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Butterflies use colour vision when searching for flowers. Unlike the trichromatic retinas of humans (blue, green and red cones; plus rods) and honeybees (ultraviolet, blue and green photoreceptors), butterfly retinas typically have six or more photoreceptor classes with distinct spectral sensitivities. The eyes of the Japanese yellow swallowtail (Papilio xuthus) contain ultraviolet, violet, blue, green, red and broad-band receptors, with each ommatidium housing nine photoreceptor cells in one of three fixed combinations. The Papilio eye is thus a random patchwork of three types of spectrally heterogeneous ommatidia. To determine whether Papilio use all of their receptors to see colours, we measured their ability to discriminate monochromatic lights of slightly different wavelengths. We found that Papilio can detect differences as small as 1–2 nm in three wavelength regions, rivalling human performance. We then used mathematical modelling to infer which photoreceptors are involved in wavelength discrimination. Our simulation indicated that the Papilio vision is tetrachromatic, employing the ultraviolet, blue, green and red receptors. The random array of three ommatidial types is a common feature in butterflies. To address the question of how the spectrally complex eyes of butterflies evolved, we studied their developmental process. We have found that the development of butterfly eyes shares its molecular logic with that of Drosophila: the three-way stochastic expression pattern of the transcription factor Spineless determines the fate of ommatidia, creating the random array in Papilio.

Several species of swallowtail butterflies (genus Papilio) are Batesian mimics that express multiple mimetic female forms, while the males are monomorphic and nonmimetic. The evolution of such sex-limited mimicry may involve sexual dimorphism arising first and mimicry subsequently. Such a stepwise scenario through a nonmimetic, sexually dimorphic stage has been proposed for two closely related sexually dimorphic species: Papilio phorcas, a nonmimetic species with two female forms, and Papilio dardanus, a female-limited polymorphic mimetic species. Their close relationship indicates that female-limited polymorphism could be a shared derived character of the two species. Here, we present a phylogenomic analysis of the dardanus group using 3964 nuclear loci and whole mitochondrial genomes, showing that they are not sister species and thus that the sexually dimorphic state has arisen independently in the two species. Nonhomology of the female polymorphism in both species is supported by population genetic analysis of engrailed, the presumed mimicry switch locus in P. dardanus. McDonald-Kreitman tests performed on SNPs in engrailed showed the signature of balancing selection in a polymorphic population of P. dardanus, but not in monomorphic populations, nor in the nonmimetic P. phorcas. Hence, the wing polymorphism does not balance polymorphisms in engrailed in P. phorcas. Equally, unlike in P. dardanus, none of the SNPs in P. phorcas engrailed were associated with either female morph. We conclude that sexual dimorphism due to female polymorphism evolved independently in both species from monomorphic, nonmimetic states. While sexual selection may drive male-female dimorphism in nonmimetic species, in mimetic Papilios, natural selection for protection from predators in females is an alternative route to sexual dimorphism.

In this study, the mitochondrial genome (mitogenome) sequence of Papilio memno is determined using next-generation sequencing. The entire mitogenome genome is determined to be 15,262 bp in length. ...

