Wetland butterfly thriving in abandoned jungle: Neptis rivularis in the Czech Republic.

With ongoing insect declines, species expanding in distribution and abundance deserve attention, as understanding their success may help design conservation strategies for less successful species. Common causes of these successes include warmer climates, novel resources, and exploiting land use change, including land abandonment. These factors affect the nymphalid butterfly Neptis rivularis, developing on Spiraea spp. shrubs and reaching the north-western limits of its trans-Palearctic distribution in Central Europe. We combined mark-recapture, behaviour analysis, and distribution modelling to study N. rivularis in wetlands of the Trebonsko Protected Landscape (IUCN category V). The long-living adults (up to 4 weeks) spent a considerable amount of time searching for partners, ovipositing and nectaring at Spiraea shrubs, alternating this with stays in tree crowns, where they located cool shelters, spent nights, and presumably fed on honeydew. They formed high-density populations (310 adults/ha), exploiting high host plant abundance. They adhered to floodplains and to conditions of relatively mild winters. The ongoing Spiraea encroachment of abandoned alluvial grasslands is, thus, a transient situation, ultimately followed by forest encroachment. Rewilding the habitats by introducing native ungulates presents an opportunity to restore the disturbance regime of the sites. The increased resource supply combined with a warming climate has opened up temperate Europe to colonization by N. rivularis.

Perishing rich, expanding poor: Demography and population genetic patterns in two congeneric butterflies.

In human-altered landscapes, specialist butterflies typically form spatially restricted populations, genetically differentiated due to dispersal restrictions. Generalists, in contrast, display minimum differentiation but high genetic diversity. While local-level actions suffice to conserve specialists and landscape-level actions are necessary for generalists, minimum information exists regarding conservation of species with intermediate features. We targeted two congeneric butterflies, the recently re-expanding Argynnis adippe and the strongly declining A. niobe, co-occurring in the pastoral landscape of the Carpathian Mountains, Czech Republic. We integrated species distribution models, mark-recapture and microsatellite analysis to compare their habitat requirements, adult demography, dispersal and genetic patterns, and expanded the genetic analysis across the Carpathian Arc and beyond to delimit spatial conservation units. In two mountain valleys, both species formed interconnected populations numbering thousands of individuals. Mobility patterns suggested the populations' interconnection across the Czech Carpathians. Genetic diversity was extremely poor in the nonthreatened A. adippe and moderate in the declining A. niobe. No population differentiation was detected within the Czech Carpathians (~1500 km2 ). Low genetic diversity and no differentiation was preserved in A. adippe across East Central Europe, whereas in A. niobe, populations from Serbia were differentiated from the Carpathian Arc + Alps. The high adult mobility linked to low differentiation probably reflects the distribution of larval resources, historically widespread but sparse and currently declining for A. niobe (grazing-disturbed grounds), while currently increasing for A. adippe (abandonment scrub, disturbed woodlands). Units as large as entire mountain systems define population boundaries, and hence conservation management units, for both species.

Molecular phylogeny and biogeography of the genus Symbrenthia (Lepidoptera, Nymphalidae) correlates with the past geography of the Oriental region.

Jesters, butterflies in the genus Symbrenthia Hübner, 1819, comprise 14 species mainly distributed in the Oriental region. Although this genus has attracted the attention of many researchers in the past, its taxonomy and biogeographic history remain unclear. In this study, we investigate phylogenetic and biogeographic relationships inferred from one mitochondrial (COI) and two nuclear genes (ArgKin, wingless), using both likelihood and Bayesian approaches. With the exception of S. hippalus, which we find to be either sister to Mynes Boisduval, 1832 or sister to Symbrenthia + Mynes + Araschnia, all species of Symbrenthia form a single monophyletic group. We describe a new genus Mynbrenthia Fric & Rindos gen. nov. to accommodate the taxon hippalus. The genus Symbrenthia splits into four sub-groups, "Brensymthia" (with S. niphanda and S. sinoides), "hypselis" (with S. hypselis, S. brabira, S. leoparda and S. doni), "intricata" (with S. intricata and S. hypatia) and "hippoclus" group (including S. platena and a complex of S. hippoclus and S. lilaea). The genus probably originated in Sundaland or continental Asia during the Eocene. The history of the genus Symbrenthia was more influenced by dispersal events and then by subsequent vicariances. Whereas the "hypselis" group colonised the Indo-Australian Archipelago from the Asian continent, the "hippoclus" group dispersed to continental Asia from the Indo-Australian Archipelago.

