Cuprininae is a small lineage within the moth family Stathmopodidae, characterized by tiny adults and larvae that feed exclusively on fern spores. This feeding habit is rare not only among butterflies and moths but among insects generally. A recent evolutionary analysis suggested that fern-spore feeding probably evolved once within Stathmopodidae and has been reported in Cuprininae and two other genera, Agrioscelis and Erineda. Cuprininae is not restricted to a single fern genus or family: larvae have been recorded from at least 11 fern families, indicating a broader but still specialized form of host use described as oligophagy.

Before this study, three Cuprininae species had been reliably recorded in Korea: Calicotis chrysoptera, C. latebrifica and Cuprina fuscella. The researchers examined material representing C. latebrifica and C. fuscella, the two previously recorded species included in their morphological and molecular comparisons, alongside the two newly described species and three new country records. C. chrysoptera was not examined in the study. Pachyrhabda benearena, recently described from Korea, was also excluded from the confirmed Cuprininae list because its taxonomic placement requires further examination.

The newly described Pediformis astathmeta is the third species recognized in its genus and can be separated from its two congeners by details of the male genitalia. Its cucullus, the foot-shaped apical portion of the valva, narrows gradually toward a rounded apex; the cucullus of P. cuneiformis is rectangular, whereas that of P. fushana ends in a pointed projection. The aedeagus of P. astathmeta is much narrower and longer than those of the other two species. Females have two signa, and the signum nearest the ductus bursae is smaller and rounder than in P. cuneiformis. The moth has a forewing span of 8.0–10.0 mm across 31 specimens, with pale wings and indistinct grey markings.

Field and laboratory observations also documented the biology of P. astathmeta. Its larvae were found feeding beneath fronds of the fern Arachniodes sporadosora, where they made tunnel-like shelters and silky galleries before forming cocoons. Adults flew in May, while larvae were observed only in June, a seasonal pattern suggesting that the species is probably univoltine, or has one generation per year. The moth shared a habitat with C. fuscella, and both occurred on A. sporadosora. However, P. astathmeta larvae were found only on that fern, whereas C. fuscella larvae consumed A. sporadosora and three additional fern species in the family Dryopteridaceae.

The second new species, Calicotis atricosta, is distinguished by a whitish forewing with a black basal section of the costa and a broad, inwardly oblique black band before the wing midpoint. Unlike many related species, it lacks dark blotches or streaks in the distal part of the forewing. Its male genitalia supply further diagnostic traits, including an oval cucullus, an oval anellar lobe and a rounded juxta. The aedeagus bears a narrow apical spatulate projection about one-third of its length. The name atricosta refers to the black basal costa. Only two male specimens were available, so no female genitalia, wing-venation slide, host plant or life-history information could be established for C. atricosta.

The first Korean record of Cuprina porphyrantha is difficult to distinguish from C. fuscella by superficial appearance, because both have dark forewings without distinct markings. Antennal coloration provides one useful difference: the scape of C. porphyrantha is ochreous-yellow dorsally, compared with fuscous to dark grey in C. fuscella, and the apical portion of its antenna is more broadly pale, although that feature can sometimes be indistinct. The basal sclerotized portion of the aedeagus is also larger in C. porphyrantha. A Korean larva was found on Arachniodes aristata, while an adult was collected in August. The species has also been reported from Japan, Russia, Sri Lanka and Taiwan.

The two newly recorded Petala species can likewise be separated through a combination of external and genital characters. Petala aedificatrix has a measured forewing span of 10.0 mm, compared with 6.5–7.5 mm for the 12 examined P. margaritacea specimens, and its forewing markings are less distinct. It also bears black bristles at the apex of the hind tibia rather than the white to pale-grey bristles of P. margaritacea. In the male genitalia, the cucullus of P. aedificatrix curves outward apically and the sacculus is indistinct; P. margaritacea has an uncurved cucullus and a sacculus with a rounded apex. The aedeagus of P. aedificatrix has a distinctive needle-like cornutus with a sharply pointed apex.

