Priors and Posteriors in Bayesian Timing of Divergence Analyses: The Age of Butterflies Revisited.

The need for robust estimates of times of divergence is essential for downstream analyses, yet assessing this robustness is still rare. We generated a time-calibrated genus-level phylogeny of butterflies (Papilionoidea), including 994 taxa, up to 10 gene fragments and an unprecedented set of 12 fossils and 10 host-plant node calibration points. We compared marginal priors and posterior distributions to assess the relative importance of the former on the latter. This approach revealed a strong influence of the set of priors on the root age but for most calibrated nodes posterior distributions shifted from the marginal prior, indicating significant information in the molecular data set. Using a very conservative approach we estimated an origin of butterflies at 107.6 Ma, approximately equivalent to the latest Early Cretaceous, with a credibility interval ranging from 89.5 Ma (mid Late Cretaceous) to 129.5 Ma (mid Early Cretaceous). In addition, we tested the effects of changing fossil calibration priors, tree prior, different sets of calibrations and different sampling fractions but our estimate remained robust to these alternative assumptions. With 994 genera, this tree provides a comprehensive source of secondary calibrations for studies on butterflies.

The fossil record and taphonomy of butterflies and moths (Insecta, Lepidoptera): implications for evolutionary diversity and divergence-time estimates.

BACKGROUND: It is conventionally accepted that the lepidopteran fossil record is significantly incomplete when compared to the fossil records of other, very diverse, extant insect orders. Such an assumption, however, has been based on cumulative diversity data rather than using alternative statistical approaches from actual specimen counts. RESULTS: We reviewed documented specimens of the lepidopteran fossil record, currently consisting of 4,593 known specimens that are comprised of 4,262 body fossils and 331 trace fossils. The temporal distribution of the lepidopteran fossil record shows significant bias towards the late Paleocene to middle Eocene time interval. Lepidopteran fossils also record major shifts in preservational style and number of represented localities at the Mesozoic stage and Cenozoic epoch level of temporal resolution. Only 985 of the total known fossil specimens (21.4%) were assigned to 23 of the 40 extant lepidopteran superfamilies. Absolute numbers and proportions of preservation types for identified fossils varied significantly across superfamilies. The secular increase of lepidopteran family-level diversity through geologic time significantly deviates from the general pattern of other hyperdiverse, ordinal-level lineages. CONCLUSION: Our statistical analyses of the lepidopteran fossil record show extreme biases in preservation type, age, and taxonomic composition. We highlight the scarcity of identified lepidopteran fossils and provide a correspondence between the latest lepidopteran divergence-time estimates and relevant fossil occurrences at the superfamily level. These findings provide caution in interpreting the lepidopteran fossil record through the modeling of evolutionary diversification and in determination of divergence time estimates.

Morphology-based phylogeny and biogeography of Wockia (Lepidoptera: Urodidae) with description of a new species from Japan and South Korea.

Wockia, one of the six genera within the lepidopteran family Urodidae, currently includes nine species distributed in Holarctic, Oriental, and Neotropical regions. A new species of the genus, W. magna sp. nov., is described from Japan and South Korea. This is the first record of Urodidae from Japan. A cladistic analysis was constructed based on 27 morphological characters from five ingroup species and two outgroup taxa. A single most parsimonious tree was found (length = 38, CI = 71, RI = 70). The resulting tree failed to recover the monophyly of Wockia chewbacca and a clade Wockia sensu stricto, including four congeners from North America, temperate East Asia and Southeast Asia. Wockia sensu stricto was resolved as monophyletic and was divided into two subgroups, one including W. asperipunctella and W. magna, and the other including W. koreana and W. variata. The sister-group relationship of W. asperipunctella and W. magna was moderately supported. Synapomorphies of Wockia sensu stricto are provided from the resulting phylogeny. Systematic definitions of Wockia and other allied genera are revised. Optimal ancestral area reconstruction implemented in DIVA resulted in different hypotheses for Wockia, depending on how to constrain the maximum number of areas. The maximum area number set as two yielded a more likely scenario suggesting that Wockia sensu stricto originated in temperate East Asia and then dispersed into the Oriental region, North America, and Europe. A working hypothesis and other equally possible alternative explanations for the biogeography of Wockia are provided.

Review of Microleon Butler, 1885 (Lepidoptera: Limacodidae) from Taiwan with description of a new species based on morphology and DNA barcoding.

