kinabalu, Mesilau, 2000 m, 14–17.
xi.2006, W. Mey & K. Ebert (MFNB).
éo Léger et al. / ZooKeys 907: 1–99 (2020)
66
Figures 76–81. Male genitalia features of Hoploscopa. 76 Hoploscopa mallyi sp. nov., paratype, TL339
♂ 77 Hoploscopa gracilis sp. nov., paratype, TL539 ♂ 78 Hoploscopa agtuuganonensis sp. nov., paratype,
TL675 ♂ 79 Hoploscopa boleta sp. nov., holotype, TL440 ♂ 80 Hoploscopa pseudometacrossa sp. nov.,
paratype, TL441 ♂ (uncus apex missing) 81 Hoploscopa metacrossa, TL443 ♂.
Twenty-six new species of Hoploscopa 67
Diagnosis. e forewings of H. mallyi sp. nov. display pale yellow crescent-shaped
median discoidal stigma lled 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.
Similar species. Hoploscopa agtuuganonensis sp. nov., H. gracilis sp. nov. In H. ag-
tuuganonensis sp. nov., median discoidal stigma together with cubital and dorsal patch-
es 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.
orax (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 eld red; median discoidal stigma crescent-shaped, pale yellow, lled 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 suused with pale yellow and iridescent scales; subter-
minal eld 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.
Male genitalia (N = 2) (Fig. 76). Uncus slender, medially narrowed, apex spatu-
late. Gnathos projection slender, ca. 2/3 of uncus length, apex blunt. Valva ventral
margin gently bent dorsad on distal 1/3, dorsal margin convex, apex rounded. Juxta
ogival, with base quadrangular, gently narrowing toward apex, apex narrow, notched.
Saccus triangular, pointing dorsad.
Female genitalia (N = 3) (Fig. 116). Anterior apophyses with dorsal bump at
posterior 1/3. Antrum sclerotisation forming a short ring. Ductus bursae long, broad,
forming two large curves. Corpus bursae small, globular, reticulate, sclerotised between
thorn and corpus opening, medially with broad faintly sclerotised band. orn sabre-
like, curved, with small dents pointing toward thorn base, basally with conspicuous
outwardly projected extension.
Distribution. Known from the slopes of Mount Kinabalu on Borneo, at altitudes
between 1,600 m and 2,000 m.
Phylogenetic relationships. is species is recovered in a clade with H. agtuu-
ganonensis sp. nov. and H. gracilis sp. nov. in the ML analysis of the COI barcode
(BS=92). e morphology of these species is in agreement with the molecular nd-
éo Léger et al. / ZooKeys 907: 1–99 (2020)
68
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.
Etymology. is species is dedicated to our colleague and friend Richard Mally, an
eminent lepidopterologist.
Hoploscopa gracilis Léger & Nuss, sp. nov.
http://zoobank.org/CDC1F535-1AB1-469F-B6DE-082810E8E28F
Figs 38, 77, 117
Material examined. Holotype: ♂, with labels: “SUMATRA-Holzweg | 25km SSW-
Pematangsiantar | Straße nach Prapat [road to Prapat], L[icht]F[ang] [light trap] |
13.ii.1996, leg. A. Kallies”; “coll[ection]. M. Nuss | Geschenk 2000 | Museum für
Tier- | kunde Dresden”; “DNA voucher | Lepidoptera | date: xi.2018 | MTD8259 |
[vertically written:] DNA- | voucher”; “TL754 | ♂”. Deposited in MTD.
Paratypes: 4 ♂, 2 ♀. INDONESIA: 2 ♂ (1 with DNA voucher MTD7876 &
genitalia on slide TL664 ♂, 1 with DNA voucher MTD LEP3203 & genitalia on
slide), North Sumatra, Sipirok, 1450 m, 27–28.i.1995, leg. A. Kallies; 1 ♂ (DNA
voucher MTD8256 & genitalia on slide TL751 ♂), North Sumatra, Mount Sibayak,
03°14'19"N, 98°29'52"E, 1900 m, 02.iii.2002, leg. U. Buchsbaum; 1 ♂ (genitalia on
slide TL539 ♂), 1 ♀ (DNA voucher MTD8263 & genitalia on slide TL758 ♀), North
Sumatra, Mount Sibayak, 03°14'13"N, 98°29'41"E, 1750 m, 07.vii.2000, leg. U. Bu-
chsbaum; 1 ♀ (DNA voucher MTD LEP3204 & genitalia on slide TL527 ♀), North
Sumatra, Dairi, near Sumbul, 2°46'N, 98°32'E, 1670 m, 20.ii.1999, leg. U. Buchsbaum.
Other specimens investigated. 1 ♀. MALAYSIA: 1 ♀ (NHMUK010923415, DNA
voucher MTD8239 & genitalia on slide TL737 ♀), West Malaysia, Cameron Highlands,
Gunung Brinchang, 1980 m, 23–31.x.1989 (G. S. Robinson & M. A. Tobin) (NHMUK).
Diagnosis. Hoploscopa gracilis sp. nov. displays pale yellow crescent-shaped median
discoidal stigma lled with reddish brown, together with median cubital patch form-
ing 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 bur-
sae 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.). Me-
dian 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 dier 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.
Twenty-six new species of Hoploscopa 69
orax (Fig. 38). Collar pale yellow. Forewing length: 9 mm (♂), 11 mm (♀); fore-
wing ground colour brown; broad elongated basal patch reddish brown with basal edge
pale yellow, distally running up to median cubital patch; costal eld reddish brown; me-
dian discoidal stigma crescent-shaped, pale yellow, lled with reddish brown; median
cubital patch streak-like, pale yellow, disrupted at 1A+2A, together with median dis-
coidal stigma forming a Y; postmedian triangular patch reddish brown, speckled with
pale yellow, with pale yellow blotch at costa; postmedian area suused with a mix of
pale yellow and iridescent scales; subterminal eld 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.
Abdomen. Male sternum A8 posterior margin notched, with short, rounded lat-
eral projections.
Male genitalia (N = 4) (Fig. 77). Uncus long and slender, gently tapering toward
apex, apex spatulate. Gnathos projection slender, ca. 2/3 of uncus length, with truncate
apex. Valva slender, ventral margin nearly straight, gently bent dorsad on distal 1/4, dor-
sal margin convex, apex pointed. Juxta with base rounded, medially straight, widening at
distal 1/3 onto duck beak-shaped apex. Saccus triangular, conspicuously pointing dorsad.
Female genitalia (N = 2) (Fig. 117). Anterior apophyses with dorsal bump at
posterior 1/3. Antrum sclerotisation as long as wide. Ductus bursae long, with one
loop, curved before corpus opening. Corpus bursae small, globular, reticulated, with
sclerotisation between thorn and corpus opening and faintly sclerotised band medially.
orn small, plump, slightly curved, with small dents pointing toward thorn base,
basally with small outwardly projected extension.
Distribution. Known from North Sumatra, at altitudes between ca. 1,200 m
to 1,900 m.
DNA barcoding. e species shows an intraspecic 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.
Hoploscopa agtuuganonensis Léger & Nuss, sp. nov.
http://zoobank.org/EA83FE68-F42D-4481-AB44-D1258E5DEBD4
Figs 39, 78, 118
Material examined. Holotype: ♀, with labels: “Philippinen | Mindanao, 1050m | Mt
Agtuuganon | 28.5–7.6.[19]96, leg. MEY”; “DNA voucher | Lepidoptera | date: i.2018
| MTD7416 | [vertically written:] DNA-voucher”; “TL | 616 ♀”. Deposited in MFNB.