Telemiades Hübner, [1819] is currently comprised of 22 species and 14 subspecies distributed exclusively in the neotropics. The genus is defined by the morphological pattern of the uncus, apiculus, labial palpi and venation. However, some species have a uniform pattern of coloration making it difficult to establish their correct identity in some cases. The aims of the present study were to present a taxonomic revision of Telemiades Hübner, [1819] in order to provide a stable taxonomic hypothesis for the group, to describe the new species and to present diagnoses to help in species identification. According to the present study, Telemiades is now composed of 41 species. Fourteen new species are recognized and described. Three subspecies are here treated as valid species: T. marpesus (Hewitson, 1876) stat. rest., T. misitheus Mabille, 1888 stat. rest. and T. pekahia (Hewitson, 1868) stat. rest. Two synonyms are treated as valid species: T. insulsus (Draudt, 1922), stat. rev. and T. xantho Hayward, 1939 stat. rev. Five new synonyms are proposed: Plesioneura lamus Mabille, 1888 syn. nov. and Telemiades antiope tosca Evans, 1953 syn. nov. of T. antiope (Plötz, 1882), Echelatus punctatus Mabille & Boullet, 1917 syn. nov. of T. epicalus Hübner, [1819], Proteides nicola Plötz, 1882 syn. nov. of T. laogonus (Hewitson, 1876) and Eudamus phlius Plötz, 1881 syn. nov. of T. nicomedes (Möschler, 1879). To provide stability for existing names, six neotypes were designated for: Proteides amphion Geyer, 1832, Plesioneura compressa Möschler, 1877, Papilio avitus Stoll, 1781, Telegonus mygdon Möschler, 1877, Proteides aesopus Plötz, 1882 and Papilio corbulo Stoll, 1781; and 19 lectotypes for: Aethilla buffumi Weeks, 1906, Eudamus marpesus Hewitson, 1876, Telemiades misitheus Mabille, 1888, Pterygospidea pekahia Hewitson, 1868, Telemiades lurideolus Mabille, 1877, Eudamus laogonus Hewitson, 1876, Eudamus penidas Hewitson, 1867, Telemiades ceramina Herrich–Schäffer, 1869, Telemiades megalloides Schaus, 1902, Pythonides hoyti Weeks, 1906, Eudamus praestes Hewitson, 1876, Telemiades epicalus Hübner, [1819], Eudamus phasias Hewitson, 1867, Telemiades arcturus Herrich–Schäffer, 1869, Echelatus punctatus Mabille & Boullet, 1917, Eudamus centrites Hewitson, 1870, Telegonus gallius Mabille, 1888, Telegonus nicomedes Möschler, 1879, and Arteurotia meris Plötz, 1886.

Uncovering whether convergent adaptations share a genetic basis is consequential for understanding the evolution of phenotypic diversity. This information can help us understand the extent to which shared ancestry or independent evolution shape adaptive phenotypes. In this study, we first ask whether the same genes underlie polymorphic mimicry in Papilio swallowtail butterflies. By comparing signatures of genetic variation between polymorphic and monomorphic species, we then investigate how ancestral variation, hybridization, and independent evolution contributed to wing pattern diversity in this group. We report that a single gene, doublesex (dsx), controls mimicry across multiple taxa, but with species-specific patterns of genetic differentiation and linkage disequilibrium. In contrast to widespread examples of phenotypic evolution driven by introgression, our analyses reveal distinct mimicry alleles. We conclude that mimicry evolution in this group was likely facilitated by ancestral polymorphism resulting from early co-option of dsx as a mimicry locus, and that evolutionary turnover of dsx alleles may underlie the wing pattern diversity of extant polymorphic and monomorphic lineages., Polymorphic mimicry in Papilio swallowtail butterflies is thought to have had multiple independent origins. Here, the authors show that the gene doublesex controls mimicry across multiple species, but with distinct alleles that may have originated from an ancestral polymorphism.

Losaria coon (Fabricius, 1793) is currently comprised of ten subspecies, which were originally described under two names, Papilio coon and P. doubledayi before 1909, when they were combined as one species. The main difference between them is the colour of abdomen and hindwing subterminal spots&mdash, yellow in coon and red in doubledayi. Wing morphology, male and female genitalia, and molecular evidence (DNA barcodes) were analysed for multiple subspecies of L. coon and three other Losaria species&mdash, rhodifer, neptunus, and palu. Our molecular data support the separation of L. coon and L. doubledayi stat. rev. as two distinct species, with L. rhodifer positioned between them in phylogenetic analyses. Wing morphology and genitalic structures also confirm the molecular conclusions. Our findings divide L. coon into two species occupying different geographic ranges: with L. coon restricted to southern Sumatra, Java, and Bawean Island, while L. doubledayi occurs widely in regions from North India to northern Sumatra, including Hainan and Nicobar Islands. Hence, future conservation efforts must reassess the status and threat factors of the two species to form updated strategies.

This study investigated the cross‐national adaptation and implementation of Papilio, a German social–emotional learning programme, in Finnish early childhood education and care (ECEC) centres. Papilio is a developmentally focused, scientifically based intervention programme focused on preventing behavioural problems and fostering social–emotional competence in children aged 3–7. The aim of this study was to investigate and evaluate the cross‐national adaptation and cross‐cultural adaptation and implementation of Papilio in the Finnish ECEC context. Results from qualitative interviews with one Finnish Papilio trainer, 11 early childhood education (ECE) teachers, two ECE special education teachers and two nursery nurses are supplemented with teachers' and nursery nurses' (N = 75) questionnaire data. Qualitative thematic analysis revealed that cultural adaptations were necessary on four levels: accommodation of materials, adaptation of the contents of the materials, structure and delivery. The materials and training contents were culturally adapted, whereas the delivery of the intervention was adapted according to Finnish ECEC practices. The structural adaptation included discarding timeout, due to opposition by some educators. The educators were committed to implementing the programme as instructed and resolving the practical difficulties they encountered. Their motivation to implement Papilio increased as they observed improvements in the children's social–emotional competence during intervention. peerReviewed