The Taiwanese species of Microleon are reviewed with morphological and DNA data. Our review recognized two congeners, including a new species, M. taiwanensis n. sp. and one known species, M. longipalpis Butler, 1885. The findings from this study request a reconsideration of the previous records of Microleon from Taiwan. The minor differences in external appearance among the species of Microleon pose a challenge for reliable identification. Our study shows that examination of the male genitalia and COI barcoding helps distinguish the species. A checklist of the world species of Microleon with their distributional range is provided.

A taxonomic review of Attevidae (Lepidoptera: Yponomeutoidea) from China with descriptions of two new species and a revised identity of the Ailanthus webworm moth, Atteva fabriciella, from the Asian tropics.

This review describes four species of Atteva (Lepidoptera: Yponomeutoidea: Attevidae) from China, including two new species: A. wallengreni n. sp. and A. yanguifella n. sp. The taxonomic identity of the Ailanthus webworm moth from South and Southeast Asia is revised with a designation of neotype for Phalaena (Tinea) fabriciella Swederus. Lectotypes of Atteva brucea Moore and A. niviguttella Walker are designated. Atteva brucea is synonymized with A. fabriciella. Synonymy of Atteva niviguttella and A. fabriciella is reconfirmed. The previous Chinese records of A. fabriciella were based on confusions with A. wallengreni n. sp. Confirmed specimens of A. fabriciella from China are reported. A pair of confused species, A. fabriciella and A. wallengreni n. sp., are distinguished by the number of white dots on the forewings and the genital features. Another confused pair, A. niveigutta and A. yanguifella n. sp., are compared by external and genital features. All type specimens of the described species are illustrated and compared with the conspecific specimens from various countries of the Asian tropics. Keys to all the species from China are provided.

A new species of Mataeomera, formerly misidentified as M. obliquisigna (Hampson, 1894) from Japan (Lepidoptera, Erebidae, Boletobiinae).

Mataeomera Butler, 1886, currently referred to the tribe Eublemmini in Boletobiinae (Holloway 2011), was proposed by Butler (1886) to accommodate an Australian species, Mataeomera dubia Butler. This genus was once regarded as a junior synonym of Autoba Walker by Poole (1989) but resurrected by Edwards (1996). Another related genus, Catoblemma Hampson, 1910 originally proposed for Catoblemma sumbavensis Hampson, 1910 from Lesser Sunda was revised to include the Australian and East and Southeast Asian species. Edwards (1996) found the Australian species of Catoblemma and Mataeomera dubia congeneric and established the synonymy of those two genera. Following Edwards (1996), Yoshimoto (1999) transferred the Asian species of Catoblemma to Mataeomera. Mataeomera currently includes 21 species (Poole 1989; Edwards 1996; Holloway 2009). Following Holloway (2009), we note that Catoblemma appears to include several unrelated species and is probably polyphyletic. In fact, the Indo-Himalayan and East Asian species currently assigned to Mataeomera are distinct from the Australian congeners in external appearance and larval feeding habits (Sohn & Ronkay 2001; Holloway 2009). As such these may merit generic status but this possibility is still under study by the first author (JCS). 

Systematics, phylogeny and biology of a new genus of Lithocolletinae (Lepidoptera: Gracillariidae) associated with Cistaceae.

The gracillariid genus Triberta gen. nov. (Lepidoptera: Gracillariidae: Lithocolletinae Stainton, 1854) is described to accommodate two species formerly assigned to the genus Phyllonorycter Hübner, 1822: Triberta helianthemella (Herrich-Schäffer, 1861) comb. nov. and T. cistifoliella (Groschke, 1944) comb. nov. Triberta cistifoliella bona sp. is restored from synonymy based on morphological characters. The new genus is biologically associated with the plant family Cistaceae of the order Malvales and is endemic to the Palaearctics. Our molecular analysis of eleven nuclear genes failed to unambiguously place Triberta in the lithocolletine phylogeny, but revealed that this genus is distinct from either clade Phyllonorycter + Cremastobombycia and Cameraria. The distinctiveness of Triberta is also supported by inferred traits in wing venation, micro morphology of the last instar larva, pupa, genital morphology of the adult and life history. A key to the species of Triberta is provided. The interspecific homogeneity in external morphology, coupled with minor differences in genital traits, an apparent narrow specialization on Cistaceae host plants, restricted geographical range and molecular evidence based on multi-nuclear genes jointly suggest that the generic diversification of Triberta is a relatively old phenomenon and driven strongly by host selection.