Radical pruning of distribution data may result in loss of knowledge (Response to Larsen & Shirey).

Larsen & Shirey (2020) criticised our analysis of latitudinal changes in butterfly phenology on the grounds of improper data management. We admit some imprecisions, but show that stringent reanalyses did not change the overall results. We also show that unreasonable treatment of data may result in critical information loss.

Phenology responses of temperate butterflies to latitude depend on ecological traits.

Global change influences species' seasonal occurrence, or phenology. In cold-adapted insects, the activity is expected to start earlier with a warming climate, but contradictory evidence exists, and the reactions may be linked to species-specific traits. Using data from the GBIF database, we selected 105 single-brooded Holarctic butterflies inhabiting broad latitudinal ranges. We regressed patterns of an adult flight against latitudes of the records, controlling for altitude and year effects. Species with delayed flight periods towards the high latitudes, or stable flight periods across latitudes, prevailed over those that advanced their flight towards the high latitudes. The responses corresponded with the species' seasonality (flight of early season species was delayed and flight of summer species was advanced at high latitudes) and oceanic vs. continental climatic niches (delays in oceanic, stability in continental species). Future restructuring of butterfly seasonal patterns in high latitudes will reflect climatic niches, and hence the evolutionary history of participating species.

Five new species of the genus Meganaclia Aurivillius, 1892 (Lepidoptera: Erebidae: Arctiinae: Syntomini).

The Afrotropical genus Meganaclia Aurivillius, 1892 is revised, five new species M. smithi sp. n., M. grehani sp. n., M. josephi sp. n., M. johannae sp. n. and M. gaerberfesti sp. n. are described with a key for identification of species based on male genitalia structures. Habitus and genitalia of all taxa are illustrated and a distribution map is provided.

More hidden diversity in a cryptic species complex: a new subspecies of Leptideasinapis (Lepidoptera, Pieridae) from Northern Iran.

A new subspecies of Leptideasinapis from Northern Iran, discovered by means of DNA barcoding, is described as Leptideasinapistabarestanassp. nov. The new subspecies is allopatric with respect to other populations of L.sinapis and is genetically distinct, appearing as a well-supported sister clade to all other populations in COI-based phylogenetic reconstructions. Details on karyotype, genitalia, ecology and behaviour for the new subspecies are given and a biogeographical speciation scenario is proposed.

Geography of Indian Butterflies: Patterns Revealed by Checklists of Federal States.

Butterflies are widely used to analyze biogeographical patterns, both at the global and regional scales. Thus far, most of the latter originated from well-surveyed northern regions, while the species-rich tropical areas lag due to a lack of appropriate data. We used checklists of 1379 butterfly species recorded in 36 federal states of the Republic of India (1) to explore the basic macroecological rules, and (2) to relate species richness and the distribution of endemics and geographic elements to geography, climate, land covers and socioeconomic conditions of the states. The area, land covers diversity and latitude did not affect species richness, whereas topographic diversity and the precipitation/temperature ratio (energy availability) were positive predictors. This is due the geographic and climatic idiosyncrasies of the Indian subcontinent, with its highest species richness in the small, densely forested mountainous northeast that receives summer monsoons. The peninsular effect that decreases the richness towards the tip of subcontinent is counterbalanced by the mountainous forested Western Ghats. Afrotropical elements are associated with savannahs, while Palearctic elements are associated with treeless habitats. The bulk of Indian butterfly richness, and the highest conservation priorities, overlap with global biodiversity hotspots, but the mountainous states of the Western Himalayas and the savannah states of peninsular India host distinctive faunas.