We obtained and phylogenetically analyzed whole genome shotgun sequences of nearly all species from the tribe Emesidini Seraphim, Freitas & Kaminski, 2018 (Riodinidae) and representatives from other Riodinidae tribes. We see that the recently proposed genera Neoapodemia Trujano, 2018 and Plesioarida Trujano & García, 2018 are closely allied with Apodemia C. & R. Felder, [1865] and are better viewed as its subgenera, new status. Overall, Emesis Fabricius, 1807 and Apodemia (even after inclusion of the two subgenera) are so phylogenetically close that several species have been previously swapped between these two genera. New combinations are: Apodemia (Neoapodemia) zela (Butler, 1870), Apodemia (Neoapodemia) ares (Edwards, 1882), and Apodemia (Neoapodemia) arnacis (Stichel, 1928) (not Emesis); and Emesis phyciodoides (Barnes & Benjamin, 1924) (not Apodemia), assigned to each genus by their monophyly in genomic trees with the type species (TS) of the genus. Surprisingly, we find that Emesis emesia Hewitson, 1867 is not grouped with Emesis, but in addition to Apodemia forms a third lineage of similar rank, here named Curvie Grishin, gen. n. (TS: Symmachia emesia Hewitson, 1867). Furthermore, we partition Emesis into 6 subgenera (4 new): Emesis (TS: Hesperia ovidius Fabricius, 1793, a subjective junior synonym of Papilio cereus Linnaeus, 1767), Aphacitis Hübner, [1819] (TS: Papilio dyndima Cramer, [1780], a subjective junior synonym of Papilio lucinda Cramer, [1775]), Poeasia Grishin, subgen. n. (TS: Emesis poeas Godman, [1901]), Mandania Grishin, subgen. n. (TS: Papilio mandana Cramer, [1780]), Brimia Grishin, subgen. n. (TS: Emesis brimo Godman & Salvin, 1889), and Tenedia Grishin, subgen. n. (TS: Emesis tenedia C. & R. Felder, 1861). Next, genomic comparison of primary type specimens suggests new status for Emesis vimena Schaus, 1928 as a subspecies of Emesis brimo Godman & Salvin, 1889, Emesis adelpha Le Cerf, 1958 with E. a. vicaria Le Cerf, 1958 are subspecies of Emesis heteroclita Stichel, 1929, and Emesis tristis Stichel, 1929 is not a synonym of E. brimo vimena but of Emesis lupina Godman & Salvin, 1886. A new status of a species is given to the following taxa: Emesis furor A. Butler & H. Druce, 1872 (not a subspecies of E. mandana (Cramer, 1780)), Emesis melancholica Stichel, 1916 (not a subspecies of E. lupina Godman & Salvin, 1886), Emesis progne (Godman, 1903) (not a subspecies of E. brimo Godman & Salvin, 1889), and Emesis opaca Stichel, 1910 (not a synonym of E. lucinda (Cramer, 1775)). Emesis castigata diringeri Gallard 2008 is a subjective junior synonym of E. opaca, new status. Finally, Xanthosa Grishin, gen. n. (TS: Charmona xanthosa Stichel, 1910) is proposed for a sister lineage of Sertania Callaghan & Kaminski, 2017 and Befrostia Grishin, gen. n. (TS: Emesis elegia Stichel, 1929) is proposed for a clade without apparent phylogenetic affinities that we place in Befrostiini Grishin, trib. n. In conclusion, genomic data reveal a number of errors in the current classification of Emesidini and allow us to confidently reclassify the tribe partitioning it in three genera: Apodemia, Curvie gen. n. and Emesis.