Paratypes: 12 ♂ , 4 ♀. PHILIPPINES: 12 ♂(1 with DNA voucher MTD8149 &
genitalia on slide TL708 ♂, 2 with genitalia on slide TL675 ♂, TL676 ♂), 4 ♀ (1 with
DNA voucher MTD7415 & genitalia on slide TL617 ♀), same data as holotype (MFNB).
Diagnosis. In H. agtuuganonensis sp. nov., median markings of the forewing form
together a pale yellow band progressively narrowing toward dorsum, faintly disrupted
éo Léger et al. / ZooKeys 907: 1–99 (2020)
70
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.).
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.
orax (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 cubi-
tal and dorsal patches abutting dorsally; costal eld reddish brown; median discoidal
stigma trapezoid, pale yellow, lled 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 suused with pale
yellow; subterminal line pale yellow; subterminal eld 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.
Abdomen. Male sternum A8 posterior margin bilobed.
Male genitalia (N = 2) (Fig. 78). Uncus long and slender, gently tapering toward
apex, narrowed at apical 1/4, apex spatulate. Gnathos projection slender, ca. 2/3 of
uncus length, apex blunt. Valva ventral margin straight, bent dorsad on distal 1/3, dor-
sal margin convex, apex rounded. Juxta with base quadrangular, with straight lateral
margin, apex blunt. Saccus triangular, conspicuously pointing dorsad.
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 dened scleroti-
sation between thorn and corpus opening. orn plump, straight, with small dents
pointing toward thorn base, basally with small outwardly projected extension.
Distribution. Known from the slopes of Mount Agtuuganon (1,660 m) on Mind-
anao Island (Philippines), at an altitude of 1,050 m.
Phylogenetic relationships. See H. mallyi sp. nov.
Etymology. Named after Mount Agtuuganon on Mindanao Island (Philippines),
where the specimens were collected.
Hoploscopa boleta Léger & Nuss, sp. nov.
http://zoobank.org/D717911D-F973-4401-BC2B-873E5ACED283
Figs 40, 79, 119
Material examined. Holotype: ♂, with labels: “Col[lection] BECKER | PNG 1562”,
“ PAPUA NEW GUINEA | Morobe, M[oun]t Kaindi | 2360m 3.x.1992 | V. O.
Twenty-six new species of Hoploscopa 71
Becker Col[lection]; “ DNA voucher | Lepidoptera | MTD2016 | [vertically written:]
no. 3160”; “TL440 | ♂”. Deposited in USNM.
Paratypes: 2 ♀. PAPUA NEW GUINEA: 1 ♀ (DNA voucher MTD LEP3168,
genitalia on slide TL448 ♀), same data as holotype; 1 ♀ (DNA voucher MTD LEP3159),
same locality as holotype, 27–28.vii.1983, leg. S.E. & P. M. Miller (USNM).
Diagnosis. e 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.
Description. Head. Antenna dorsally striped with ochreous and bronze scales.
Proboscis pale yellow, basally brown. Maxillary palpi brown, base and inner side pale
yellow. Labial palpi brown, ventro-basally pale yellow.
orax (Fig. 40). Collar pale yellow. Forewing length: 9–10 mm (♂ & ♀); fore-
wing ground colour brown; basal dash dark brown, distally pale yellow; basal and
distal discoidal patches quadrangular, dark brown, basally and distally edged with pale
yellow; median discoidal stigma there between trapezoid, light brown; dorsal median
patch broad, pale yellow; postmedian patch light brown, with pale yellow blotch at
costa; postmedian area suused with pale yellow near costa; subterminal line dark
brown, diuse; fringe brown, with pale yellow dots. Hindwing pale yellow, darker at
apex. Forelegs brown, tarsi bronze. Mid- and hindlegs with femur brown, tibia pale
yellow, speckled with brown, tarsi bronze.
Abdomen. Male sternum A8 posterior margin broadly indented.
Male genitalia (N = 1) (Fig. 79). Uncus slender, conspicuously narrowed at api-
cal 1/4, apex duck beak-shaped, bearing thick setae, ventrally with three small ridges.
Gnathos projection ca. 1/3 of the uncus length, gently narrowing toward apex, apex
tongue-shaped. Valva ventral margin nearly straight, dorsal margin slightly convex,
apex slightly blunt. Juxta mushroom-shaped, with base rounded, laterally with two
conspicuous projections, abruptly narrowed at basal 1/3, apex spatulate. Saccus small,
pointing dorsad.
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. orn straight, with small dents pointing
toward thorn base.
éo Léger et al. / ZooKeys 907: 1–99 (2020)
72
Distribution. Known from Mount Kaindi in the Morobe Province (Papua New
Guinea), at an altitude of 2,360 m.
Phylogenetic relationships. is species displays an uncus apex similar to those
of H. jubata sp. nov., H. metacrossa, and H. pseudometacrossa sp. nov. and is possibly
related to them.
Etymology. From the Latin boletus, in reference to the mushroom shape of the juxta.
Hoploscopa pseudometacrossa Léger & Nuss, sp. nov.
http://zoobank.org/E21452AB-8784-4A67-B270-1802A4D7293A
Figs 41, 80, 120
Material examined. Holotype: ♀, with labels: “PAPUA NEW GUINEA: W[estern].
H[ig]hl[and] | Pr[o]v[ince], n[ea]r Mt. Hagen, Kuk Ag | Res[earch]. Sta[tion]., 1600m,
UV Lite | 19–20 August 1983 | Scott E. & Pamela Miller”; “DNA voucher | Lepidop-
tera | date: i.2018 | MTD 7898 | [vertically written:] DNA-voucher”; “TL689 | ♀”.
Deposited in USNM.
Paratypes: 1 ♂, 1 ♀. PAPUA NEW GUINEA: 1 ♂, 1 ♀ (♂ with DNA voucher
MTD LEP3161 & genitalia on slide TL441 ♂; ♀ with DNA voucher MTD LEP3155
& genitalia on slide TL435 ♀), same data as holotype (USNM).
Diagnosis. e 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 thin-
ner, 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.
orax (Fig. 41). Collar pale yellow. Forewing length: 9–10 mm (♂ & ♀); fore-
wing 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 suused 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.
Twenty-six new species of Hoploscopa 73
Abdomen. Male sternum A8 posterior margin broadly indented, with short,
rounded lateral projections.
Male genitalia (N = 1) (Fig. 80). Uncus slender, narrowed on apical 1/4 which
bears thick setae, apex missing on slide examinated. Gnathos projection thumb-shaped,
ca. 1/4 of uncus length. Valva ventral margin bent dorsad on distal half, dorsal margin
medially angled, apex roughly rounded. Juxta with base rounded, medially narrowed,
apex notched. Saccus broad, triangular, pointing dorsad.
Female genitalia (N = 2) (Fig. 120). Anterior apophyses with dorsal bump at
posterior 1/3. Antrum sclerotisation twice as long as broad. Ductus bursae short, more
or less straight, bent before corpus opening. Corpus bursae large, posterior half re-
ticulated, medially covered with erect acanthae, anterior half membranous, with weak
sclerotisation at thorn base. orn straight, with small dents pointing toward thorn
base, basally with small outwardly projected extension.
Distribution. Known from Mount Hagen in the Morobe Province (Papua New
Guinea), at an altitude of 1,600 m.
Phylogenetic relationships. e hindwing scent scales observed in males of H.
jubata sp. nov., H. pseudometacrossa sp. nov., and H. metacrossa suggest a close relation-
ship between these three species.