To test the classifications, the team extracted DNA from one or two legs of dried adults and from larvae, then analyzed the mitochondrial protein-coding gene cytochrome c oxidase subunit I together with four nuclear genes: EF-1α, carbamoyl-phosphate synthase domain, histone H3 and wingless. The concatenated alignment was 2,578 base pairs long and included sequences representing 37 additional Cuprininae species and the stathmopodid outgroup Stathmopoda auriferella. Maximum-likelihood and Bayesian analyses produced largely congruent trees, placing all seven Korean taxa in their respective genera. P. astathmeta occurred within a strongly supported Pediformis clade, with an ultrafast bootstrap value of 96 and a Bayesian posterior probability of 1, while C. atricosta joined Calicotis with bootstrap support of 100 and posterior probability of 1.

The molecular results also supported the species-level identities of the three new Korean records. Petala aedificatrix, P. margaritacea and Cuprina porphyrantha grouped with previously identified sequences from the same species in other regions, providing an independent line of evidence for their placement. Confidence is lower for some deeper relationships within Cuprininae. Both analyses placed Calicotis and Cuprina as sister lineages, with Trigonoda outside that pair, but the bootstrap support was moderate and this topology differed from one analysis in an earlier family-level study. The present dataset focused on Cuprininae and verification of Korean taxa, and several nuclear genes were missing for some specimens, so broader sampling is needed to clarify the subfamily’s deeper evolutionary relationships.

Subject of Research: Taxonomic and molecular study of seven Cuprininae moth species collected on Jeju Island, South Korea

Article Title: Two new species and three new records of Cuprininae ( Lepidoptera , Stathmopodidae ) from Jeju Island, Korea, with molecular evidence for their placement

Article References: Lee, G.-E., & Terada, T. (2026). Two new species and three new records of Cuprininae (Lepidoptera, Stathmopodidae) from Jeju Island, Korea, with molecular evidence for their placement. ZooKeys, 1287, 253-272. https://doi.org/10.3897/zookeys.1287.194473

Image Credits: AI Generated

DOI: 10.3897/zookeys.1287.194473

Keywords: Cuprininae, Stathmopodidae, moth taxonomy, new species, Jeju Island, phylogenetics, fern-spore feeding, Korean fauna, DNA barcoding, species records, species, three

The name bellatula is a diminutive form of the Latin adjective bellatulus , meaning beautiful. Six Epicroesa species were recognized before this description: E. ambrosia , E. calliteucha , E. chromatorhoea , E. metallifera , E. thiasarcha and E. lylae . Most heliodinid sun moths occur in the New World, whereas Epicroesa is an Old World genus.

Before the new Taiwanese taxon was separated, its members were documented from Australia, New Guinea, Japan and Tahiti. The addition of E. bellatula raises the number of recognized species in the genus to seven and extends its documented distribution to southern Taiwan. Like its Australian relative, E. bellatula has metallic dark-brown forewings, orange distal areas with silver markings and elaborate coremata containing lanceolate scales.

Adults of both species also feed as larvae on the same plant, Ceodes umbellifera . That plant was formerly called Pisonia umbellifera before a reclassification separated the broadly defined Pisonia group into Pisonia , Rockia and Ceodes . The first recorded larval-host association for Epicroesa therefore links an Australian moth and its newly distinguished Taiwanese counterpart through the same member of Nyctaginaceae.

Adult morphology nevertheless revealed a series of consistent differences. The forewing of E. bellatula measures 2.76–3.69 mm, with a mean of 3.39 ± 0.23 mm in a sample of 30, compared with 3.05–3.94 mm and a mean of 3.55 ± 0.17 mm for 30 Australian specimens. In the Taiwanese species, the dark proximal band on the upper forewing bears an orange patch, whereas the Australian species has several narrow orange streaks.

The distal orange area contains three metallic markings in E. bellatula but four in E. thiasarcha . Metallic scaling over the basal half of the forewing and head is yellow with a green tinge in the new species, but green with a blue tinge in the Australian taxon. Behind the compound eye, the Taiwanese moth has creamy-yellow, spatulate scales; the Australian moth has dark-brown, elongate, lanceolate scales.

