Discovery of a peculiar insular race of Ravenna nivea (Nire, 1920) (Lepidoptera: Lycaenidae) endemic to Yinggeling Mountain of Hainan, suggesting heterogeneous geological history of mountain formation of the island.

A peculiar population of Ravenna nivea (Nire, 1920) was discovered from the Yinggeling Mountain Mass of central Hainan. Its wing pattern and COI barcode data show considerable distinction from other geographic populations of R. nivea, including that of Bawangling, approximately only 40 km away and also located in Hainan. The p-distance value of the COI barcode between the Yinggeling and Bawangling populations was 1.1%, considerably higher than the value (0.6%) between Bawangling population and populations in eastern China, where the subspecific name howarthi Saigusa, 1993 applies. The population is regarded as a distinct subspecies ngiunmoiae Lo & Hsu, subsp. nov. The distinctness and high degree of COI haplotype diversity of R. nivea found in Hainan and Taiwan suggest continental islands may serve as glacial refugees for the butterfly and other organisms during previous glaciations, and the presence of the relict populations of montane butterflies like R. nivea may provide useful clues towards a better understanding of the geological history of mountain formation within islands.

Comprehensive phylogeny of Pieridae butterflies reveals strong correlation between diversification and temperature.

Temperature is thought to be a key factor influencing global species richness patterns. We investigate the link between temperature and diversification in the butterfly family Pieridae by combining next generation DNA sequences and published molecular data with fine-grained distribution data. We sampled nearly 600 pierid butterfly species to infer the most comprehensive molecular phylogeny of the family and curated a distribution dataset of more than 800,000 occurrences. We found strong evidence that species in environments with more stable daily temperatures or cooler maximum temperatures in the warm seasons have higher speciation rates. Furthermore, speciation and extinction rates decreased in tandem with global temperatures through geological time, resulting in a constant net diversification.

Systematics and evolutionary dynamics of insect-fern interactions in the specialized fern-spore feeding Cuprininae (Lepidoptera, Stathmopodidae).

Fern-spore-feeding (FSF) is rare and found in only four families of Lepidoptera. Stathmopodidae is the most speciose family that contains FSF species, and its subfamily Cuprininae exclusively specializes on FSF. However, three species of Stathmopodinae also specialize on FSF. To better understand the evolutionary history of FSF and, more generally, the significance of specialization on a peculiar host, a phylogenetic and taxonomic revision for this group is necessary. We reconstructed the most comprehensive molecular phylogeny, including one mitochondrial and four nuclear genes, of Stathmopodidae to date, including 137 samples representing 62 species, with a particular focus on the FSF subfamily, Cuprininae, including 33 species (41% of named species) from 6 of the 7 Cuprininae genera. Species from two other subfamilies, Stathmopodinae and Atkinsoniinae, were also included. We found that FSF evolved only once in Stathmopodidae and that the previous hypothesis of multiple origins of FSF was misled by inadequate taxonomy. Moreover, we showed that (1) speciation/extinction rates do not differ significantly between FSF and non-FSF groups and that (2) oligophage is the ancestral character state in Cuprininae. We further revealed that a faster rate of accumulating specialists over time, and thus a higher number of specialists, was achieved by a higher transition rate from oligophagages to specialists compared to the transition rate in the opposite direction. We finish by describing three new genera, Trigonodagen. nov., Petalagen. nov., and Pediformisgen. nov., and revalidating five genera: Cuprina, Calicotis, Thylacosceles, Actinoscelis, Thylacosceloides in Cuprininae, and we provide an updated taxonomic key to genera and a revised global checklist of Cuprininae.

To and fro in the archipelago: Repeated inter-island dispersal and New Guinea's orogeny affect diversification of Delias, the world's largest butterfly genus.

The world's largest butterfly genus Delias, commonly known as Jezebels, comprises ca. 251 species found throughout Asia, Australia, and Melanesia. Most species are endemic to islands in the Indo-Australian Archipelago or to New Guinea and nearby islands in Melanesia, and many species are restricted to montane habitats over 1200 m. We inferred an extensively sampled and well-supported molecular phylogeny of the group to better understand the spatial and temporal dimensions of its diversification. The remarkable diversity of Delias evolved in just ca. 15-16 Myr (crown age). The most recent common ancestor of a clade with most of the species dispersed out of New Guinea ca. 14 Mya, but at least six subsequently diverging lineages dispersed back to the island. Diversification was associated with frequent dispersal of lineages among the islands of the Indo-Australian Archipelago, and the divergence of sister taxa on a single landmass was rare and occurred only on the largest islands, most notably on New Guinea. We conclude that frequent inter-island dispersal during the Neogene-likely facilitated by frequent sea level change-sparked much diversification during that period. Many extant New Guinea lineages started diversifying 5 Mya, suggesting that orogeny facilitated their diversification. Our results largely agree with the most recently proposed species group classification system, and we use our large taxon sample to extend this system to all described species. Finally, we summarize recent insights to speculate how wing pattern evolution, mimicry, and sexual selection might also contribute to these butterflies' rapid speciation and diversification.