Etymology. e name is made by the apposition of the prex pseudo- from
the Greek pseudes, false and metacrossa, referring to the resemblance of this species
with H. metacrossa.
Hoploscopa metacrossa (Hampson, 1917)
Figs 42, 45, 81, 84, 86, 121
Material examined. Holotype: ♂, with labels: “Holo- | type” [round label, red ringed];
“Fak-Fak | Dutch New Guinea | Dec’[19]07 | 1700f[ee]t | (Pratt)”; “1913-216”;
“Scoparia | metacrossa | type ♂. H[a]mps[o]n.” [handwritten]; “Pyralidae | Brit[ish].
Mus[eum]. | Slide N°. | 3612”; "NHMUK 010923297” [barcode appended]. Depos-
ited in NHMUK.
Other specimens examined. 12 ♂, 38 ♀. PAPUA NEW GUINEA: 10 ♂ (3 with
DNA vouchers MTD7899, MTD7900, MTD7901 and genitalia on slides TL690 ♂,
TL691 ♂, TL692 ♂ respectively; 1 with genitalia on slide TL482 ♂), 25 ♀ (1 with
genitalia on slide TL481 ♀), Morobe Province, near Bulolo, Mt Susu National Re-
serve, Araucaria forest, 975m, 27–28.viii.1983, leg. S. Miller; 1 ♂ (genitalia on slide
TL472♂), 9 ♀ (2 with DNA vouchers MTD LEP3166, MTD LEP3167 and genitalia
on slides TL446 ♀, TL447 ♀), Morobe Province, Wau, Wau Ecological Institute, mon-
tane forest, 1200 m, 12–24.vii.1983 (2 ♀), 25–31.vii.1983 (1 ♂, 3 ♀), 1–10.viii.1983
(1 ♀), 23–31.viii.1983 (3 ♀), leg. S. E. & P. M. Miller; 4 ♀, same collecting data except
“trap at zoo pond”, 25–27.vii.1983; 1 ♂ (DNA voucher MTD LEP3163 & genitalia on
slide TL443 ♂), Morobe Province, Wau, 1200 m, 8–14.xii.1976, leg. G. F. Hevel & R.
E. Dietz (USNM).
éo Léger et al. / ZooKeys 907: 1–99 (2020)
74
Diagnosis. e forewings of H. metacrossa display Y-shaped median and a post-
median pale yellow patches observed in several other Hoploscopa species (e.g., H. da-
naoensis sp. nov.). Male hindwing displays an androconial organ on the dorsum. e
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, tongue-
shaped 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. kinabaluen-
sis 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 bur-
sae 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.
orax (Fig. 42, 45). Collar pale yellow. Forewing length: 9–10 mm (♂ & ♀);
forewing ground colour brown; basal dark brown spot distally pale yellow; median
discoidal stigma trapezoid, pale yellow, together with cubital pale yellow patches
forming a Y; postmedian patch pale yellow speckled with brown, distal edge dark
brown; postmedian line marked on costal half; postmedian area variously suused
with pale yellow; subterminal line pale yellow; fringes 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 pale yel-
low, speckled with brown; tarsi brown.
Abdomen (Fig. 84, 86). Male sternum A8 posterior margin broadly indented, with
short, rounded lateral projections.
Male genitalia (N = 3) (Fig. 81). Uncus slender, narrowed at apical 1/4, apex
spatulate, bearing thick setae, ventrally with ve small ridges. Gnathos projection ca.
1/4 of uncus length, slender, tongue-shaped. Valva ventral margin curved dorsad on
distal half, dorsal margin conspicuously convex, apex rounded. Juxta with base round-
ed or slightly quadrangular, narrowing at basal 1/4, apex tongue-shaped. Saccus tri-
angular, conspicuously pointing dorsad. Phallus with elongated, at, spatula-shaped
cornutus with subapical tip.
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. orn
straight, with small dents pointing toward thorn base, basally with small outwardly
projected extension.
Twenty-six new species of Hoploscopa 75
Distribution. Known from the Papua (Indonesia) and the Morobe Provinces
(Papua New Guinea) in New Guinea, at altitudes between 600 and 1,200 m.
Phylogenetic relationships. See H. boleta sp. nov. and H. pseudometacrossa sp. nov.
Remarks. Nuss (1998) transferred this species from Eudorina to Hoploscopa.
Hoploscopa jubata Léger & Nuss, sp. nov.
http://zoobank.org/F81F0837-5324-4B81-AC72-B0519F850301
Figs 2, 43, 44, 82, 122
Material examined. Holotype: ♂, with labels: “PAPUA NEW GUINEA | Morobe
Prov[ince]., n[ea]r Bulolo | Mt Susu Nat[ional]. Res[erve]., 975m | 27–28 Aug.1983,
S. Miller | UV Lite, Araucaria For[est].”; “ DNA voucher | Lepidoptera | MTD2016 |
[vertically written:] no. 3164”; “TL444 | ♂”. Deposited in USNM.
Paratypes: 4 ♂, 24 ♀. PAPUA NEW GUINEA. 4 ♂ (1 with genitalia on slide
TL447 ♂, 1 with genitalia on slide TL542 ♂, 1 with wing preparation TL706), 21 ♀ (1
with DNA voucher MTD LEP3165 & genitalia on slide TL445 ♂, 2 with genitalia on
slide TL476 ♀ & TL541 ♀), same data as holotype. 1 ♀, Morobe Province, Wau, Wau
Ecological Institute, 12–24.vii.1983; 1 ♀, same data except 1–10.viii.1983; 1 ♀, Morobe
Province, Wau, 1200 m, 8–14.xii.1976, at black light, leg. G. F. Hevel & R. E. Dietz
(USNM).
Diagnosis. is species shows a strong sexual dimorphism. In males, median and
postmedian markings are white, and postmedian area is broadly suused 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 in-
dented 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.
orax (Figs 2, 43, 44). Collar brown. Forewing length: 9–10 mm (♂ & ♀); fore-
wing ♂ ground colour brown; basal narrow white streak, distally marked with yellow
and orange; median discoidal stigma white, lled 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 suusion; 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, lled 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-
éo Léger et al. / ZooKeys 907: 1–99 (2020)
76
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
eld marked with reddish orange and yellow; fringe brown with pale yellow dots. Hind-
wing pale yellow; in males, presence of an androconial organ on the dorsum of the hind-
wing. 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 modied scales. Ster-
num A8 posterior margin straight.
Male genitalia (N = 3) (Fig. 82). Uncus long and slender, basally narrow, medially
slightly widened, narrowed at apical 1/4, apex spatula-shaped, ventrally with eight small
ridges. Gnathos projection slender, ca. 1/3 of the uncus length. Valva gently bending
dorsad from distal 1/3, dorsal margin slightly convex, apex roughly truncate, forming a
furrow ring. Juxta mushroom-shaped, with base roughly rounded, slightly concave on
its middle, with two conspicuous lateral extensions pointing outward, conspicuously
narrowed at basal 1/4, apex bilobed. Saccus broad, triangular, pointing dorsad.
Female genitalia (N = 3) (Fig. 122). Anterior apophyses with dorsal bump at
posterior 1/3. Antrum sclerotisation short. Ductus bursae very long, forming four con-
spicuous loops bent before corpus opening. Corpus bursae small, globular, reticulated,
medially with light broad sclerotised patch, with weak rounded sclerotisation at corpus
opening. orn gently curved, with small dents pointing toward thorn base, basally
with small outwardly projected extension.
Distribution. Known from Wau and from Mount Susu in the Morobe Province
(Papua New Guinea), at altitudes between 950 m and 1,200 m.