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.

New and less known Notodontidae from Manipur, India (Lepidoptera: Noctuoidea).

The present study discusses the systematic accounts of 66 species of the family Notodontidae recorded from the Indian state of Manipur. There are some records of the moths from other places including Sikkim (India) and Nepal in addition to Shirui Hill, Manipur. Here, we report, (1) three new species to science viz. Cleapa ukhrulensis Irungbam & Schintlmeister sp. nov., Spatalina rimbiensis Irungbam & Schintlmeister sp. nov. and Hexafrenum tangkhula Irungbam & Schintlmeister sp. nov.; (2) seventeen species for the first time from India: Hupodonta corticalis Butler, 1877, Chalepa binotata Kiriakoff, 1959, Besaia isis Schintlmeister, 1997, Ogulina. argentilinea Cai, 1982, Bireta juncturina (Kiriakoff, 1959), Spatalina melanopa Schintlmeister, 2007, S. desiccata desiccata (Kiriakoff, 1963), S. umbrosa (Leech, 1898), Phalera albocalceolata (Bryk, 1950), Neodrymonia albinomarginata Schintlmeister, 2007, Syntypistis wunna (Schintlmeister, 1997), S. scensus (Schintlmeister, 1997), Harpyia nadiae Morozov, 2013, Hexafrenum viola (Schintlmeister, 1997), Rachiades lichenicolor siamensis Sugi, 1993, Pheosiopsis norina Schintlmeister, 1989, and Periphalera spadixa Wu & Fang, 2003; and (3) one genus for the first time from India: Rachiades Kiriakoff, 1967. Thus, the present study updates the Indian Notodontidae richness to a total of 281 species (including nominotypical subspecies) and 16 subspecies under 120 genera. Here, we provide species diagnoses along with habitus, and genitalia images for each of these newly recorded species.

Microhabitat Selection for Overwintering: Overwintering Conditions of Three Jumping Spiders (Pellenes tripunctatus, P. nigrociliatus, and Attulus penicillatus) Living in Terrestrial Shells in the Czech Republic.

Taking the regular overwintering of spider species in land snail shells as a model, we studied environmental conditions affecting the choice of overwintering sites in three jumping spider species: Pellenes tripunctatus, Pellenes nigrociliatus, and Attulus penicillatus. The research was conducted at 11 steppe localities on calcareous bedrock with abundant empty shells (mainly Caucasotachea vindobonensis and Xerolenta obvia). We documented 889 shells and collected 186 of them, of which 113 were inhabited by 146 spider individuals (13 species). Our three focal species made up 81.5% of these. We found different environmental preferences between the sexes in P. tripunctatus and P. nigrociliatus. These females preferred shells with more vegetation nearby. In the case of P. tripunctatus, these were shells with a higher proportion of herbs, whereas P. nigrociliatus selected for a higher proportion of moss. In the immediate vicinity of the shells, environmental conditions did not differ significantly. We found insufficient A. penicillatus to determine any preferences. We also recorded six overwintering P. tripunctatus individuals in a single shell (in environmental conditions preferred by females), consisting of five females and one male, which indicated an unusual social behavior for these spider species.

Wolbachia affects mitochondrial population structure in two systems of closely related Palaearctic blue butterflies.

The bacterium Wolbachia infects many insect species and spreads by diverse vertical and horizontal means. As co-inherited organisms, these bacteria often cause problems in mitochondrial phylogeny inference. The phylogenetic relationships of many closely related Palaearctic blue butterflies (Lepidoptera: Lycaenidae: Polyommatinae) are ambiguous. We considered the patterns of Wolbachia infection and mitochondrial diversity in two systems: Aricia agestis/Aricia artaxerxes and the Pseudophilotes baton species complex. We sampled butterflies across their distribution ranges and sequenced one butterfly mitochondrial gene and two Wolbachia genes. Both butterfly systems had uninfected and infected populations, and harboured several Wolbachia strains. Wolbachia was highly prevalent in A. artaxerxes and the host's mitochondrial structure was shallow, in contrast to A. agestis. Similar bacterial alleles infected both Aricia species from nearby sites, pointing to a possible horizontal transfer. Mitochondrial history of the P. baton species complex mirrored its Wolbachia infection and not the taxonomical division. Pseudophilotes baton and P. vicrama formed a hybrid zone in Europe. Wolbachia could obscure mitochondrial history, but knowledge on the infection helps us to understand the observed patterns. Testing for Wolbachia should be routine in mitochondrial DNA studies.