Author(s): Grehan, John R | Abstract: Biogeographic patterns are reviewed for four widespread Southeast Asia butterfly groups in the superfamily Papilionoidea: Papilio subgenus Achillides Hubner, 1819 (Papilionidae), the birdwing butterflies (Papilionidae tribe Troidini), Genus Polyura Billberg, 1820 (Nymphalidae), and Genus Vanessa Fabricius, 1807 (Nymphalidae). The patterns of allopatry and sympatry are shown to be consistent with the vicariance of widespread ancestors with distributions including parts of Asia and Australasia, followed by secondary range expansion. Aspects of the distributions that are correlated with tectonic structures provide evidence of the age and origin of these butterflies in South-east Asia and Australasia. The transpacific affinities of the Troidini are consistent with a Pacific ancestry linked with former Cretaceous landscapes. The multi-island ranges of many of the butterfly species in Southeast Asia represent examples of metapopulation structure in which groups survive and persist in a region over long periods of time, even where individual islands are ephemeral.

A 3D image adds realism in viewing experience and can assist in simplifying the graphical displays. A Third dimension supplement to the input can improve pattern recognition, and can be used for 3D scene reconstruction and robot navigation. Recently popularity of 3D hardware is also increased which makes it a hot topic. The production of content as 3D is not matching with its need so there is scope of improvement of these 3D contents. Monocular cues give profundity data when seeing a scene with one eye. When a spectator moves, the evident relative movement of a few stationary articles against a foundation gives indicates about their relative separation. Depth estimation from monocular cues is a difficult task because single image lacks prior information like depth information, motion information etc. In Depth using scene features depth is estimated by exploring the features like shape, edges, color, texture and as well as an analysis of the environment of the scene that are of interest with respect to the target. Different objects have different hue and value and hence color is useful for depth estimation. Shape and texture provides disparity which is used to estimate depth. The main problem in converting a 2 dimensional to 3 dimensional images using single image is that it lacks information required for reconstruction in 3D data. While doing conversion by taking different cues or combination of multiple cues from scene conversion has been done e.g. structure form shape, motion, defocus etc. But such methods work for restricted scenarios not for global scenes. For instance, outdoor algorithms worked poor for indoor algorithms. Here we have implemented automatic conversion of 2 dimensional to 3 dimensional images using monocular image which can convert global images in visually comfortable 3D image.

This article documents 91 species-group names of Butterflies (Lepidoptera: Papilionoidea) for which either primary or secondary type material is present in the collections of the National Museums of Scotland, Edinburgh. The describers and the 76 nominal taxa they descibed are as follows: H. Druce (1846–1913) Euphaedra COOKSONI, Mycalesis HAROLDI ; H.H. Druce (1869–1922) Cyclopides COOKSONI, Spindasis KALLIMON ; H.J. Elwes (1846–1922) Parnassius delphius ssp. INFERNALIS ; J.C. Fabricius (1745–1808) Papilio COCALIA (“neotype”), Papilio MARDANIA (“neotype”); H. Fruhstorfer (1866–1922) Parnassius mnemosyne ssp. MELAINA ; D.R. Gifford (1918–1981) Leptomyrina HANDMANI, Alaena LAMBORNI, Deudorix (Virachola) MAGDA, Papilio ophidicephalus ssp. MKUWADZI ; J.B. Godart (1775–1825) Idea AGELIA, Danais ALCATHOE, Cethosia ALIPHERA, Danais ALOPIA, Danais BAUDINIANA, Papilio BITIAS, Argynnis BRIAREA, Heliconia CLEOBAEA, Danais CLEOPHILE, Danais CLEOTHERA, Heliconia CYRBIA, Pieris DOXO, Pieris EPICHARIS, Pieris ERIPHIA, Heliconia ETHILLA, Heliconia EUCLEA, Papilio EURYMAS, Pieris GIDICA, Argynnis HEGEMONE, Satyrus HYSIUS, Papilio IMERIUS, Acraea JANISCA, Pieris JOSEPHINA, Vanessa LAODORA, Papilio LEUCASPIS, Papilio LYCORAEUS, Vanessa LYTREA, Heliconia MEGARA, Heliconia MELPHIS, Libythea MYRRHA, Acraea OZOMENE, Pieris PHISADIA, Papilio POLYMETUS, Danais PROTHOE, Argynnis PYGMAEA, Pieris PYRO , Pieris SALACIA, Acraea SERVONA, Papilio TEMENES, Papilio TEREAS, Libythea TERENA, Biblis THADANA, Argynnis THAROSSA, Papilio TRIOPAS, Pieris VENILIA, Vanessa VULCANIA, Acraea ZETHEA, Acraea ZIDORA, Acraea ZOSTERIA ; C.W.N. Holmes (1916–2018) Bebearia paludicola ssp. BLANDI , Bebearia cocalioides ssp. HECQI, Bebearia orientis ssp. MALAWIENSIS, Bebearia PALUDICOLA ; E.G. Honrath (1837–1893) Parnassius bremeri ssp. GRAESERI, Parnassius nordmanni var. MINIMA ; W.J. Kaye (1875–1967) Hesperocharis LAMONTI ; F. Moore (1830–1907) Neptis ADARA , Abisara ANGULATA, Lebadea ATTENUATA, Ixias CITRINA, Euploea LIMBORGII, Papilio ONPAPE , Cirrhochroa SURYA ; W.F.H. Rosenberg & G. Talbot (1914) Dismorphia orise ssp. DENIGRATA. The secondary type material (number of species in brackets) originates from H. Carcasson (1), M. Cock (1), J. B. Godart (7), W. J. Kaye (1), O. Kudrna (3) and M. B. Usher (2).