Phylogenetic relationships. See H. boleta sp. nov. and H. pseudometacrossa sp. nov.
Etymology. From the Latin jubatus, having a mane, referring to the dense hair
covering on the inner side of the valva in male genitalia.
Remarks. is species is the only one known from the genus to exhibit a pro-
nounced sexual dimorphism.
Misplaced species
Hoploscopa mediobrunnea (De Joannis, 1929) from Vietnam has been provisionally placed
in Hoploscopa by Nuss (1998). He justied his choice as follows: ““Since Eudorina Snellen
is a homonym of Eudorina Ehrenberg, 1832 (Protozoa), I place E. mediobrunnea here pre-
liminarily 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. e 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 dierent from all Hoploscopa species we have exam-
ined to date, and we therefore assume that H. mediobrunnea belongs to a dierent genus.
Twenty-six new species of Hoploscopa 77
Figures 82–86. Male genitalia and abdomen features of Hoploscopa. 82 Hoploscopa jubata sp. nov.,
paratype, TL542 ♂ 83 Tympanal organs of Hoploscopa sumatrensis sp. nov., paratype, TL538 ♂ 84 Male
tympanal organs of Hoploscopa metacrossa, TL472 ♂ 85 Sternite VIII of Hoploscopa sp. near isarogensis,
TL626 ♂ 86 Sternite VIII of Hoploscopa metacrossa, TL443 ♂. Abbreviations: tb (tympanic bridge), td
(tympanic drum), tdp (tympanic depression), tm (tympanum), tr (transverse ridge).
éo Léger et al. / ZooKeys 907: 1–99 (2020)
78
Figures 87–90. Female genitalia features of Hoploscopa. 87 Hoploscopa albipuncta sp. nov., paratype,
TL609 ♀ 88 Hoploscopa matheae sp. nov., paratype, TL599 ♀ 89 Hoploscopa sepanggi sp. nov., paratype,
TL547 ♀ 90 Hoploscopa cynodonta sp. nov., paratype, TL625 ♀. Abbreviations: aa (anterior apophyses),
at (antrum), cb (corpus bursae), cs (corpus sclerotization), db (ductus bursae), ds (ductus seminalis), pa
(posterior apophyses), pp (papillae anales), segm. VIII (segment VIII), t (thorn).
Twenty-six new species of Hoploscopa 79
Figures 91–94. Female genitalia features of Hoploscopa. 91 Hoploscopa parvimacula sp. nov., paratype,
TL551 ♀ 92 Hoploscopa kinabaluensis sp. nov., TL674 ♀ 93 Hoploscopa luteomacula, paratype, GU prep.
Nuss 743 ♀ 94 Hoploscopa obliqua, paratype, TL656 ♀.
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80
Figures 95–98. Female genitalia features of Hoploscopa. 95 Hoploscopa marijoweissae sp. nov., paratype,
TL709 ♀ 96 Hoploscopa gombongi sp. nov., paratype, TL654 ♀ 97 Hoploscopa pangrangoensis sp. nov.,
paratype, TL627 ♀ 98 Hoploscopa ypsilon sp. nov., paratype, TL620 ♀.
Twenty-six new species of Hoploscopa 81
Figures 99–102. Female genitalia features of Hoploscopa. 99 Hoploscopa danaoensis sp. nov., paratype,
TL618 ♀ 100 Hoploscopa isarogensis sp. nov., holotype, TL514 ♀ 101 Hoploscopa tonsepi sp. nov., para-
type, TL658 ♀ 102 Hoploscopa aurantiacalis, lectotype, Pyralidae Brit. Mus. Slide N° BMNH20246 ♀.
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82
Figures 103–106. Female genitalia features of Hoploscopa. 103 Hoploscopa brunnealis, lectotype, Pyrali-
dae Brit. Mus. Slide N° BMNH20247 ♀ 104 Hoploscopa ocellata, holotype, Pyralidae Brit. Mus. Slide N°
BMNH20258 ♀ 105 Hoploscopa quadripuncta, TL713 ♀ 106 Hoploscopa semifascia, holotype, Pyralidae
Brit. Mus. Slide N° BMNH20253 ♀.
Twenty-six new species of Hoploscopa 83
Figures 107–110. Female genitalia features of Hoploscopa. 107 Hoploscopa astrapias, TL23 ♀ 108Hop-
loscopa anamesa, TL718 ♀ 109 Hoploscopa subvariegata, lectotype, Pyralidae Brit. Mus. Slide N°
BMNH20254 ♀ 110 Hoploscopa triangulifera, holotype, Pyralidae Brit. Mus. Slide N° BMNH20256 ♀.
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84
Figures 111–114. Female genitalia features of Hoploscopa. 111 Hoploscopa anacantha sp. nov., paratype,
TL732 ♀ 112 Hoploscopa kelama sp. nov., paratype, TL665 ♀ 113 Hoploscopa ignitamaculae sp. nov.,
paratype, TL364 ♀ 114 Hoploscopa albomaculata sp. nov., paratype, TL529 ♀.
Twenty-six new species of Hoploscopa 85
Figures 115–118. Female genitalia features of Hoploscopa. 115 Hoploscopa sumatrensis sp. nov., para-
type, TL530 ♀ 116 Hoploscopa mallyi sp. nov., paratype, TL513 ♀ 117 Hoploscopa gracilis sp. nov.,
paratype, TL527 ♀ 118 Hoploscopa agtuuganonensis sp. nov., holotype, TL616 ♀.
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86
Figures 119–122. Female genitalia features of Hoploscopa. 119 Hoploscopa boleta sp. nov., paratype,
TL448 ♀ 120 Hoploscopa pseudometacrossa sp. nov., paratype, TL435 ♀ 121 Hoploscopa metacrossa, para-
type, TL446 ♀ 122 Hoploscopa jubata sp. nov., paratype, TL445 ♀.
Twenty-six new species of Hoploscopa 87
Figure 123. RaxML analysis of the COI barcode dataset partitionned after codon positions. Node sup-
port 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.
éo Léger et al. / ZooKeys 907: 1–99 (2020)
88
Discussion
Our iterative taxonomic revision resulted in the discovery and description of 26 new
species as well as the redescription of 15 species. irty of 38 morphospecies were
concordant with species delimitation based on the COI barcode. Distinct genetic
divergence (i.e., more than 2%) among populations of dierent islands, with minor
morphological dierences in wing pattern or male genitalia, suggests reproductive iso-
lation through allopatric eects. 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 Penin-
sula 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 dierent Philippines islands display
distinct genetic dierentiation suggesting allopatric dierentiation. Cryptic diversity
among populations of dierent 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 dierences in male genitalia in each case. is suggests potential
east-west dierentiation 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 intraspecic and interspecic variation (Bergsten et al. 2012, Talavera et al. 2013).
Two cases of high genetic divergence found in sympatric populations deserve
closer attention. e rst case involves H. sumatrensis sp. nov. from north Sumatra,
where two mitochondrial lineages dier 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 dierences in
either case. Inclusion of nuclear markers is needed in order to test if these mitochon-
drial lineages reect dierent 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).
e investigated material more than doubled the number of Hoploscopa species.
With 41 species now described, we still have not reached the saturation phase in spe-
cies discovery for this group. For example, morphology of the female genitalia and
molecular data support specimens MTD8238 and MTD8243 as dierent 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 unde-
scribed 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. e presumed
Twenty-six new species of Hoploscopa 89
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 altitu-
dinal 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. nautico-
rum). 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 diversication of Hoploscopa. Isolation through
elevation promotes species diversity and endemism (Steinbauer et al. 2016). e 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 mor-
phological 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 rep-
resents 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 spe-
cialists 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 ve 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. e Hoploscopa larva described by Mally et al. (2017)
and collected during a eld trip on Mount Kinabalu did not match any of the eight spe-
cies 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.
