Wing Pattern Research Articles

A wing-flapping robot with a bio-inspired folding mechanism derived from the beetle's hind wing.

When the beetle lands on the target, the hind wings fold regularly to form smaller wing packages and are hidden on the ventral side of the elytra due to the interaction between the elytra and abdomen. Its complex folding pattern is attributed to the flexibility of the hind wings, the super-elasticity of the folding joints, and the special geometric morphology of the veins. The corrugation and folding pattern of the hind wings can provide new insights for the design of folding anti-collision mechanisms and the improvement of aerodynamic performance of ornithopter. This paper first proposes a beetle-type ornithopter with foldable wings based on the folding mechanism and kinematic characteristics of the beetle's hind wings. Subsequently, a series of numerical simulations were conducted on flapping wing robot to explore its flapping kinematics, folding stability, structural stiffness. Finally, the force generation of flapping wings was tested on the fabricated prototype.

Dorsoventral comparison of intraspecific variation in the butterfly wing pattern using a convolutional neural network.

Butterfly wing patterns exhibit notable differences between the dorsal and ventral surfaces, and morphological analyses of them have provided insights into the ecological and behavioural characteristics of wing patterns. Conventional methods for dorsoventral comparisons are constrained by the need for homologous patches or shared features between two surfaces, limiting their applicability across species. We used a convolutional neural network (CNN)-based analysis, which can compare images of the two surfaces without focusing on homologous patches or features, to detect dorsoventral bias in two types of intraspecific variation: sexual dimorphism and mimetic polymorphism. Using specimen images of 29 species, we first showed that the level of sexual dimorphism calculated by CNN-based analysis corresponded well with traditional assessments of sexual dissimilarity, demonstrating the validity of the method. Dorsal biases were widely detected in sexual dimorphism, suggesting that the conventional hypothesis of dorsally biased sexual selection can be supported in a broader range of species. In contrast, mimetic polymorphism showed no such bias, indicating the importance of both surfaces in mimicry. Our study demonstrates the potential versatility of CNN in comparing wing patterns between the two surfaces, while elucidating the relationship between dorsoventrally different selections and dorsoventral biases in intraspecific variations.

First record of Atypophthalmus umbratus (Diptera, Limoniidae) from Central Europe, a species introduced accidentally throughout global trade of exotic plants

Here we report the first data of the exotic Atypophthalmus umbratus (de Meijere, 1911) from two Central European countries, Romania and Hungary. This is the first time that the presence of an introduced exotic species of Limoniidae (Diptera) has been reported in the area. The above-mentioned species was first observed in Cluj, Romania, on tropical plant specimens (e.g. Alocasia x mortfontanensis ‘Polly’), which we bought from a large store selling tropical plants, from which both male and female specimens were collected. After that, checking some citizen science platforms of entomological interest, we also noticed record of the species in Hungary, based on their conspicuous wing pattern and general habitus. The presence of this accidentally introduced species far from its native tropical environment highlights the growing intensity of the global exotic plant market and the importance of citizen science in early warning systems for biological invasions.

Description of a new subspecies of Lethe gemina Leech, 1891 from northern Guangdong of China (Lepidoptera, Nymphalidae, Satyrinae)

Based upon morphological and molecular evidences, a new subspecies of Lethe gemina Leech, 1891, namely L. gemina bicolor subsp. nov., is described from northern Guangdong Province of China. The adults and genitalia of both sexes are illustrated. Differences in wing patterns of this new taxon and the seven known subspecies of L. gemina are discussed.

Contributions of the Dachsous intracellular domain to Dachsous-Fat signaling.