A global phylogeny of butterflies reveals their evolutionary history, ancestral hosts and biogeographic origins.

Butterflies are a diverse and charismatic insect group that are thought to have evolved with plants and dispersed throughout the world in response to key geological events. However, these hypotheses have not been extensively tested because a comprehensive phylogenetic framework and datasets for butterfly larval hosts and global distributions are lacking. We sequenced 391 genes from nearly 2,300 butterfly species, sampled from 90 countries and 28 specimen collections, to reconstruct a new phylogenomic tree of butterflies representing 92% of all genera. Our phylogeny has strong support for nearly all nodes and demonstrates that at least 36 butterfly tribes require reclassification. Divergence time analyses imply an origin ~100 million years ago for butterflies and indicate that all but one family were present before the K/Pg extinction event. We aggregated larval host datasets and global distribution records and found that butterflies are likely to have first fed on Fabaceae and originated in what is now the Americas. Soon after the Cretaceous Thermal Maximum, butterflies crossed Beringia and diversified in the Palaeotropics. Our results also reveal that most butterfly species are specialists that feed on only one larval host plant family. However, generalist butterflies that consume two or more plant families usually feed on closely related plants.

Rediscovering a species not seen for a hundred years, Stathmopodatacita (Meyrick, 1913) (Lepidoptera, Stathmopodidae), with its unusual fern-spore-feeding life history.

Background: Despite being the second largest group of vascular plants, ferns are scarcely reported being fed by insects when compared to angiosperms. Within these fern-feeding insects, lepidopterans are poorly represented and are restricted only to specific groups in this speciose order. The consumers specialising on fern spores are even scarcer in the order, with the majority being consumers of vegetative structures. Amongst the fern-spore-feeding Lepidoptera, Stathmopodidae is the family with the highest species diversity, even with a subfamily, Cyprininae Sinev, 2015, specialising on fern spores. However, fern-spore-feeding habit is not restricted to this subfamily. To understand the evolution of fern-spore-feeding within this family and to increase our knowledge of insect-fern evolution, detailed studies on fern-spore feeding stathmopodids are essential. New information: The present study rediscovered a rare, fern-spore-feeding, stathmopodid micro-moth, Stathmopodatacita (Meyrick, 1913), which has not been formally recorded or identified for more than 100 years. We documented the life history of this species and identified several species of Pyrrosia (Polypodiaceae, Platycerioideae) as host for the moth's larvae. A re-description of the fern-feeding moth is also provided as the original description is obscure in terms of character diagnosis.

Alien species revises systematic status: integrative species delimitation of two similar taxa of Symbrenthia Hübner, [1819] (Lepidoptera, Nymphalidae).

Introduction of organisms to new range may impose detrimental effects on local organisms, especially when closely related species are involved. Species delimitation employing an integrative taxonomy approach may provide a quick assessment for the species status between taxa of interest, and to infer ecological competition and/or introgression that may be associated with the introduction. A nymphalid butterfly, Symbrenthia lilaea lunica, was recently introduced to Taiwan, where a closely related local taxon, S. l. formosanus, can be found. We employed multiple species delimitation methods to study the species status between the two taxa, and the results revealed that they can be recognized as two distinct species, revised to S. l. lilaea (syn. nov.) and S. formosanus (stat. rev.) respectively. We further performed a niche modeling approach to investigate the ecological interaction between the two species. The taxonomic status of the two taxa, now elevated to species, has been revised and conservation facing rapid expansion of the introduced species discussed.

A new species of the genus Synanthedon Hbner [1819] (Lepidoptera Sesiidae) from Taiwan.

A new species of clearwing moth, Synanthedon suhua sp. nov., is described from Taiwan in this article. Adults and genitalia of both sexes are illustrated, DNA barcodes provided, and potential damage to Quercus longinux (Fagaceae) discussed.

Local adaptation and migratory habits balance spatial-genetic structure between continental and insular chestnut tiger butterflies in East Asia.