Differences in wing venation and genitalic structures supplied additional diagnostic characters. The hindwing vein CuA2 is present in E. bellatula but vestigial in E. thiasarcha . In males, the valva of the Taiwanese species forms an elongate, ovate lobe, while the corresponding structure in the Australian species is triangular and axe-shaped.

Female E. bellatula has two V-shaped signa, each armed with prominent spines, whereas the Australian species has an elongate, bifurcate sclerotized band accompanied by a smaller sclerite bearing short cones. These distinctions occur across multiple anatomical systems rather than being confined to a single visible wing mark. The immature stages also separated the two taxa.

Eggs of E. bellatula measured 0.35 ± 0.03 mm long and 0.20 ± 0.01 mm in diameter in a sample of six. They are ovoid and have fine sculpturing on a thin chorion. Mature larvae averaged 5.94 ± 0.39 mm long in 22 individuals.

Both species have a rounded head with a deep V-shaped medial notch, but the head capsule of E. bellatula has a Y-shaped dark-brown marking and thick lateral bands. The Australian larva instead has a pair of prominent, straight dark-brown bands. Their pupae are flattened and tapered in both species, yet differ in proportion: the mean length-to-width ratio at abdominal segment A3 was approximately 2.75 for E. bellatula and 2.58 for E. thiasarcha , based on 30 individuals of each species.

For molecular comparison, the researchers extracted genomic DNA from one adult specimen of each species and amplified a 665-base-pair fragment of the mitochondrial cytochrome c oxidase subunit I gene. The amplified products were purified, sequenced, visually checked and edited, then aligned with MUSCLE. Genetic differentiation was estimated as pairwise distances among ten individuals under the Kimura two-parameter model in MEGA 12.

The resulting COI p-distance between the Australian and Taiwanese samples was 9.75 percent, represented by GenBank accessions PZ166696 and PZ166697. This value was well above the 3 percent divergence often cited as a general species-delimitation criterion for Lepidoptera. The barcode result, together with the morphological and early-stage differences, supported recognition of the Taiwanese population as a separate species rather than an extension of the known range of E. thiasarcha .

Larvae of E. thiasarcha construct a loose silk shelter on the underside of a C. umbellifera leaf, skeletonize the leaf by consuming mesophyll tissue and deposit frass nearby. Pupation occurs beneath the leaf in a loosely constructed cocoon. Eggs of E. bellatula were recorded on leaves, young twigs and flower buds.

Its larvae also make loose silk shelters, after which they bore into flower buds or feed on the undersides of leaves. The pupa is usually attached to the underside of a leaf. Adults and immature stages have been observed at different times of the year in both species, leading the researchers to describe them as apparently multivoltine.

Although C. umbellifera supports both species, their currently known populations are separated by more than 6,000 km: E. thiasarcha is known from Queensland, Australia, while E. bellatula is known only from southern Taiwan. The authors identify two possible explanations: each species may have a broader distribution than current records show, or additional Epicroesa taxa may remain undescribed between the two regions. The authors also caution that the presence or absence of hindwing vein CuA2 should not, by itself, be treated as a reliable diagnostic character for the genus Epicroesa , because previous comparative work indicates that this feature can be lost independently in heliodinid lineages.

Subject of Research: Taxonomic identification of the Taiwanese sun moth Epicroesa bellatula and its larval association with Ceodes umbellifera .

Article Title: Sun moths in the genus Epicroesa ( Lepidoptera , Heliodinidae ) associated with Ceodes umbellifera , with description of a new species from Taiwan

Article References: Hsu, Y.-F. (2026). Sun moths in the genus Epicroesa (Lepidoptera, Heliodinidae) associated with Ceodes umbellifera, with description of a new species from Taiwan. ZooKeys, 1289, 203-214. https://doi.org/10.3897/zookeys.1289.194186

Image Credits: AI Generated

DOI: 10.3897/zookeys.1289.194186

Keywords: new species, Epicroesa bellatula, Epicroesa thiasarcha, sun moths, Heliodinidae, DNA barcoding, Ceodes umbellifera, moth taxonomy, Taiwan, Australia, moths, genus