Insect (epi)cuticular lipids characterize sex and species and often play an important role in mating behavior. We previously revealed that two black-colored swallowtail butterflies, Papilio polytes and Papilio protenor, show sexual dimorphism and species specificity in cuticular lipid composition and that P. polytes males use specific monoene components for mate discrimination. These findings suggest that their closely related species may have different profiles of cuticular lipids. We examined the cuticular lipid compositions of five Papilio species (P. bianor, P. maackii, P. helenus, P. macilentus, and P. memnon), closely related and sympatric to P. polytes and P. protenor, and discussed whether it is possible to discriminate between sexes, and between species based on their chemical profiles. The cuticular lipids consist mainly of C23–C31 aliphatic hydrocarbons, in which n-tricosane, n-heptacosane, and n-nonacosane are predominant. Several aliphatic ketones, aliphatic acids, and oxygenated terpenoids were also identified as major components shared by several species. There were no components exclusive to a particular species. Conspecific males and females shared most of the components but were mostly distinguishable based on their composition. Moreover, P. helenus males, P. polytes females, and P. protenor females had two different phenotypes of lipid composition. Although unrelated to the genetic lineage, the seven species were classified into four clusters based on their lipid profiles. The first cluster was composed of only P. memnon. The other six species were broadly classified into three clusters consisting of subclusters for each species: 1) P. polytes, P. helenus, P. macilentus, and several P. protenor females; 2) P. bianor and P. protenor; and 3) P. maackii and several P. helenus males. These results indicate that cuticular lipid profiles characterize the species and sex of the Papilio species and may be responsible for mate discrimination among them.

Lepidoptera is one of the largest orders of the class Insecta which includes butterflies and moths and is economically, biologically and aesthetically an important order. Chromosome structure and size are of great significance in karyological studies and have relevance to evolution, speciation and chromosome organization. Karyotypic studies in Lepidoptera has been a difficult task due to small dot- like chromosomes of similar sizes. The chromosome cytology of Indian Lepidoptera is very much limited. Chromosome studies in three species of butterflies ( Papilio polytes, Papilio demoleus, Papilio glaucus ) revealed 60 as diploid chromosome number.

Background and aims – Large-celled Eunotia species from tropical central Africa are documented in order to define morphological variation within species and to describe new or poorly documented taxa.Methods – Samples were collected from the Congo River between Kisangani and Bumba in the Democratic Republic of the Congo. These samples were prepared for diatom analysis and examined according to standard methods and documented using both light and scanning electron microscopy.Key results – A single population of Eunotia zygodon is used to demonstrate the high degree of change in gross cell morphology during its life cycle, changes previously observed in cultures of this genus. Cells of this taxon from each extremity of the reproductive cycle could be, and probably often were, placed in discrete taxa during sample analysis, especially as this taxon occurs rather infrequently. We also describe Eunotia leonardii as new to science. It has an undulate valve margin and is closely related to Eunotia serra and Eunotia georgii but differs in respect to overall cell size and the structure of the terminal raphe endings. The bi-undulate Eunotia papilio var. africana is raised to the rank of species and named Eunotia fuseyi as it is clearly a discrete taxon from E. papilio s. str.