Geographic and climatic differences between islands and continents may affect the evolution of their biota, and promote divergent selection in species distributed in both landscapes. To assess spatial-genetic structure, we genotyped 18 expressed sequence tag-simple sequence repeat (EST-SSR) loci and sequenced two mtDNA markers (ND5 and COI) and one nuclear marker (EF1α) in two subspecies of the butterfly Parantica sita. Compared with nuclear markers, mtDNA had a stronger signal of population structure. Approximate Bayesian computation (ABC) suggested that a continuous-gene-flow model best described the data. According to this model, the two subspecies diverged approximately 23.1 kya, with 10 times more introgression from the continental (ssp. sita) to the insular subspecies (ssp. niphonica) than vice versa. Ecological niche modelling was performed to predict the paleo- and current potential distributions and elucidate the geohistorical process, which revealed a northeastern, insular origin. Winter precipitation and annual temperature range were the main determinants of the subspecies distributions. Maximum-likelihood population-effects models showed that the population differentiation of the insular and continental subspecies was primarily affected by environmental resistance and local climate. Sex-biased migration capacity and long-term precipitation-driven divergence between the continental and insular lineages shaped the current genetic structure of P. sita. Evidence from the nuclear markers confirmed intersubspecific gene flow despite adaptive divergence between the subspecies. These results imply that the continental subspecies is still capable of returning to the island and introgressing with the insular subspecies.

The Newly Recorded Fern-spore Feeding Moths in the Genus Calicotis, Meyrick 1889 (Lepidoptera: Stathmopodidae) from Taiwan, with Notes on Life History of Three Species.

Three newly recorded species of the genus Calicotis, Meyrick 1889 are reported from Taiwan: C. attiei (Guillermet, 2011), C. rotundinidus Terada, 2016, and C. exclamationis Terada, 2016. C. biserraticola Terada, 2016 is treated as a junior subjective synonym for C. attiei based on both morphological and molecular data. The life history of these three species is presented as well as the first observation of fern-feeding stathmopodid eggs in the world.

Systematic position of Bombyx incomposita (Lepidoptera: Bombycidae), with notes on its immature biology and hostplant association.

The immature stages, morphology, and systematic position of Bombyx incomposita (van Eecke, 1929) are reported. The morphology of larva was investigated using scanning electron microscopy, and the phylogeny was inferred using six genes (COI, CAD, EF-1, GAPDH, RpS5, and wgl). The molecular phylogeny strongly supported that B. incomposita is sister to B. huttoni Westwood, 1847. The divergence of the COI barcode, phylogenetic reconstruction, and difference in immature stages support the current classification, in which the species incomposita belongs to the genus Bombyx. We give illustrations of male and female genitalia and briefly discuss the utility of the chaetotaxy of the first instar larvae on taxonomy of silkmoths. The SV group bears six setae in B. mandarina, but eight in B. incomposita. The L1 bears three setae in Rotunda, but just single seta in Bombyx.

On a New Insular Subspecies of Chrysozephyrus mushaellus (Matsumura) (Lepidoptera: Lycaenidae) Discovered from Hainan.

Chrysozephyrus mushaellus paolongkoui, ssp. nov. is described and illustrated from Hainan island of southern China. This new subspecies possesses diagnosable characters different from previously recognized subspecies of C. mushaellus in wing marking and genitalia of both sexes. It utilizes Lithocarpus litseifolius (Fagaceae) as the larval hostplant.

A new genus and species of slug caterpillar (Lepidoptera: Limacodidae) from Taiwan.

The genus Epsteinius Lin, Braby Hsu gen. nov. is established to accommodate a new, diminutive species of limacodid moth, E. translucidus Lin, sp. nov., from central Taiwan. Although the new species is the sister taxon to Microleon Butler, 1885 according to molecular phylogenetic data, it differs fundamentally from that genus in adult morphology, wing color pattern, male and female genitalia, and larval morphology. Larvae of E. translucidus Lin, sp. nov. represent an interesting case in which crypsis is based on transparency through its body color, resting behavior, and feeding pattern.

The types of skippers described by Shu-Iti Murayama (Lepidoptera: Hesperiidae).

Fifteen holotypes of Asian Hesperiidae taxa described by Shu-iti Murayama were examined, and their taxonomic status is discussed. We confirm that five are valid names while the rest are synonyms as indicated partially by previous authors. Valid species-level names are Aeromachus matudai (Murayama), Aeromachus bandaishanus Murayama Shimonoya, Coladenia pinsbukana (Shimonoya Murayama), and Sebastonyma suthepiana Murayama Kimura. Valid subspecies-level name is Ochlodes yuchingkina Murayama Shimonoya. In order to settle the taxonomic status of Pedesta masuriensis cuneomaculata Murayama, masuriensis and tali were studied morphologically and molecularly. As a result, we consider that masuriensis and tali are two different species and treat cuneomaculata as a junior subjective synonym of tali.