P. barrassoi was described in 2006 from a single male collected at 2,530 metres in the Maiella Mountains of Abruzzo, Italy. That specimen became the holotype. A female originally associated with the species was subsequently found to be a misidentified and unusually small specimen of Colostygia aqueata. A male collected on Monte Terminillo in 2004, about 100 kilometres from the type locality, was barcoded in work reported in 2019 and showed 8.9% sequence divergence from P. incultaria. That isolated genetic result supported recognition of a separate Central Italian taxon but did not establish its wider distribution. The later study brought the type anatomy, mitochondrial barcodes, nuclear genomic data, adult morphology and ecological records together to reassess populations assigned to the P. incultaria–P. barrassoi complex.

Researchers combined external characters with dissections of male and female genitalia, observations of phenology and vertical distribution, and DNA evidence. They measured anatomy with a calibrated microscope eyepiece and examined prepared genitalia and abdomens. For barcoding, they extracted DNA from one or two legs of dried specimens and analyzed a 658-base-pair region of the mitochondrial cytochrome c oxidase I gene, or COI. Sequence divergences were calculated with the Kimura two-parameter model and analyzed using a neighbor-joining algorithm. High-throughput sequencing was also used for older museum material. In parallel, ddRAD sequencing generated genome-wide single-nucleotide polymorphisms, or SNPs, for maximum-likelihood phylogenetic analysis and population-clustering tests. The authors used these independent evidence streams together because the two moths have no constant and reliable external wing feature that permits separation in the field.

The mitochondrial results divided the tested specimens into two clearly separated groups. The Monte Terminillo specimen produced a 658-base-pair sequence assigned to BOLD bin ACJ5976. Six specimens from Austria, southwestern Bavaria and northern Italy shared that bin and were only 1.1% distant from the Central Italian sequence. Their nearest species was P. incultaria, represented in the comparison by 19 specimens from Germany, Austria and northern Italy in bin AAF5044, with a reported barcode distance of 8.5%. Two specimens attributed to P. incultaria from the vicinity of Gastein in Salzburg were also successfully sequenced using high-throughput methods and fell unambiguously into the P. incultaria bin. Thus, the barcode evidence separated the taxa while also connecting Alpine populations with the Central Italian reference material.

Male genital anatomy strongly supported the DNA assignments. The newly attributed Alpine and Pyrenean populations closely matched the Italian holotype of P. barrassoi. In this species, the valvae are comparatively narrow, with their dorsal and ventral margins approximately parallel, and the costa narrows toward a finely tapered apex. Its aedeagus bears five to nine small cornuti measuring approximately 0.1 to 0.25 millimetres, none expanded at the base. P. incultaria has a much broader valval costa that dilates toward its apex and terminates in a small, beak-shaped sclerotized process bearing a conspicuous tuft of setae. Its aedeagus has five to seven stouter cornuti, with the distalmost generally the largest, curved and dilated at the base. The diagnostic contrasts were assessed in 7 male P. barrassoi and 23 male P. incultaria dissections.

Female genitalia supplied additional but subtler distinctions. In the six P. barrassoi females examined, the apophyses are fine and tapered, while the elongated colliculum is almost parallel-sided for most of its length. Its reported length was 0.37–0.5 millimetres in most specimens, with one measuring 0.6 millimetres. The corpus bursae is small and often globular, and the scobinate signum is small and round, with weak lateral spinules measuring 0.10–0.17 millimetres in diameter. Female P. incultaria has a generally longer colliculum of 0.6–0.7 millimetres and a larger, more elongate and often triangular signum of 0.13–0.3 millimetres, accompanied by stout lateral spinules. The sexes therefore differed in several reproductive structures, although the strongest visible separation was in the male genitalia.