Swallowtail diversity sampling in Okomu National Park was carried between July 2012 and June 2014 using butterfly nets and pollard methods (Pollard, 1977) along 1 km by 0.005 km of eight (8) transects in the various ranges (Igwonwan, Arakhuan, Julius Creek, Babui) of the park. Five hundred and fifty five (551) individuals grouped into two genera of twelve species were collected from the ranges of the park. Four rare species notably: Papilio sosia sosia, Papilio nierus nireus, Papilio garienus and Graphium angolanus (Family: Papilionidae) were observed in the park. Papilio garienus and Graphium angolanus recorded for the first time in the park. Most of the Graphium species confined in large congregations around areas with streams and mud puddling. The MOF transect had the highest diversity and was closely related to the CC transect. While the SHR transect had the lowest diversity (H) and was closely related to GRA transect. Swallowtail butterflies were found throughout the months of sampling and significantly higher (p

A variety of arthropods employ camouflage as bird droppings to avoid predation. We briefly review these cases and describe two new species of the buprestid genus Agrilus Curtis, 1825 that we hypothesize to be among them. Agrilus ornithocoprus Hespenheide andWestcott, new species, and Agrilus avifaecus Hespenheide and Westcott, new species, are described from specimens collected in Mexico. Agrilus albicollis (Waterhouse, 1887) from Jamaica also resembles a bird dropping.

International audience; The previously recognised closely related species Graphium (Pazala) mandarinus (Oberthür, 1879) and G. (P.) sichuanica(Koiwaya, 1993) are shown to comprise seven species as a result of both molecular and morphological analysis. Molecular dating analysis is also performed on the mandarinus group in order to investigate the divergence time of the taxa. Two HU ET AL. 402·Zootaxa 4441 (3) © 2018Magnolia Presstaxa, G. (P.) garhwalica (Katayama, 1988) stat. nov. and G. (P.) paphus (de Nicéville, 1886) stat. nov., are raised from sub-specific to specific status; G. (P.) hoeneanus Cotton & Hu nom. nov., stat. rev. is separated from sichuanica at species level; and two previously unrecognised new species, G. (P.) daiyuanae Hu, Zhang & Cotton sp. nov. and G. (P.) confucius Hu, Duan & Cotton sp. nov. are described from Vietnam and China respectively, the latter being sympatric with nominate G.(P.) mandarinus. The identity of the lectotype of G. (P.) mandarinus is confirmed and a lectotype is designated for the tax-on Papilio Glycerion Gray, 1831. A new subspecies of G. (P.) mandarinus is described from western Yunnan and northern Myanmar, G. (P.) mandarinusstilwelli Cotton & Hu ssp. nov.

The discovery of a new species of satyrine butterfly, Magneuptychia louisammour Benmesbah & Zacca, sp. n., that is phenotypically similar to a sympatric species widely known as Magneuptychia ocypete (Fabricius, 1776), led to a review of the taxonomic status of M. ocypete . A neotype for Papilio ocypete Fabricius, 1776 is designated and its redescription is provided. Neonympha sabina C. Felder & R. Felder, 1867 and Euptychia helle var. olivacea Aurivillius, 1929 are treated as junior subjective synonyms of P. ocypete and a lectotype is designated for each name . A neotype for the preoccupied name Papilio helle Cramer, 1779, and its replacement name Magneuptychia fugitiva Lamas, [1997], is also designated, and a redescription of this name and discussion of its taxonomic status are provided. We describe an additional phenotypically similar species, Magneuptychia kamel Benmesbah & Zacca , sp. n. from the western Amazon and raise the name Magneuptychia opima sheba Brevignon & Benmesbah, 2012 to species rank (stat. rev.) on the basis of morphological and DNA sequence evidence. We also discuss the difficulties regarding taxonomic and geographical delimitation in these complex species groups. Observations on Euptychiina behaviour are also provided. Finally, based on external characters and male genitalia, a proposed preliminary arrangement of two species groups is proposed, including other species of Magneuptychia Forster, 1964 and Cissia Doubleday, 1848 .

Spectrally opponent responses, that is, wavelength-dependent inversions of response polarity, have been observed at the level of photoreceptors in butterflies. As inter-photoreceptor connections have been found in the butterfly Papilio xuthus, and histamine is the only neurotransmitter so far identified in insect photoreceptors, we hypothesize that histaminergic sign-inverting synapses exist in the lamina between different spectral receptors as a mechanism for spectral opponency as in the medulla of Drosophila. Here, we localized two histamine-gated chloride channels, PxHCLA (Drosophila Ort homolog) and PxHCLB (Drosophila HisCl1 homolog), in the visual system of Papilio xuthus by using specific antisera. The antiPxHCLA labeling was associated with the membrane of nonphotoreceptor cells that are postsynaptic to photoreceptors, while the antiPxHCLB labeling overlapped with photoreceptor axons, indicating that PxHCLB is expressed at inter-photoreceptor synapses: PxHCLB is likely involved in producing spectral opponency at the first visual synapses. Color processing in Papilio may appear earlier than previously hypothesized in insect visual systems.

