Dissecting supergene: evidence for recombination suppression among multiple functional loci within inversions.

Supergenes are genetic architectures associated with discrete and concerted variation in multiple traits. It has long been suggested that supergenes control these complex polymorphisms by suppressing recombination between set of coadapted genes. However, because recombination suppression hinders the dissociation of the individual effects of genes within supergenes, there is still little evidence that supergenes evolve by tightening linkage between coadapted genes. Here, combining an landmark-free phenotyping algorithm with multivariate genome wide association studies, we dissected the genetic basis of wing pattern variation in the butterfly Heliconius numata. We showed that the supergene controlling the striking wing-pattern polymorphism displayed by this species contains many independent loci associated with different features of wing patterns. The three chromosomal inversions of this supergene suppress recombination between these loci, supporting the hypothesis that they may have evolved because they captured beneficial combinations of alleles. Some of these loci are associated with colour variations only in morphs controlled by inversions, indicating that they were recruited after the formation of these inversions. Our study shows that supergenes and clusters of adaptive loci in general may form via the evolution of chromosomal rearrangements suppressing recombination between co-adapted loci but also via the subsequent recruitment of linked adaptive mutations.

FIRST RECORD OF THE GENUS NIPPOPTILIA MATSUMURA (LEPIDOPTERA: PTEROPHORIDAE) FROM RUSSIA

A plume moth Nippoptilia regula (Meyrick, 1906) is found in the southern part of Primorsky krai. The specimens examined are characterized by typical for this species wing pattern, male and female genitalia. The genus Nippoptilia Matsumura 1931 is recorded from Russia for the first time.

Taxonomic notes on the genus Elaphromyia Bigot (Diptera: Tephritidae: Tephritinae: Pliomelaenini) in India, with description of a new species

A new species of Elaphromyia, E. juncta David, Hancock & Sachin, sp. n. is described from India. It can be differentiated from the morphologically similar E. siva Frey and E. pterocallaeformis (Bezzi) by the wing pattern, epandrial characters, morphology of spicules on the eversible membrane and spermathecal shape. Elaphromyia siva Frey and E. yunnanensis Wang are recorded for the first time from India. Records of E. pterocallaeformis (Bezzi) from southern India are regarded as misidentifications. A key to the 7 known non-African species is included.

Unraveling a complex problem: Dichrorampha velata sp. nov., a new species from the Alps hitherto confounded with D. alpestrana ([Zeller], 1843) sp. rev. = D. montanana (Duponchel, 1843) syn. nov. (Lepidoptera, Tortricidae)

Analysis of wing pattern, genital morphology and results of DNA barcoding indicates that the name Dichrorampha montanana sensu auct. actually comprises two species. D. alpestrana ([Zeller], 1843) sp. rev. is considered as senior synonym of D. montanana (Duponchel, 1843) syn. nov., and a lectotype is designated for the latter name to fix the identity. After thorough search for possible synonyms, Dichrorampha velatasp. nov. is described and differentiated morphologically and with DNA barcodes from D. alpestrana and adults and genitalia of both species are figured. Dichrorampha velatasp. nov. is restricted to the European Alps and adjacent regions whereas D. alpestrana is more widespread with likely arctic-alpine disjunction and records from the Alps and the northern part of Great Britain.

Sibling species as a result of microevolution (a case study of Lepidoptera)

This article is devoted to the problems of diagnosis and taxonomy of sibling species in Lepidoptera of different taxonomic groups. The most important advances in understanding the reality of species include: species are real and objectively exist; each species has specific spatial characteristics and its own biological time, which may not coincide with the physical one. The main criteria for the reality of a species include the following: a certain stability in space and time, isolation from the surrounding world, a certain opposition to the environment; material continuity over time; a certain degree of indivisibility; the presence of special, distinctive properties in relation to other similar systems; the presence of a certain number of degrees of freedom in relation to higher taxa; the presence of a given set of individuals of a lower order, a certain hierarchy of individuals; continuity in space and time. Through the construction of a natural system of phenotypes of the wing pattern, it is possible to build a system of groups of populations of a species and create a population taxonomy, while the phenotype should act as a reliable diagnostic feature of a specific population or groups of populations. In the landscape complex of Lepidoptera populations of a particular geographic region, as a rule, only one of the available forms is the most often found, which is also the most often collected and described as typical. All other forms are found in populations with a lower frequency, so they can be described as atypical, up to giving them the status of new species. This applies primarily to European populations of Lepidoptera, individuals of which were once accepted by taxonomists as typical and therefore received the status of nominative species and subspecies. Individuals from populations at the border of ranges, for example, from mountainous areas, are most often described as new taxa. Quite often, polytypical Lepidoptera species have a wide phenotypic diversity of the wing pattern, which makes rare forms in central populations often found in peripheral populations of the range. Therefore, the species of Lepidoptera should be considered in the light of a biological concept. Based on an integrated approach to studying the phenotypic variability of the wing pattern, it is possible to carry out taxonomic studies of Lepidoptera populations and the separation of sibling species.