Acknowledgements
We are grateful to Ulf Buchsbaum (ZSM), David Lees (NHMUK), Wolfram Mey
(MFNB) and Scott Miller (USNM) for the specimens loaned for investigation. TL is
éo Léger et al. / ZooKeys 907: 1–99 (2020)
90
grateful to David Lees and Geo Martin for the assistance provided during his stay
at the NHMUK. For assistance provided on the 2015 eld work on Mount Kinabalu
(Malaysia, Sabah), TL would like to thank Stephen Sutton, Richard Mally, Ching Y.
Chung (Sabah Biodiversity Center), and the sta of the Sabah Parks. Manuela Bartel
(MTD) performed PCR and sample preparation for several specimens. We thank the
SGN-SNSD-Mol-Lab for allowing access to their facilities for the molecular work.
We are very grateful to Richard Mally and Nathalie Yonow for carefully reviewing our
manuscript. Material for Hoploscopa gombongi sp. nov., H. obliqua sp. nov., and H.
tonsepi sp. nov. originates from the Binatang Research Center program led by Vojtěch
Novotný (Institute of Entomology, Czech Academy of Sciences, České Budějovice,
Czech Republic), George Weiblen (Bell Museum of Natural History, University of
Minnesota, USA), Yves Basset (Smithsonian Tropical Research Institute, Panama),
and Scott Miller (National Museum of Natural History, Smithsonian Institution,
Washington, DC, USA), and was supported by the US National Science Founda-
tion (grants DEB-0515678, 0841885 and others) and the Czech Science Foundation
(grants 206/09/0115, 14-04258S and others). DNA barcodes for these specimens were
provided by Paul Hebert (Department of Integrative Biology, University of Guelph,
Ontario, Canada) through a grant from Genome Canada and the Ontario Genomics
Institute in support of the iBOL project. We thank Vojtěch Novotný, Gibson Mayiah,
Stephen Sutton, and Francesca Vegliante for suggestions of names for some of the
newly described species.
References
Bergsten J, Bilton DT, Fujisawa T, Elliott M, Monaghan MT, Balke M, Hendrich L, Geijer J,
Herrmann J, Foster GN (2012) e eect of geographical scale of sampling on DNA bar-
coding. Systematic biology 61: 851–869. https://doi.org/10.1093/sysbio/sys037
Cong Q, Shen J, Borek D, Robbins RK, Opler PA, Otwinowski Z, Grishin NV (2017) When
COI barcodes deceive: complete genomes reveal introgression in hairstreaks. Proceed-
ings of the Royal Society B: Biological Sciences 284: 20161735. https://doi.org/10.1098/
rspb.2016.1735
Dincă V, Montagud S, Talavera G, Hernández-Roldán J, Munguira ML, García-Barros E,
Hebert PDN, Vila R (2015) DNA barcode reference library for Iberian butteries ena-
bles a continental-scale preview of potential cryptic diversity. Scientic Reports 5: 12395.
https://doi.org/10.1038/srep12395
Dumas P, Barbut J, Le Ru B, Silvain J-F, Clamens A-L, d’Alençon E, Kergoat GJ (2015) Phylo-
genetic Molecular Species Delimitations Unravel Potential New Species in the Pest Genus
Spodoptera Guenée, 1852 (Lepidoptera, Noctuidae). PLOS ONE 10: 1–20. https://doi.
org/10.1371/journal.pone.0122407
Ebihara A, Kuo L-Y (2012) East and Southeast Asian pteridophyte ora and DNA barcoding.
In: e Biodiversity Observation Network in the Asia-Pacic Region. Springer, Tokyo,
321–327. https://doi.org/10.1007/978-4-431-54032-8_23
Twenty-six new species of Hoploscopa 91
Ezard T, Fujisawa T, Barraclough TG (2009) Splits: species’ limits by threshold statistics. R
package version 1.0-14/ r31. http://R-Forge.R-project.org/projects/splits/
Feldhaar H, Gadau J, Fiala B (2010) Speciation in obligately plant-associated Crematogaster
Ants: host distribution rather than adaption towards specic hosts drives the process.
In: Glaubrecht M (Ed.) Evolution in Action. Springer, Berlin, Heidelberg. https://doi.
org/10.1007/978-3-642-12425-9_10
Francis CM, Borisenko AV, Ivanova NV, Eger JL, Lim BK, Guillén-Servent A, Kruskop SV,
Mackie I, Hebert PDN (2010) e role of DNA barcodes in understanding and conserva-
tion of mammal diversity in southeast Asia. PloS ONE 5: e12575–e12575. https://doi.
org/10.1371/journal.pone.0012575
Fujisawa T, Barraclough TG (2013) Delimiting species using single-locus data and the General-
ized Mixed Yule Coalescent approach: a revised method and evaluation on simulated data
sets. Systematic biology 62: 707–724. https://doi.org/10.1093/sysbio/syt033
Funk DJ, Omland KE (2003) Species-level paraphyly and polyphyly: frequency, causes, and conse-
quences, with insights from animal mitochondrial DNA. Annual Review of Ecology, Evolution,
and Systematics 34: 397–423. https://doi.org/10.1146/annurev.ecolsys.34.011802.132421
Gaskin DE (1985) Morphology and Reclassication of the Australasian, Melanesian and Poly-
nesian Glaucocharis Meyrick (Lepidoptera: Crambinae: Diptychophorini). Australian Jour-
nal of Zoology, Supplementary Series 33: 1–75. https://doi.org/10.1071/AJZS115
Hajibabaei M, Smith MA, Janzen DH, Rodriguez JJ, Whiteld JB, Hebert PD (2006) A mini-
malist barcode can identify a specimen whose DNA is degraded. Molecular Ecology Notes
6: 959–964. https://doi.org/10.1111/j.1471-8286.2006.01470.x
Harrison RG, Larson EL (2014) Hybridization, introgression, and the nature of species bound-
aries. Journal of Heredity 105: 795–809. https://doi.org/10.1093/jhered/esu033
Hausmann A, Miller SE, Holloway JD, Dewaard JR, Pollock D, Prosser SW, Hebert PD (2016)
Calibrating the taxonomy of a megadiverse insect family: 3000 DNA barcodes from ge-
ometrid type specimens (Lepidoptera, Geometridae). Genome 59: 671–684. https://doi.
org/10.1139/gen-2015-0197
Hebert PD, Zakharov EV, Prosser SW, Sones JE, McKeown JT, Mantle B, La Salle J (2013) A
DNA “Barcode Blitz”: Rapid digitization and sequencing of a natural history collection.
PLoS One 8: e68535. https://doi.org/10.1371/journal.pone.0068535
Heppner JB (1998) Classication of Lepidoptera. Part. 1 Introduction. Holarctic Lepidoptera
5, Suppl. 1: iv + 148 + 6 pp.