Batesian mimicry protects animals from predators when mimics resemble distasteful models. The female-limited Batesian mimicry in Papilio butterflies is controlled by a supergene locus switching mimetic and nonmimetic forms. In Papilio polytes, recent studies revealed that a highly diversified region (HDR) containing doublesex (dsx-HDR) constitutes the supergene with dimorphic alleles and is likely maintained by a chromosomal inversion. In the closely related Papilio memnon, which exhibits a similar mimicry polymorphism, we performed whole-genome sequence analyses in 11 butterflies, which revealed a nearly identical dsx-HDR containing three genes (dsx, Nach-like, and UXT) with dimorphic sequences strictly associated with the mimetic/nonmimetic phenotypes. In addition, expression of these genes, except that of Nach-like in female hind wings, showed differences correlated with phenotype. The dimorphic dsx-HDR in P. memnon is maintained without a chromosomal inversion, suggesting that a separate mechanism causes and maintains allelic divergence in these genes. More abundant accumulation of transposable elements and repetitive sequences in the dsx-HDR than in other genomic regions may contribute to the suppression of chromosomal recombination. Gene trees for Dsx, Nach-like, and UXT indicated that mimetic alleles evolved independently in the two Papilio species. These results suggest that the genomic region involving the above three genes has repeatedly diverged so that two allelic sequences of this region function as developmental switches for mimicry polymorphism in the two Papilio species. The supergene structures revealed here suggest that independent evolutionary processes with different genetic mechanisms have led to parallel evolution of similar female-limited polymorphisms underlying Batesian mimicry in Papilio butterflies.

Papilio alexanor Esper, 1800 —A species that has been rarely reported, but with several reliable records from coastal areas of Montenegro. It was first mentioned by Nicholl (1899) who collected two specimens from an unspecified site near Cetinje. Rebel, however, cited the small village of Boljevići near Skadar Lake as the locality, but it remains unclear as to how Rebel obtained the precise data regarding Nicholl’s exact location. The species was later reported by Carnelutti & Michieli (1958) and Švara et al. (2015) in the area around Stari Bar (old town of Bar), during the month of May. In addition, Kühnert (1971) observed a female of this species in the vicinity of Igalo at the beginning of June, while Mihljević (Sijarić 1991) and Bollino & Sala (2004) collected a male above Kotor. It seems that P. alexanor is widespread in the mountainous coastal region of Montenegro, although it remains a very local species, and it is rarely observed. The host plant in Montenegro is probably Opopanax chironium Koch, as this plant is reported as the sole host in neighbouring Croatia (Verovnik & Švara 2016)., Published as part of Franeta, Filip, 2018, Checklist of the butterflies (Lepidoptera: Papilionoidea) of Montenegro, pp. 128-148 in Zootaxa 4392 (1) on page 136, DOI: 10.11646/zootaxa.4392.1.6, http://zenodo.org/record/1195491, {"references":["Nicholl, M. de la B. (1899) Butterfly hunting in Dalmatia, Montenegro, Bosnia and Herzegovina. Entomologists Record, 11, 1 - 8.","Carnelutti, J. & Michieli, S. (1958) i. Beitrag zur Lepidopterenfauna der Crna Gora. Fragmenta Balcanica, 10 (2), 67 - 81.","Kuhnert, H. (1971) Die Tagfalterbeobachtungen im Gebiet von Igalo (Montenegro). Entomologische Zeitschrift Frankfurt am Main, 81 (8), 73 - 93.","Sijaric, R. (1991) Katalog naucne zbirke Lepidoptera (Insecta) donatora Bore Mihljevica iz Sarajeva. Glasnik Zemaljskog muzeja Bosne i Hercegovine, Prirodne Nauke, Nova serija 30, 169 - 360.","Bollino, M. & Sala, G. (2004) Synopsis of Papilio alexanor Esper, 1799 (An updated review of its taxonomy and biology). Natura edizioni scientifiche, Bologna, 64 pp."]}