A new species of the genus Teratozephyrus Sibatani, 1946 from China (Lepidoptera, Lycaenidae, Theclinae)

A new species, T. yaolihuoi Huang, Jiang & Song sp. nov. of the genus Teratozephyrus Sibatani, 1946 is described from southeastern China. Among the other representatives of the genus, the new species resembles only T. hinomaru Fujioka, 1994 from southwestern China, from which the new species can be separated easily by examining details in both wing pattern and the male genitalia. Wing patterns and genitalia of the new species and T. hinomaru are illustrated and compared.

The Taxa of the Hyponephele lycaon – H. lupina Species Complex (Lepidoptera, Nymphalidae, Satyrinae): Deep DNA Barcode Divergence despite Morphological Similarity

The genus Hyponephele includes about 40 species distributed throughout the southern part of the Palaearctic area. Within this genus, the taxa of the H. lycaon – H. lupina species complex are similar with respect to the wing pattern and genitalia structure. Here we revise this group using analysis of butterfly morphology, DNA barcodes, and study of the type material. We show that, with a few exceptions, the species in this group are allopatric in distribution. Allopatry in combination with phenotypic similarity may be theoretically interpreted as evidence for the conspecifity of these taxa. Here we falsify this hypothesis by using DNA barcode analysis. We show that the species of this complex are genetically very distant and cannot be combined together as a polytypic species. We also demonstrate that H. lupina consists of two deeply diverged allopatric clades, H. lupina s. s. and H. mauritanica comb. & stat. nov. The barcode p-distance between these taxa (3.4-4.9%) is significantly higher than the generally accepted 'standard' minimum interspecific divergence (2.0-3.0% ) threshold. These two clades can also be distinguished by the color of the upperside of the wing in males (brown with conspicuous golden reflection in H. lupina ; dark brown without golden reflection in H. mauritanica) and by details in male genitalia and male androconia structures. Syntypes of Hyponephele sifanica, H. cheena cheena, H. cheena iskander, and H. cheena kashmirica are studied and figured.

A new species of Dichagyris Lederer, 1857 from China (Lepidoptera, Noctuidae)

Dichagyris is a diverse genus with a Holarctic distribution. Only a few taxa of the subgenus Albocosta Fibiger & Lafontaine, 1997 occur in the northwesternmost part of the Oriental region. The genus is most diverse in Turkey, Iran and the Central Asiatic high mountains. However, there is a distinct Sino-Tibetan group of species, possibly representing an undescribed subgenus. However, this idea is provisional and awaits revision of the genus. For present purposes, we assign these species to the subgenus Dichagyris. They share the following putative synapomorphies: black, dark brown or dark greyish forewing ground colour (with one exception), with obscure, reduced wing pattern and a broad-based, short, conical harpe in the male genitalia. Species of this group can be subdivided to two species-groups. In the astigmata-group which was mentioned for the first time by Hreblay et al. (1998) but without description or diagnosis from the minuta-group, the vesica is elongate and tubular (males), the appendix bursae and corpus bursae are long, sack–like, almost equal in size (females). In the minuta-group, the vesica is more ample than in the astigmata-group, and coiled, and the appendix bursae is globular and much shorter than the corpus bursae. The astigmata-group includes Dichagyris astigmata (Hampson, 1906), D. gansuensis Hreblay & Ronkay, 1998, D. geochroides (Boursin, 1948) and D. vargazoli (Gyulai & Ronkay, 2001). The minuta-group comprises D. minuta Hreblay & Plante, 1998 and D. kormos Gyulai & Ronkay, 2001. During a research expedition in Sichuan in 2019, the existence of a third species in the minuta-group was recognized, which is described here. In addition, the female and its genitalia of D. gansuensis, which was described by Hreblay & Ronkay (1998) based on a single male, are illustrated here for the first time.

Minoa lutea Schwingenschuss, 1954 (Lepidoptera: Geometridae: Larentiinae) recognized as bona species

The species Minoa murinata (Scopoli, 1763) sensu lato is examined throughout its distribution range. Specimens from central Europe (Germany, Italy, France) are compared with those from Serbia, Croatia, Slovenia and newly collected specimens from eastern Turkey, Armenia, Georgia and Russia. The study is based on a combination of behavioural observations, morphological characters (size, wing coloration, structure of male and female genitalia) as well as genetic data (DNA barcoding). The taxon Minoa murinata var. monochroaria Herrich-Schäffer, 1848 is downgraded from subspecies rank to synonymy of M. murinata. Morphological study of the populations from eastern Turkey, Armenia, Georgia and Russia confirm the taxon Minoa murinata f./ssp. lutea Schwingenschuss, 1954 as a bona species. It is herewith upgraded from synonymy of M. murinata to species level. The results of DNA barcoding are discussed. Wing pattern, male and female genitalia of both species are illustrated.