Conserved ancestral tropical niche but different continental histories explain the latitudinal diversity gradient in brush-footed butterflies.

The global increase in species richness toward the tropics across continents and taxonomic groups, referred to as the latitudinal diversity gradient, stimulated the formulation of many hypotheses to explain the underlying mechanisms of this pattern. We evaluate several of these hypotheses to explain spatial diversity patterns in a butterfly family, the Nymphalidae, by assessing the contributions of speciation, extinction, and dispersal, and also the extent to which these processes differ among regions at the same latitude. We generate a time-calibrated phylogeny containing 2,866 nymphalid species (~45% of extant diversity). Neither speciation nor extinction rate variations consistently explain the latitudinal diversity gradient among regions because temporal diversification dynamics differ greatly across longitude. The Neotropical diversity results from low extinction rates, not high speciation rates, and biotic interchanges with other regions are rare. Southeast Asia is also characterized by a low speciation rate but, unlike the Neotropics, is the main source of dispersal events through time. Our results suggest that global climate change throughout the Cenozoic, combined with tropical niche conservatism, played a major role in generating the modern latitudinal diversity gradient of nymphalid butterflies.

AN INVENTORY OF THE BUTTERFLIES OF MANIPUR, INDIA (INSECTA: LEPIDOPTERA).

The butterfly fauna of Manipur is poorly known, and a few sporadic studies were carried out decades ago. In this study, butterflies were photographed and/or collected in 80 localities including revisiting of 12 historical localities in the hills and valleys from Manipur state, India. Butterflies were regularly sampled between 2010 and 2019. In the present checklist, we have included both previously published and recently recorded species: 798 species belong to six families; Papilionidae (52 sp.), Pieridae (39 sp.), Riodinidae (17 sp.), Lycaenidae (225 sp.), Nymphalidae (270 sp.), and Hesperiidae (195 sp.). Eight species were rediscovered during the study; Byasa latreillei kabrua, Papilio machaon suroia, Lamproptera meges indistincta, Bhutanitis lidderdalii lidderdalii, Lethe kangjupkula, Una usta usta, Arhopala hellenore hellenore and Celaenorrhinus munda maculicornis. Thirty-two species were new records to Manipur; Papilio agestor agestor, Lamproptera curius curius, Appias albina darada, Artogeia erutae montana, Miletus mallus, Flos fulgida fulgida, Cigaritis nipalicus, Rapala rectivitta, Heliophorus kohimensis, H. tamu, Jamides caeruleus, Tarucus venosus, Everes huegelii dipora, Talicada nyseus nyseus, Lestranicus transpectus, Euploea radamanthus radamanthus, Lethe distans, L. dura gammiei, L. latiaris latiaris, L. sura, Neope pulaha, N. yama yama, Zipaetis scylax, Algia fasciata fasciata, Athyma opaline, Pantoporia paraka paraka, Kallima knyvettii, Celaenorrhinus asmara, Hyarotis adrastus praba, Erionota torus, Baoris penicillata chapmani, and Potanthus mingo ajax of which, Miletus mallus was a new record for India. Ninety species are legally protected in schedules (I, II IV) of Indian (Wildlife) Protection Act, 1972.

Cross-continental phylogeography of two Holarctic Nymphalid butterflies, Boloria eunomia and Boloria selene.