Hundsdoerfer AK, Päckert M, Kehlmaier C, Strutzenberger P, Kitching IJ (2017) Museum
archives revisited: Central Asiatic hawkmoths reveal exceptionally high late Pliocene spe-
cies diversication (Lepidoptera, Sphingidae). Zoologica Scripta 46: 552–570. https://doi.
org/10.1111/zsc.12235
Hurst GD, Jiggins FM (2005) Problems with mitochondrial DNA as a marker in population, phy-
logeographic and phylogenetic studies: the eects of inherited symbionts. Proceedings of the
Royal Society B: Biological Sciences 272: 1525–1534. https://doi.org/10.1098/rspb.2005.3056
Joseph L, Slikas B, Alpers D, Schodde R (2001) Molecular systematics and phylogeography of
New Guinean logrunners (Orthonychidae). Emu 101: 273–280. https://doi.org/10.1071/
MU01008
éo Léger et al. / ZooKeys 907: 1–99 (2020)
92
Kekkonen M, Hebert PD (2014) DNA barcode-based delineation of putative species: e-
cient start for taxonomic workows. Molecular ecology resources 14: 706–715. https://
doi.org/10.1111/1755-0998.12233
Knölke S, Erlacher S, Hausmann A, Miller MA, Segerer AH (2004) A procedure for combined
genitalia dissection and DNA extraction in Lepidoptera. Insect Systematics & Evolution
35: 401–409. https://doi.org/10.1163/187631204788912463
Kristensen NP (2003) 4. Skeleton and muscles: adults. Handbook of Zoology, Vol. IV, Part 36,
Lepidoptera, Moths and Butteries, Vol. 2, Morphology, Physiology, and Development.
De Gruyter, Berlin, 39–131.
Landry B (1995) A phylogenetic analysis of the major lineages of the Crambinae and of the
genera of Crambini of North America (Lepidoptera:Pyralidae). Memoirs on Entomology
International. Associated Publishers, Gainesville, 242 pp.
Lanfear R, Calcott B, Ho SYW, Guindon S (2012) Partitionnder: combined selection of parti-
tioning schemes and substitution models for phylogenetic analyses. Molecular Biology and
Evolution 29: 1695–1701. https://doi.org/10.1093/molbev/mss020
Lanfear R, Frandsen PB, Wright AM, Senfeld T, Calcott B (2017) PartitionFinder 2: New
methods for selecting partitioned models of evolution for molecular and morphologi-
cal phylogenetic analyses. Molecular Biology and Evolution 34: 772–773. https://doi.
org/10.1093/molbev/msw260
Leraut P (1980) Liste systématique et synonymique des Lépidoptères de France, Belgique et
Corse. Alexanor and Bulletin de la Société entomologique de France Supplement: 1–334.
Léger T, Landry B, Nuss M (2019) Phylogeny, character evolution and tribal classication in
Crambinae and Scopariinae (Lepidoptera, Crambidae). Systematic Entomology 44: 757–
776. https://doi.org/10.1111/syen.12353
Lohman DJ, de Bruyn M, Page T, von Rintelen K, Hall R, Ng PK, Shih H-T, Carvalho GR,
von Rintelen T (2011) Biogeography of the Indo-Australian archipelago. Annual Review
of Ecology, Evolution, and Systematics 42: 205–226. https://doi.org/10.1146/annurev-
ecolsys-102710-145001
Lohman DJ, Ingram KK, Prawiradilaga DM, Winker K, Sheldon FH, Moyle RG, Ng PK, Ong
PS, Wang LK, Braile TM (2010) Cryptic genetic diversity in “widespread” Southeast Asian
bird species suggests that Philippine avian endemism is gravely underestimated. Biological
Conservation 143: 1885–1890. https://doi.org/10.1016/j.biocon.2010.04.042
Maddison WP, Maddison DR (2017) Mesquite: a modular system for evolutionary analysis –
Version 3.31. http://www.mesquiteproject.org
Mally R, Huemer P, Nuss M (2018) Deep intraspecic DNA barcode splits and hybridisation
in the Udea alpinalis group (Insecta, Lepidoptera, Crambidae) – an integrative revision.
ZooKeys 746: 51–90. https://doi.org/10.3897/zookeys.746.22020
Mally R, Léger T, Vairappan CS, Sutton S, Nuss M (2017) Discovery of another fern-feeding
group of moths: the larvae of Hoploscopini (Insecta: Lepidoptera: Pyraloidea) from Bor-
neo. Raes Bulletin of Zoology 65: 100–108.
Merckx VSFT, Hendriks KP, Beentjes KK, Mennes CB, Becking LE, Peijnenburg KTCA,
Afendy A, Arumugam N, de Boer H, Biun A, Buang MM, Chen P-P, Chung AYC, Dow
R, Feijen FAA, Feijen H, Soest CF, Geml J, Geurts R, Gravendeel B, Hovenkamp P, Im-
bun P, Ipor I, Janssens SB, Jocqué M, Kappes H, Khoo E, Koomen P, Lens F, Majapun
Twenty-six new species of Hoploscopa 93
RJ, Morgado LN, Neupane S, Nieser N, Pereira JT, Rahman H, Sabran S, Sawang A,
Schwallier RM, Shim P-S, Smit H, Sol N, Spait M, Stech M, Stokvis F, Sugau JB, Sulei-
man M, Sumail S, omas DC, van Tol J, Tuh FYY, Yahya BE, Nais J, Repin R, Lakim M,
Schilthuizen M (2015) Evolution of endemism on a young tropical mountain. Nature 524:
347. https://doi.org/10.1038/nature14949
Miller MA, Pfeier W, Schwartz T (2010) Creating the CIPRES Science Gateway for inference
of large phylogenetic trees. Proceedings of the Gateway Computing Environments Work-
shop (GCE), New Orleans, 1–8. https://doi.org/10.1109/GCE.2010.5676129
Miller SE, Rosati ME, Gewa B, Novotny V, Weiblen GD, Hebert PD (2015) DNA Barcodes
of Lepidoptera Reared from Yawan, Papua New Guinea. Proceedings of the Entomologi-
cal Society of Washington 117: 247–250. https://doi.org/10.4289/0013-8797.117.2.247
Minet J (1982) Les Pyraloidea et leurs principales divisions systématiques (Lep. Ditrysia). Bul-
letin de la Société entomologique de France 86: 262–280.
Mitchell A (2015) Collecting in collections: a PCR strategy and primer set for DNA barcod-
ing of decades-old dried museum specimens. Molecular ecology resources 15: 1102–1111.
https://doi.org/10.1111/1755-0998.12380
Müller J, Müller K, Neinhuis C, Quandt D (2005) PhyDE-Phylogenetic Data Editor.
Munroe E, Solis MA (1998) e Pyraloidea. Lepidoptera, Moths and Butteries. Vol 1: Evolu-
tion, Systematics, and Biogeography. Handbook of Zoology, Berlin, 233–256.
Myers N, Mittermeier RA, Mittermeier CG, Da Fonseca GA, Kent J (2000) Biodiversity hot-
spots for conservation priorities. Nature 403: 853. https://doi.org/10.1038/35002501
Nakahara S, Lamas G, Tyler S, Marín MA, Huertas B, Willmott KR, Mielke OHH, Espe-
land M (2019) A revision of the new genus Amiga Nakahara, Willmott & Espeland, gen.
n., described for Papilio arnaca Fabricius, 1776 (Lepidoptera, Nymphalidae, Satyrinae).
ZooKeys 821: 85–152. https://doi.org/10.3897/zookeys.821.31782
Nuss M (1998) e Scopariinae and Heliothelinae stat. rev. (Lepidoptera: Pyraloidea, Crambidae)
of the Oriental Region – a revisional synopsis with descriptions of new species from the Phil-
ippines and Sumatra. Nachrichten des entomologischen Vereins Apollo suppl. 17: 475–528.