The protocadherins Fat and Dachsous regulate organ growth, shape, patterning, and planar cell polarity. Although Dachsous and Fat have been described as ligand and receptor, respectively, in a signal transduction pathway, there is also evidence for bidirectional signaling. Here, we assess signaling downstream of Dachsous through analysis of its intracellular domain. Genomic deletions of conserved sequences within dachsous identified regions of the intracellular domain that contribute to Dachsous activity. Deletion of the A motif increased Dachsous protein levels and decreased wing size. Deletion of the D motif decreased Dachsous levels at cell membranes, increased wing size, and disrupted wing, leg and hindgut patterning and planar cell polarity. Co-immunoprecipitation experiments established that the D motif is necessary and sufficient for association of Dachsous with key partners, including Lowfat, Dachs, Spiny-legs, Fat and MyoID. Subdivision of the D motif identified distinct regions that preferentially contribute to different Dachsous activities. Our results identify motifs that are essential for Dachsous function and are consistent with the hypothesis that the key function of Dachsous is regulation of Fat.

Description of a new species of the genus Aeromachus Nicéville (Lepidoptera, Hesperiidae, Hesperiinae) from Southwestern China.

A new species of the genus Aeromachus, A. angustifascia sp. nov. is described from western Guizhou Province, southwestern China. The new species is superficially similar to A. catocyanea (Mabille, 1876) but can be distinguished from the latter by the details in wing pattern and male genitalia as well as COI genetic difference. The adults and genitalia of the aforementioned taxa are illustrated.

A new species of Plutella Schrank (Lepidoptera, Plutellidae) from the Andes of northern Chile.

The moth genus Plutella Schrank, 1802 (Lepidoptera, Plutellidae) includes 26 described species. In addition to the diamondback moth, P.xylostella (Linnaeus, 1758), which is an important and widely distributed pest of cruciferous crops, only two species have been previously recorded in Chile, both with distribution ranges restricted to the southern temperate rainforests. Plutellacopaquillaensis sp. nov. is described and illustrated, based on adults reared from larvae collected on Neuontobotryslanata (Walp.) Al-Shehbaz (Brassicaceae) in the arid highlands of the Andes of northern Chile. The new species resembles P.xylostella in wing pattern, but clearly differs in genitalia morphology.

Robust biomimetic strain sensor based on butterfly wing-derived skeleton structure

A biomimetic strain sensor was designed and constructed based on Ir nanoparticles-modified multi-wall carbon nanotubes (Ir NPs@MWCNTs) and parallel Pt layer/dragon skin with carbonized butterfly wing patterns. This sensor exhibits high gauge factor (∼515.4), extensive tensile range (0%–96%), and swift response (∼300 ms), especially remarkable stability up to 60 000 cycles. The work mechanism has been proposed based on the experimental test and finite-element method. Some important applications such as human motion and micro-expression recognition have been confirmed using 3 × 3 flexible biomimetic sensor array.

Discovery of a new gall-inducing species, Aciurinaluminaria (Insecta, Diptera, Tephritidae) via multi-trait integrative taxonomy.

Integrative taxonomic practices that combine multiple lines of evidence for species delimitation greatly improve our understanding of intra- and inter-species variation and biodiversity. However, extended phenotypes remain underutilized despite their potential as a species-specific set of extracorporeal morphological and life history traits. Primarily relying on variations in wing patterns has caused taxonomic confusion in the genus Aciurina, which are gall-inducing flies on Asteraceae plants in western North America. However, species display distinct gall morphologies that can be crucial for species identification. Here we investigate a unique gall morphotype in New Mexico and Colorado that was previously described as a variant of that induced by Aciurinabigeloviae (Cockerell, 1890). Our analysis has discovered several consistent features that distinguish it from galls of A.bigeloviae. A comprehensive description of Aciurinaluminaria Baine, sp. nov. and its gall is provided through integrative taxonomic study of gall morphology, host plant ecology, wing morphometrics, and reduced-representation genome sequencing.