Pleistocene glaciations had significant effects on the distribution and evolution of species inhabiting the Holarctic region. Phylogeographic studies concerning the entire region are still rare. Here, we compared global phylogeographic patterns of one boreo-montane and one boreo-temperate butterflies with largely overlapping distribution ranges across the Northern Hemisphere, but with different levels of range fragmentation and food specialization. We reconstructed the global phylogeographic history of the boreo-montane specialist Boloria eunomia (n = 223) and of the boreo-temperate generalist Boloria selene (n = 106) based on mitochondrial and nuclear DNA markers, and with species distribution modelling (SDM). According to the genetic structures obtained, both species show a Siberian origin and considerable split among populations from Nearctic and Palaearctic regions. According to SDMs and molecular data, both butterflies could inhabit vast areas during the moderate glacials. In the case of B. selene, high haplotype diversity and low geographic structure suggest long-lasting interconnected gene flow among populations. A stronger geographic structuring between populations was identified in the specialist B. eunomia, presumably due to the less widespread, heterogeneously distributed food resources, associated with cooler and more humid climatic conditions. Populations of both species show opposite patterns across major parts of North America and in the case of B. eunomia also across Asia. Our data underline the relevance to cover entire distribution ranges to reconstruct the correct phylogeographic history of species.

Distribution of Ultraviolet Ornaments in Colias Butterflies (Lepidoptera: Pieridae).

Ultraviolet patterns in butterflies have been recognized and studied for many years. They are frequently involved in both intraspecific and interspecific interactions. Only a handful of studies, however, have investigated possible links between ultraviolet (UV) reflectance and ecological properties in some genera of the Lepidoptera as a whole. This study examines the impact of habitat and distribution on UV reflectance patterns on the wings of 106 species and subspecies of Colias butterflies. Based on standardized digital photographs, we performed a multivariate analysis of relations between UV reflectance, preferred habitat (alpine, arctic, dry grasslands, humid, forest, and ubiquitous), and distribution area (Afrotropical, Nearctic, Neotropical, European, Caucaso-Anatolian, boreal Eurasian, Central Asian mountains, northern China and Japan, and northern Oriental region). UV patterns occur more frequently in the male (60 taxa) than in female (25 taxa) Coliads. This difference in presence of UV patterns is used for differentiating between the males and females of a given species or subspecies. Further possible explanations of this phenomenon are also discussed. This study also shows that particular configurations of UV patterns are significantly associated with particular distribution areas. This relation is relatively strong but overall trends remain unclear. Based on the results of this study, it can be concluded that there exists a significant difference in the configuration of UV reflectance between the sexes, and that the configuration of UV reflectance significantly interacts with the geographical distribution of Colias species and subspecies.

An updated checklist of the European Butterflies (Lepidoptera, Papilionoidea).

This paper presents an updated checklist of the butterflies of Europe, together with their original name combinations, and their occurrence status in each European country. According to this checklist, 496 species of the superfamily Papilionoidea occur in Europe. Changes in comparison with the last version (2.6.2) of Fauna Europaea are discussed. Compared to that version, 16 species are new additions, either due to cryptic species most of which have been discovered by molecular methods (13 cases) or due to discoveries of Asian species on the eastern border of the European territory in the Ural mountains (three cases). On the other hand, nine species had to be removed from the list, because they either do not occur in Europe or lost their species status due to new evidence. In addition, three species names had to be changed and 30 species changed their combination due to new evidence on phylogenetic relationships. Furthermore, minor corrections were applied to some authors' names and years of publication. Finally, the name Polyommatusottomanus Lefèbvre, 1831, which is threatened by its senior synonym Lycaenalegeri Freyer, 1830, is declared a nomen protectum, thereby conserving its name in the current combination Lycaenaottomana.

Life History Traits Reflect Changes in Mediterranean Butterfly Communities Due to Forest Encroachment.