Genome-wide data independently recovered the same two lineages. The ddRAD workflow generated an average of 1.54 million reads per individual, of which 84.8% remained after stringent quality filtering. A de novo assembly recovered 1,042 putative orthologues shared by more than three samples, covering 193,994 base pairs in total. This dataset contained 1,678 SNPs, including 200 parsimony-informative sites. A reference assembly mapped against the genome of the distantly related moth Operophtera brumata yielded a smaller final dataset of 50 retained loci across more than three individuals. Maximum-likelihood trees constructed independently from the de novo and reference datasets produced congruent arrangements, with the P. incultaria and P. barrassoi lineages receiving 100% bootstrap support. Population-clustering analysis likewise identified two discrete genetic groups corresponding to the two species. Only one of the eight individuals tested carried the bacterial parasite Wolbachia.

The revision also addressed the uncertain name-bearing material of P. incultaria. Its original description was based on two males and one female from the Alps. Two collectors were mentioned, with Gastein in Salzburg associated with the material, but the type specimens were presumed destroyed and could not be traced in museums in London or Berlin. Because P. incultaria cannot be separated externally from P. barrassoi and both may occur around Gastein, the authors designated a neotype to preserve nomenclatural stability. The chosen male was collected at Bad Hofgastein, Schlossalm, at 2,000 metres on 6 August 1975. High-throughput sequencing recovered a 658-base-pair fragment that clustered unambiguously with P. incultaria, fixing the name to a documented, genetically identified individual.

Ecological and seasonal records suggest reasons why P. barrassoi may have escaped attention. It is a montane, univoltine species recorded from approximately 700 metres in Bavarian and Austrian valleys to 2,500 metres in the Maiella Mountains. Other sites span about 1,000–1,300 metres in the Bavarian and Austrian Alps, 1,500–2,100 metres in the French Alps and Pyrenees, 1,800 metres on Monte Terminillo and 2,450 metres on the Adamello. Its central Italian specimens were recorded in mid- and late July, and its Pyrenean and French Alpine flight period extends from early July to early August. Alpine records, however, range from mid-April to late May, with one eastern Tyrolean specimen from mid-June and northern Italian specimens from late June. Where the two species coexist, P. barrassoi generally flies earlier than P. incultaria, whose Alpine populations usually fly from early June to early August. The larval stages and host plants of P. barrassoi remain unknown, although its sister species feeds on Primulaceae in early larval stages and on Saxifragaceae later, and representatives of both plant families occur at P. barrassoi collection sites.

On the present evidence, P. barrassoi is confirmed in the central Apennines, the Pyrenees, the French Alps, the Adamello area of northern Italy, eastern and northern Tyrol in Austria, and southernmost Bavaria in Germany. These confirmed records consist of distinct subareas, and the authors considered incomplete recording a more likely explanation than a genuinely fragmented distribution. They predicted that additional populations could remain in the northern Apennines and the western Alps, including Switzerland, Piedmont and Valle d’Aosta, and noted that the species’ early flight period and close resemblance to P. incultaria could make it easy to overlook. That wider range remains a prediction rather than a documented finding. Within the studied material, however, the combined barcode, nuclear-genomic, anatomical and ecological evidence establishes P. barrassoi as a distinct species occurring alongside P. incultaria across previously unrecognized parts of Central and Western Europe.

Subject of Research: Taxonomic identification and European distribution of the geometrid moth Perizoma barrassoi

Article Title: DNA barcoding and genomics reveal Perizoma barrassoi Zahm, Cieslak & Hausmann, 2006 as new for the fauna of Central Europe ( Lepidoptera , Geometridae , Larentiinae )

Article References: Hausmann, A., Huemer, P., Lee, K. M., & Mutanen, M. (2021). DNA barcoding and genomics reveal Perizoma barrassoi Zahm, Cieslak & Hausmann, 2006 as new for the fauna of Central Europe (Lepidoptera, Geometridae, Larentiinae). Nota Lepidopterologica, 44, 17-28. https://doi.org/10.3897/nl.44.58871

Image Credits: AI Generated

DOI: 10.3897/nl.44.58871

Keywords: Perizoma barrassoi, DNA barcoding, ddRAD sequencing, genome-wide SNPs, cryptic species, Geometridae, Lepidoptera, moth taxonomy, genital morphology, European fauna, barcoding, genomics