On systematic status of Spindasis syama Horsfield, [1829] in Taiwan and the Philippines (Lepidoptera: Lycaenidae: Aphnaeini).

Based upon both morphological and gene barcode evidence, the population of Spindasis syama in Taiwan is described as Spindasis syama lamuae, ssp. nov., and the taxon negrita Felder, 1862 is revised as a species endemic to the Philippines (stat. rev.).

Revision of Pazala Moore, 1888: The Graphium (Pazala) alebion and G. (P.) tamerlanus Groups, with Notes on Taxonomic and Distribution Confusions (Lepidoptera: Papilionidae).

Three Graphium species belonging to two species groups of the subgenus Pazala, the alebion and tamerlanus groups, were examined in molecular and morphological studies, and their female genitalia are reported for the first time. Their relationship with other species groups within the subgenus is assessed and their divergence times are estimated. We find that G. (P.) alebion is the first lineage to diverge within Pazala in the early Miocene (20 Ma) and that G. (P.) tamerlanus and G. (P.) parus are sister species and diverged from each other in the late Miocene (7 Ma). A revision of the four recognised taxa belonging to three species is presented, and historical misidentification of these taxa and their geographic ranges are explained.

The identity of Alfred Wallace's mysterious butterfly taxon Lycaena nisa solved: Famegana nisa comb. nov., a senior synonym of F. alsulus (Lepidoptera, Lycaenidae, Polyommatinae).

Lycaena nisa Wallace, 1866 was described from Formosa (Taiwan) and is here recognized as a senior subjective synonym of Lycaena alsulus Herrich-Schäffer, 1869. It is resurrected to serve as the valid name, Famegana nisa (Wallace, 1866), comb nov. of the species commonly known as Famegana alsulus. The name Zizera taiwana Sonan, 1938 (syn. nov.), also described from Formosa, is recognized as a junior subjective synonym of L. nisa. Another name, Zizeeria alsulus eggletoni Corbet, 1941 (syn. nov.), described from Hong Kong is also considered a junior subjective synonym of L. nisa. Moreover, all former synonyms of alsulus automatically become new junior synonyms for nisa. This species occurs in the Oriental and Australian regions and western Pacific.

The fern-feeding genus Cuprina Sinev, 1988 (Lepidoptera, Stathmopodidae), new for Taiwan, with descriptions of two new species.

Two new species of Stathmopodidae are described from Taiwan: Cuprina atayalica Shen & Hsu, sp. nov., reared from larvae on Microsorum brachylepis, and C. insolita Hsu & Shen, sp. nov., reared from larvae on Tectaria subtriphylla and T. harlandii. Diagnostic characters for both species are provided. Larval host plants and the biology of the immature stages of both new species are documented.

Vibrational communication between a myrmecophilous butterfly Spindasis lohita (Lepidoptera: Lycaenidae) and its host ant Crematogaster rogenhoferi (Hymenoptera: Formicidae).

Ants are a dominant insect group in terrestrial ecosystems and many myrmecophilous species evolve to associate with ants to gain benefits. One iconic example is myrmecophilous butterflies that often produce ant-mimicking vibrational calls to modulate ant behaviors. Despite its popularity, empirical exploration of how butterflies utilize vibrational signals to communicate with ants is scarce. In this study, we reported that the myrmecophilous butterfly Spindasis lohita produce three types of larval calls and one type of pupal call, while its tending ant, Crematogaster rogenhoferi emit a single type of call. The results of discriminant analysis revealed that calls of the two species are quantitatively similar in their signal attributes; the potential role of butterfly calls are further confirmed by the playback experiments in which certain ant behaviors including antennation, aggregation, and guarding were induced when one of the butterfly calls was played to C. rogenhoferi workers. The findings in the current study represent the very first evidence on vibrational communication between Spindasis and Crematogaster and also imply that S. lohita may have been benefited from ant attendance due to the ability to produce similar calls of the ant C. rogenhoferi.

On revised systematic status of Mycalesis suaveolens kagina Fruhstorfer, 1911, with notes on its immature biology and hostplant associations.

Mycalesis kagina Fruhstorfer, 1911 is separated from Mycalesis suaveolens Wood-Mason de Nicéville, 1883 to represent a species endemic to Taiwan based upon COI barcode divergence, morphological diagnosis of larva, and genitalia of both sexes. Both kagina and suaveolens are confirmed as members of the genus Mycalesis in Mycalesina. Immature morphology, biology, and hostplant associations for both species are given for the first time. Larvae of both species are recognized as specialists on Zingiberaceae, a plant family rarely used by satyrid butterflies.