The biodiversity of the Southern Balkans, part of the Mediterranean global biodiversity hot-spot, is threatened by land use intensification and abandonment, the latter causing forest encroachment of formerly open habitats. We investigated the impact of forest encroachment on butterfly species richness, community species composition and the representation of life history traits by repeated seasonal visits of 150 one-hectare sites in five separate regions in three countries-Greece, Bulgaria, and the Republic of Macedonia (FYROM-the Former Yugoslav Republic of Macedonia)- 10 replicates for each habitat type of grasslands, open formations and scrub forest within each region. Grasslands and open formations sites hosted in average more species and more red-listed species than scrub forest, while no pattern was found for numbers of Mediterranean species. As shown by ordination analyses, each of the three habitat types hosted distinct butterfly communities, with Mediterranean species inclining either towards grasslands or open formations. Analysing the representation of life history traits revealed that successional development from grasslands and open formations towards scrub forest shifts the community composition towards species overwintering in earlier stages, having fewer generations per year, and inhabiting large European or Eurosiberian (e.g. northern) ranges; it decreases the representation of Mediterranean endemics. The loss of grasslands and semi-open formations due to forest encroachment thus threatens exactly the species that should be the focus of conservation attention in the Mediterranean region, and innovative conservation actions to prevent ongoing forest encroachment are badly needed.

Phylogeny of Maculinea blues (Lepidoptera: Lycaenidae) based on morphological and ecological characters: evolution of parasitic myrmecophily.

A phylogeny of blue butterflies of the genus Maculinea and related genera (Lycaenidae) is proposed, based on 91 morphological and ecological characters. The resulting tree shows that: (1) Phengaris is a derived group nested within Maculinea; (2) the Maculinea-Phengaris clade is probably nested within Glaucopsyche; (3) there are three well supported groups within the Maculinea-Phengaris clade: (alcon group ((teleius group) (arion-Phengaris group))). Some species (M. alcon, M. arionides) appear to be non-monophyletic and require reclassification. The two alternative strategies of parasitic myrmecophily in the Maculinea-Phengaris clade, viz., "predatory" and "cuckoo", seem to be derived characters of the alcon group, and of the teleius and arion-Phengaris groups, respectively. The common ancestor of Maculinea used dorsal nectary organ secretions for ant attraction, while this trait was reduced in the ancestor of the alcon group and in M. nausithous (of the teleius group). The three recent Maculinea lineages utilize taxonomically diverse host plants, the asterid families Gentianaceae (alcon and arion-Phengaris groups), Lamiaceae (arion-Phengaris group), Campanulaceae (arion-Phengaris group), and the rosid family Rosaceae (teleius group).

Wolbachia infections mimic cryptic speciation in two parasitic butterfly species, Phengaris teleius and P. nausithous (Lepidoptera: Lycaenidae).

Deep mitochondrial divergence within species may result from cryptic speciation, from phylogeographic isolation or from endosymbiotic bacteria like Wolbachia that manipulate host reproduction. Phengaris butterflies are social parasites that spend most of their life in close relationship with ants. Previously, cryptic speciation has been hypothesised for two Phengaris species based on divergent mtDNA sequences. Since Phengaris species are highly endangered, the existence of cryptic species would have drastic consequences for conservation and management. We tested for cryptic speciation and alternative scenarios in P. teleius and P. nausithous based on a comprehensive sample across their Palaearctic ranges using COI gene sequences, nuclear microsatellites and tests for Wolbachia. In both species a deep mitochondrial split occurring 0.65-1.97 myrs ago was observed that did not correspond with microsatellite data but was concordant with Wolbachia infection. Haplotypes previously attributed to cryptic species were part of the Wolbachia-infected clades. In both species remaining phylogeographic structure was largely consistent between mitochondrial and nuclear genomes. In P. teleius several mitochondrial and nuclear groups were observed in East Asia while a single haplogroup and nuclear cluster prevailed across continental Eurasia. Neutrality tests suggested rapid demographic expansion into that area. In contrast, P. nausithous had several mitochondrial and nuclear groups in Europe, suggesting a complex phylogeographic history in the western part of the species range. We conclude that deep intraspecific divergences found in DNA barcode studies do not necessarily need to represent cryptic speciation but instead can be due to both infection by Wolbachia and phylogeographic structure.