The ivory lncRNA regulates seasonal color patterns in buckeye butterflies

Long noncoding RNAs (lncRNAs) are transcribed elements increasingly recognized for their roles in regulating gene expression. Thus far, however, we have little understanding of how lncRNAs contribute to evolution and adaptation. Here, we show that a conserved lncRNA, ivory, is an important color patterning gene in the buckeye butterfly Junonia coenia. ivory overlaps with cortex, a locus linked to multiple cases of crypsis and mimicry in Lepidoptera. Along with a companion paper by Livraghi et al., we argue that ivory, not cortex, is the color pattern gene of interest at this locus. In J. coenia, a cluster of cis-regulatory elements (CREs) in the first intron of ivory are genetically associated with natural variation in seasonal color pattern plasticity, and targeted deletions of these CREs phenocopy seasonal phenotypes. Deletions of different ivory CREs produce other distinct phenotypes as well, including loss of melanic eyespot rings, and positive and negative changes in overall wing pigmentation. We show that the color pattern transcription factors Spineless, Bric-a-brac, and Ftz-f1 bind to the ivory promoter during wing pattern development, suggesting that they directly regulate ivory. This case study demonstrates how cis-regulation of a single noncoding RNA can exert diverse and nuanced effects on the evolution and development of color patterns, including modulating seasonally plastic color patterns.

Harnessing Nature's ingenuity to engineer butterfly-wing-inspired photoactive nanofiber patches for advanced postoperative tumor treatment

Postoperative tumor treatment necessitates a delicate balance between eliminating residual tumor cells and promoting surgical wound healing. Addressing this challenge, we harness the innovation and elegance of nature's ingenuity to develop a butterfly-wing-inspired photoactive nanofiber patch (WingPatch), aimed at advancing postoperative care. WingPatch is fabricated using a sophisticated combination of electrostatic spinning and spraying techniques, incorporating black rice powder (BRP) and konjac glucomannan (KGM) into a corn-derived polylactic acid (PLA) nanofiber matrix. This fabrication process yields a paclitaxel-infused porous nanofiber architecture that mirrors the delicate patterns of butterfly wings. Meanwhile, all-natural composites have been selected for their strategic roles in postoperative recovery. BRP offers the dual benefits of photothermal therapy and antibacterial properties, while KGM enhances both antibacterial effectiveness and tissue regeneration. Responsive to near-infrared light, WingPatch ensures robust tissue adhesion and initiates combined photothermal and chemotherapeutic actions to effectively destroy residual tumor cells. Crucially, it simultaneously prevents infections and promotes wound healing throughout the treatment process. Its effectiveness has been confirmed by animal studies, and WingPatch significantly improves treatment outcomes in both breast and liver tumor models. Thus, WingPatch exemplifies our dedication to leveraging natural world's intricate patterns and inventiveness to propel postoperative care forward.

A long noncoding RNA at the cortex locus controls adaptive coloration in butterflies

Evolutionary variation in the wing pigmentation of butterflies and moths offers striking examples of adaptation by crypsis and mimicry. The cortex locus has been independently mapped as the locus controlling color polymorphisms in 15 lepidopteran species, suggesting that it acts as a genomic hotspot for the diversification of wing patterns, but functional validation through protein-coding knockouts has proven difficult to obtain. Our study unveils the role of a long noncoding RNA (lncRNA) which we name ivory, transcribed from the cortex locus, in modulating color patterning in butterflies. Strikingly, ivory expression prefigures most melanic patterns during pupal development, suggesting an early developmental role in specifying scale identity. To test this, we generated CRISPR mosaic knock-outs in five nymphalid butterfly species and show that ivory mutagenesis yields transformations of dark pigmented scales into white or light-colored scales. Genotyping of Vanessa cardui germline mutants associates these phenotypes to small on-target deletions at the conserved first exon of ivory. In contrast, cortex germline mutant butterflies with confirmed null alleles lack any wing phenotype and exclude a color patterning role for this adjacent gene. Overall, these results show that a lncRNA gene acts as a master switch of color pattern specification and played key roles in the adaptive diversification of wing patterns in butterflies.

Integrative description of Thaeides ramoni sp. nov. (Lepidoptera: Lycaenidae), a sympatric sibling species of Thaeides theia (Hewitson, 1870) found in the Sierra Nevada de Santa Marta, Colombia.