A new species of Eucalantica Busck (Lepidoptera, Yponomeutidae) poses potential concerns on blueberry industry in Mexico.

A new species of Eucalantica, E. nestori n. sp. is reported on the basis of 13 type specimens collected from blueberry farms in Atzingo and Xoxonacatla, Puebla, Mexico. Habitus and genital features of the species are described and illustrated. Larval feeding habit of E. nestori n. sp. is demonstrated from field observations. Damage of the larvae was observed from the leaves, flowers and shoots of blueberries. Agricultural concerns involving E. nestori n. sp. are discussed.

Review of Calauta Solovyev amp; Witt, 2009 (Lepidoptera: Limacodidae) with description of a new species from Korea.

The genus Calauta Solovyev Witt, 2009 is reviewed with four included species. It is critically compared to other similar genera in Limacodidae based on external and genital morphology. A new species, Calauta koreana sp. nov. is described from the Republic of Korea. A taxonomic transfer of Aphendala notoseusa Wu, 2020 to Calauta is proposed. A distributional range of C. obscura Solovyev, 2017 is expanded with new distributional data. A diagnostic key of all the congeners is given. Photographs of habitus and genitalia are provided for all four species of Calauta.

The first complete mitochondrial genome in the family Attevidae (Attevaaurea) of the order Lepidoptera.

The superfamily Yponomeutoidea, one of the early-derived groups in the order Lepidoptera, consists of 11 families. However, mitochondrial genome (mitogenome) sequences, popularly used for phylogeny and evolutionary tracing, are available for only seven species across six genera and five families. Thus, a larger variety of mitogenome sequences in Yponomeutoidea are required to improve our understanding of lepidopteran phylogeny and genomic evolution. In this study, we present the complete mitogenome of Attevaaurea (Fitch, 1856), the first species in the family Attevidae (superfamily Yponomeutoidea, order Lepidoptera) to be sequenced. The complete mitogenome comprises 16,329 bp and contains a typical set of genes and one non-coding region. Within Yponomeutoidea, the mitogenome of A.aurea has a unique trnI-trnM-trnQ arrangement at the A + T-rich region and ND2 junction and trnA-ND3 arrangement at the trnG and trnR junction. Twelve of the 13 protein-coding genes (PCGs) of A.aurea have a typical ATN starting codon, whereas COI has the atypical CGA codon, which is frequently found in the starting region of lepidopteran COI. Phylogenetic analyses, based on the concatenated sequences of 13 PCGs and two rRNA genes, using the Maximum Likelihood method, revealed a sister relationship between Attevidae and Praydidae with moderately low nodal support (bootstrap support = 64%).

Review of Lypusinae (Lepidoptera: Gelechioidea: Lypusidae) from Korea with a description of a new species of Agnoea Walsingham, 1907.

The subfamily Lypusinae of Korea is reviewed with three species including a new species, Agnoea danguni Sohn, sp. nov. belonging to nominotypical subgenus, and two species of Agnoea belonging to the subgenus Tubuliferodes Toll, 1956: A. (T.) digitiella Kim, 2020 and A. (T.) josephinae (Toll, 1956). The female characters of A. digitiella are described for the first time. Additional collecting records are provided for A. digitiella and A. josephinae. Photographs of the habitus and the genitalia are provided for all three species of Agnoea known so far from Korea.

Eidophasia assmanni sp. nov., the first alpine representative of the genus, detected in the Russian Altai Mountains (Lepidoptera, Plutellidae).

Eidophasia assmanni sp. nov., a new species of Plutellidae from the alpine zone of Russian Altai Mountains, is described from diagnostic morphology and DNA barcodes. Male adult and genitalia are illustrated, whereas the female sex remains unknown. The species inhabits alpine scree with patchy herbaceous plants and is considered as possible endemic species of the Altai Mountains. An updated checklist of the 13 global Eidophasia Stephens, 1842 species is provided. The likely polyphyly of the genus is discussed from molecular data of the barcode region of the mt COI gene.

A new genus of Pexicopiini (Lepidoptera: Gelechiidae) for "Gelechia" acanthopis Meyrick, 1932, with review of functional morphology of male genitalia in allied genera.