Pons J, Barraclough TG, Gomez-Zurita J, Cardoso A, Duran DP, Hazell S, Kamoun S, Sumlin WD,
Vogler AP (2006) Sequence-based species delimitation for the DNA taxonomy of undescribed
insects. Systematic biology 55: 595–609. https://doi.org/10.1080/10635150600852011
Prosser SW, deWaard JR, Miller SE, Hebert PD (2016) DNA barcodes from century-old type
specimens using next-generation sequencing. Molecular Ecology Resources 16: 487–497.
https://doi.org/10.1111/1755-0998.12474
Puillandre N, Lambert A, Brouillet S, Achaz G (2012) ABGD, Automatic Barcode Gap Dis-
covery for primary species delimitation. Molecular Ecology 21: 1864–1877. https://doi.
org/10.1111/j.1365-294X.2011.05239.x
Quek S-P, Davies SJ, Ashton PS, Itino T, Pierce NE (2007) e geography of diversication
in mutualistic ants: a gene’seye view into the Neogene history of Sundaland rain forests.
Molecular Ecology 16: 2045–2062. https://doi.org/10.1111/j.1365-294X.2007.03294.x
Rambaut A, Drummond AJ, Xie D, Baele G, Suchard MA (2018) Tracer. http://tree.bio.ed.ac.
uk/software/tracer/
Robinson GS (1976) e preparation of slides of Lepidoptera genitalia with special reference to
the Microlepidoptera. Entomologist’s Gazette 27: 127–132.
éo Léger et al. / ZooKeys 907: 1–99 (2020)
94
Robinson GS, Tuck KR, Shaer M (1994) A eld guide to the smaller moths of South-East
Asia. Natural History Museum, London, 308 pp.
RodríguezCastañeda G, Dyer LA, Brehm G, Connahs H, Forkner RE, Walla TR (2010)
Tropical forests are not at: how mountains aect herbivore diversity. Ecology letters 13:
1348–1357. https://doi.org/10.1111/j.1461-0248.2010.01525.x
Schlick-Steiner BC, Steiner FM, Seifert B, Stauer C, Christian E, Crozier RH (2010) Integra-
tive taxonomy: a multisource approach to exploring biodiversity. Annual review of ento-
mology 55: 421–438. https://doi.org/10.1146/annurev-ento-112408-085432
Schulze CH (2000) Auswirkungen anthropogener Störungen auf die Diversität von Herbi-
voren: Analyse von Nachtfalterzönosen entlang von Habitatgradienten in Ost-Malaysia.
Universität Bayreuth.
Schulze CH, Fiedler K (2003) Vertical and temporal diversity of a species-rich moth taxon in
Borneo. In: Basset Y, et al. (Eds) Arthropods of Tropical Forests: Spatio-temporal Dynam-
ics and Resource Use in the Canopy. Cambridge University Press, 69–85.
Song H, Buhay JE, Whiting MF, Crandall KA (2008) Many species in one: DNA barcoding
overestimates the number of species when nuclear mitochondrial pseudogenes are coam-
plied. Proceedings of the national academy of sciences 105: 13486–13491. https://doi.
org/10.1073/pnas.0803076105
Stamatakis A (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses
with thousands of taxa and mixed models. Bioinformatics 22: 2688–2690. https://doi.
org/10.1093/bioinformatics/btl446
Steinbauer MJ, Field R, Grytnes J-A, Trigas P, AhPeng C, Attorre F, Birks HJB, Borges PA,
Cardoso P, Chou C-H (2016) Topographydriven isolation, speciation and a global in-
crease of endemism with elevation. Global Ecology and Biogeography 25: 1097–1107.
https://doi.org/10.1111/geb.12469
Suchard MA, Lemey P, Baele G, Ayres DL, Drummond AJ, Rambaut A (2018) Bayesian phylo-
genetic and phylodynamic data integration using BEAST 1.10. Virus Evolution 4: vey016.
https://doi.org/10.1093/ve/vey016
Sutton S, Barlow H, Whitaker T (2015) An exciting new book. Pyraloid Planet 9: 2–3.
Talavera G, Dincă V, Vila R (2013) Factors aecting species delimitations with the GMYC
model: insights from a buttery survey. Methods in Ecology and Evolution 4: 1101–1110.
https://doi.org/10.1111/2041-210X.12107
Wahlberg N, Wheat CW (2008) Genomic outposts serve the phylogenomic pioneers: design-
ing novel nuclear markers for genomic DNA extractions of Lepidoptera. Systematic Biol-
ogy 57: 231–242. https://doi.org/10.1080/10635150802033006
Weintraub JD, Lawton JH, Scoble MJ (1995) Lithinine moths on ferns: a phylogenetic
study of insect-plant interactions. Biological Journal of the Linnean Society 55: 239–
250. https://doi.org/10.1111/j.1095-8312.1995.tb01062.x
Yeates DK, Seago A, Nelson L, Cameron SL, Joseph LEO, Trueman JW (2011) Integrative
taxonomy, or iterative taxonomy? Systematic Entomology 36: 209–217. https://doi.
org/10.1111/j.1365-3113.2010.00558.x
Zakharov EV, Lobo NF, Nowak C, Hellmann JJ (2009) Introgression as a likely cause of
mtDNA paraphyly in two allopatric skippers (Lepidoptera: Hesperiidae). Heredity 102:
590pp. https://doi.org/10.1038/hdy.2009.26
Twenty-six new species of Hoploscopa 95
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.