The systematic status of Gelechia acanthopis Meyrick, 1932 is reviewed. The general and functional morphology of the male genitalia merits a new genus in Pexicopiini, Sitotrogoides gen. nov., for G. acanthopis. The musculoskeletal apparatus of the male genitalia in Sitotrogoides is described for the first time and compared with allied genera. The taxonomic position of the genus Sitotrogoides is determined within the tribe Pexicopiini. The lack of the muscle m3 is suggested as a characteristic associating Sitotrogoides with Sitotroga and differing from all other genera in Pexicopiini. The type species of Sitotrogoides, Gelechia acanthopis Meyrick is redescribed. Our new records of Sitotrogoides acanthopis from Korea suggest its extended distribution out of Japan.

Dataset for effect of elevation on the insect herbivory of Mongolian oaks in the high mountains of southern South Korea.

The data presented in this article are related to the research article "Effect of elevation on the insect herbivory of Mongolian oaks in the high mountains of southern South Korea" (Sohn et al., 2019). We collected oak leaves occurring in two mountains: Jirisan Mountain on the mainland of Korea (12 September 2015) and Hallasan Mountain on Jejudo Island (21 September 2015). From three randomly-chosen trees, we sampled six branches with breast height with all leaves attached. Feeding traces associated with feeding activities of insects and mites on the leaves were recorded. The underlying data of that research article are presented here: Feeding damage type per surveyed leaf at four research sites of southern South Korea; the geographic location of the study sites on each mountain and the abbreviation of the feeding types and guilds.

New fossils of Elateridae (Insecta, Coleoptera) from Early Cretaceous Jinju Formation (South Korea) with their implications to evolutionary diversity of extinct Protagrypninae.

Two new genera and species of Elateridae, Megalithomerus magohalmii gen. et sp. nov. and Koreagrypnus jinju gen. et sp. nov., are described based on two pairs of fossils from the late Early Cretaceous Jinju Formation in Jinju City, South Korea. Both Megalithomerus and Koreagrypnus represent the youngest occurrences of an extinct elaterid subfamily, Protagrypninae. Megalithomerus magohalmii is the largest known fossil elaterid. These newly described elaterids provide a better understanding of the morphological diversity and occurrence of Protagrypninae through geologic time.

Two new species of Gisilia Kasy, 1968 (Lepidoptera, Cosmopterigidae) from Korea with first report of piercing oviscapts in Gelechioidea.

Two new species of Gisilia Kasy, 1968, G. melanobasis n. sp. and G. tamrae n. sp., are described from Korea. Ascalenia thoracista (Meyrick, 1915) is transferred to Gisilia and compared to two Korean congeners. External and genital features of two new species and G. thoracista are illustrated. Piercing oviscapts are reported from Gelechioidea for the first time. The occurrences of oviscapts in Lepidoptera are reviewed.

A new species of Naryciodes Matsumura, 1931 (Lepidoptera: Limacodidae) from Korea.

The genus Naryciodes was originally proposed as a monobasic genus (type species: Naryciodes posticalis Matsumura) by Matsumura (1931) in Tineidae. About fifty years later, the genus was transferred to Limacodidae by Inoue (1982) because the larvae possess an autapomorphic character of Limacodidae: the lack of crochets on the prolegs (Epstein et al. 1999).

Revised status of Oridryas Meyrick, 1938: new synonyms of Ypsolopha Latreille, 1796, and Ypsolopha parenthesella (Linnaeus, 1761 (Yponomeutoidea: Ypsolophidae).

The eminent entomologist Aristide Caradja (1861-1955) published between 1925 and 1939, on his own or jointly with Edward Meyrick, a series of taxonomic papers on Chinese microlepidoptera including the descriptions of 18 new species in Yponomeutidae (Hyponomeutidae of authors). Their type specimens are currently deposited in two institutions, the Natural History Museum in London (BMNH: formerly British Museum of Natural History), as previously incorporated in the Meyrick collection, and the "Grigore Antipa" National Museum of Natural History in Bucharest, Romania (MGAB), as incorporated in the Caradja collection. The BMNH specimens were examined and illustrated by Clarke (1965). The type specimens of Oridryas kept in MGAB have never been studied since the original descriptions.

Revised taxonomic status of Eidophasia zukowskyi Amsel, 1938 (Lepidoptera, Plutellidae) with first description of its male and female genitalia.

We revise the taxonomic status of Eidophasia zukowskyi Amsel, 1938, and we provide the first descriptions of the genitalia of both sexes. A diagnostic summary is given with illustrations for a similar congener, Eidophasia syenitella (Herrich-Schäffer, [1854]).