Species Latitude Longitude Geographical
location
Country Detailed locality Remarks
H. agtuuganonensis 7.78 126.21 Mindanao Philippines Mount Agtuuganon
H. albipuncta 6.042 116.596Borneo Malaysia Kopogon
6.045 116.595 Borneo Malaysia Mesilau Nature Resort
6.028 116.5483 Borneo Malaysia Timpohon Gate, 300m after Ligawu trail start
6.0277 116.5497 Borneo Malaysia Timpohon Gate, 700m after Ligawu trail start
H. albomaculata 0.76 124.46 North Sulawesi Indonesia Lake Danau Mooat
H. anacantha 0.56 123.68 North Sulawesi Indonesia Dumoga-Bone NP [Bogani Nani Wartabone NP]
0.76 124.46 North Sulawesi Indonesia Lake Danau Mooat
0.45 123.93 North Sulawesi Indonesia Mount Mogogonipa
H. anamesa -20.2 169.81 Aneityum Vanuatu Aneityum
-20.23 169.76 Aneityum Vanuatu Anelgauhat
-19.5 169.3 Tanna Vanuatu Tanna
-17.56 177.96 Fiji Fiji Nandarivatu
-17.82 178.32 Fiji Fiji Vunîdawa
H. aurantiacalis -7.175 107.57 Pengaleng Indonesia Pengaleng is place could not be assessed with
condence. We refer here to West Java
H. boleta -7.35 146.68 Papua Papua New Guinea Mount Kaindi
H. brunnealis -7.175 107.57 Pengaleng Indonesia Pengaleng is place could not be assessed with
condence. We refer here to West Java
H. cynodonta 4.555 115.154 Borneo Brunei Ulu Temburong
6.009 116.537 Borneo Malaysia Kiau View-Pandanus Trail
5.48 116.56 Borneo Malaysia Mount Monkobo
4.467 117.917 Borneo Malaysia Tawau Hills HQ
6.0277 116.5497 Borneo Malaysia Timpohon Gate, 700m after Ligawu trail start
H. danaoensis 11.07 124.69 Leyte Philippines Lake Danao
H. diusa -9.5 150.66 Papua Papua New Guinea Fergusson Island
H. gombongi -6.2 146.1 Papua Papua New Guinea Yawan village
H. gracilis 2.77 98.53 Sumatra Indonesia Dairi
3.24 98.51 Sumatra Indonesia Mount Sibayak
2.755 99 Sumatra Indonesia Road from Pematangsiantar to Prapat original label: “Holzweg 2, 25km SSW-
Pematangsiantar, Straße nach Prapat”
éo Léger et al. / ZooKeys 907: 1–99 (2020)
96
Species Latitude Longitude Geographical
location
Country Detailed locality Remarks
H. gracilis 1.617 99.267 Sumatra Indonesia Sipirok
H. ignitamaculae 0.56 123.68 North Sulawesi Indonesia Dumoga-Bone NP [Bogani Nani Wartabone NP]
0.76 124.46 North Sulawesi Indonesia Lake Danau Mooat
0.45 123.93 North Sulawesi Indonesia Mount Mogogonipa
1.51 125.18 North Sulawesi Indonesia Tangkoko Batuangus Nature Reserve
H. isarogensis 13.66 123.33 Luzon Philippines Mount Isarog
H. jubata -7.22 146.61 Papua Papua New Guinea Mount Susu NP
-7 146 Papua Papua New Guinea Wau Ecology Institute
H. kelama 0.56 123.68 North Sulawesi Indonesia Dumoga-Bone NP [Bogani Nani Wartabone NP]
0.76 124.46 North Sulawesi Indonesia Lake Danau Mooat
H. kinabaluensis 6.009 116.537 Borneo Malaysia Kiau View-Pandanus Trail
6.0116 116.5344 Borneo Malaysia Kinabalu Mountain Lodge
6.028 116.5483 Borneo Malaysia Timpohon Gate, 300m after Ligawu trail start
6.0277 116.5497 Borneo Malaysia Timpohon Gate, 700m after Ligawu trail start
H. luteomacula -0.4 100.35 Sumatra Indonesia Mount Singgalang
2.755 99 Sumatra Indonesia Road from Pematangsiantar to Prapat original label: “Holzweg 2, 25km SSW-
Pematangsiantar, Straße nach Prapat”
H. mallyi 6.0394 116.598 Borneo Malaysia Mesilau logging site
6.045 116.595 Borneo Malaysia Mesilau Nature Resort
6.009 16.5388 Borneo Malaysia Pandanus trail, 250m after start
6.01 116.54 Borneo Malaysia Sabah, Kundasang, Kinabalu NP Headquarters
6.028 116.5483 Borneo Malaysia Timpohon Gate, 300m after Ligawu trail start
H. marijoweissae -4.4 139.7 Papua Indonesia Mount Goliath [Mount Yamin]
H. matheae 4.33 115.29 Borneo Brunei Bukit Retak
6.0116 116.5344 Borneo Malaysia Kinabalu Mountain Lodge
4.467 117.917 Borneo Malaysia Tawau Hills HQ
H. metacrossa -2.91 132.3 Papua Indonesia Fak-Fak
-7.22 146.61 Papua Papua New Guinea Mount Susu NP
-7 146 Papua Papua New Guinea Wau Ecology Institute
H. nauticorum -13.9 -171.75 Upolu Samoa Malololelei
-13.92 171.75 Upolu Samoa Tiavi
H. niveofascia -7.22 146.61 Papua Papua New Guinea Mount Susu NP
H. obliqua -4.48 137.34 Papua Indonesia Utakwa [Oetakwa] river
-5.25 145.283 Papua Papua New Guinea Wanang village
-0.61 127.3 Moluccas Indonesia Batchian [Bacan islands]
Twenty-six new species of Hoploscopa 97
Species Latitude Longitude Geographical
location
Country Detailed locality Remarks
H. pangrangoensis -6.77 106.96 Java Indonesia Mount Pangrango
H. parvimacula 6.0116 116.5344 Borneo Malaysia Kinabalu Mountain Lodge
6.0394 116.598 Borneo Malaysia Mesilau logging site
6.045 116.595 Borneo Malaysia Mesilau Nature Resort
6.028 116.5483 Borneo Malaysia Timpohon Gate, 300m after Ligawu trail start
H. persimilis -4.48 137.34 Papua Indonesia Utakwa [Oetakwa] river
H. pseudometacrossa -5.58 144.07 Papua Papua New Guinea Mount Hagen
H. quadripuncta -3.08 129.53 Moluccas Indonesia Mount Kobipoto
-4.48 137.34 Papua Indonesia Utakwa [Oetakwa] river
-9 148.37 Papua Papua New Guinea Hydrographers Range
H. semifascia -2.91 132.3 Papua Indonesia Fak-Fak
-6.58 142.83 Papua Papua New Guinea Bosavi (Mount)
-7 146 Papua Papua New Guinea Wau Ecology Institute
H. sepanggi 6.0394 116.598 Borneo Malaysia Mesilau logging site
6.045 116.595 Borneo Malaysia Mesilau Nature Resort
6.028 116.5483 Borneo Malaysia Timpohon Gate, 300m after Ligawu trail start
H. subvariegata -8.8 146.5 Papua Papua New Guinea Angabunga river
-7.35 146.68 Papua Papua New Guinea Mount Kaindi
H. sumatrensis 2.755 99 Sumatra Indonesia Road from Pematangsiantar to Prapat original label: “Holzweg 2, 25km SSW-
Pematangsiantar, Straße nach Prapat”
H. titika 2.755 99 Sumatra Indonesia Road from Pematangsiantar to Prapat original label: “Holzweg 2, 25km SSW-
Pematangsiantar, Straße nach Prapat”
H. tonsepi -6.2 146.1 Papua Papua New Guinea Yawan village
H. triangulifera -4.63 145.97 Papua Papua New Guinea Dampier Island [Karkar]
-9.5 150.66 Papua Papua New Guinea Fergusson Island
-8.45 147.81 Papua Papua New Guinea Mambare River
H. ypsilon 17.04 121.1 Luzon Philippines Barlig (Chatol)
16.97 121.03 Luzon Philippines Mount Polis
16.16 120.94 Luzon Philippines Santa Fe
éo Léger et al. / ZooKeys 907: 1–99 (2020)
98
Supplementary material 1
Figure S1. Scheme summarising the approach followed for amplication of the
COI barcode from old material
Authors: éo Léger, Christian Kehlmaier, Charles S. Vairappan, Matthias Nuss
Data type: molecular data
Copyright notice: is dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Link: https://doi.org/10.3897/zookeys.907.36563.suppl1
Supplementary material 2
Table S1. Primers sequences with characteristics and chemical properties
Authors: éo Léger, Christian Kehlmaier, Charles S. Vairappan, Matthias Nuss
Data type: molecular data
Copyright notice: is dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Link: https://doi.org/10.3897/zookeys.907.36563.suppl2
Supplementary material 3
Table S2. List of DNA samples of Hoploscopa moths and allies used in this study
Authors: éo Léger, Christian Kehlmaier, Charles S. Vairappan, Matthias Nuss
Data type: molecular data
Copyright notice: is dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Link: https://doi.org/10.3897/zookeys.907.36563.suppl3
Twenty-six new species of Hoploscopa 99
Supplementary material 4
Table S3. K2P-distances calculated on the ABGD online server (https://bioinfo.
mnhn.fr/abi/public/abgd/abgdweb.html) using the Kimura 2-parameters algorithm
Authors: éo Léger, Christian Kehlmaier, Charles S. Vairappan, Matthias Nuss
Data type: molecular data
Copyright notice: is dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Link: https://doi.org/10.3897/zookeys.907.36563.suppl4
Available via license: CC BY
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