We present evidence from DNA barcodes, wing pattern and distribution to support the hypothesis that an entity flying in sympatry with Thaeides theia in the Sierra Nevada de Santa Marta, is an undescribed biological species named herein as Thaides ramoni Prieto, sp. nov. Adult specimens and genital structures are illustrated for both species along with molecular and morphological diagnostic characters. In addition, we present some considerations about the taxonomy of the species in the Thaeides muela group.

Evolution of Opsin Genes in Caddisflies (Insecta: Trichoptera).

Insects have evolved complex and diverse visual systems in which light-sensing protein molecules called "opsins" couple with a chromophore to form photopigments. Insect photopigments group into three major gene families based on wavelength sensitivity: long wavelength (LW), short wavelength (SW), and ultraviolet wavelength (UV). In this study, we identified 123 opsin sequences from whole-genome assemblies across 25 caddisfly species (Insecta: Trichoptera). We discovered the LW opsins have the most diversity across species and form two separate clades in the opsin gene tree. Conversely, we observed a loss of the SW opsin in half of the trichopteran species in this study, which might be associated with the fact that caddisflies are active during low-light conditions. Lastly, we found a single copy of the UV opsin in all the species in this study, with one exception: Athripsodes cinereus has two copies of the UV opsin and resides within a clade of caddisflies with colorful wing patterns.

First record of the genus Sergeya Ponomarenko, 2008 (Lepidoptera, Gelechiidae) from the Afrotropical and Oriental regions, with description of ten new species.

Genus Sergeya Ponomarenko, 2008 is recorded here for the first time from the Afrotropical and Oriental regions. Ten species are described as new: S. hackeri sp. nov. (Yemen), S. olei sp. nov. (Thailand), S. chuii sp. nov. (DR Congo), S. bunsoensis sp. nov. (Ghana), S. palescens sp. nov. (Uganda, South Africa), S. malawica sp. nov. (Malawi), S. korongotaji sp. nov. (Uganda), S. lobata sp. nov. (South Africa, Uganda, Ethiopia), S. davidi sp. nov. (Kenya), S. harambee sp. nov. (Kenya). The genus is redescribed, and its differences from most related Afrotropical genus Lanceopenna Janse, 1950, as well as Oriental Neolitinia Ponomarenko, Omelko & Omelko, 2021 and Parateleiopsis Ponomarenko, Omelko & Omelko, 2021 are discussed. Additionally, Lanceopenna prominula (Meyrick, 1913) (Aristotelia) is transferred from the genus Lanceopenna to Sergeya: S. prominula (Meyrick, 1913) comb. nov. and is recorded for the first time from Zimbabwe. All Afrotropical species of Sergeya are diagnosed, and an identification key to all species of the genus is provided. Wing pattern and genitalia of all species are illustrated. A lectotype is designated for Aristotelia prominula Meyrick, 1913. An updated check-list of species of Sergeya is provided.

Evolutionary patterns and functional effects of 3D chromatin structures in butterflies with extensive genome rearrangements

Chromosome rearrangements may distort 3D chromatin architectures and thus change gene regulation, yet how 3D chromatin structures evolve in insects is largely unknown. Here, we obtain chromosome-level genomes for four butterfly species, Graphium cloanthus, Graphium sarpedon, Graphium eurypylus with 2n = 30, 40, and 60, respectively, and Papilio bianor with 2n = 60. Together with large-scale Hi-C data, we find that inter-chromosome rearrangements very rarely disrupted the pre-existing 3D chromatin structure of ancestral chromosomes. However, some intra-chromosome rearrangements changed 3D chromatin structures compared to the ancestral configuration. We find that new TADs and subTADs have emerged across the rearrangement sites where their adjacent compartments exhibit uniform types. Two intra-chromosome rearrangements altered Rel and lft regulation, potentially contributing to wing patterning differentiation and host plant choice. Notably, butterflies exhibited chromatin loops between Hox gene cluster ANT-C and BX-C, unlike Drosophila. Our CRISPR-Cas9 experiments in butterflies confirm that knocking out the CTCF binding site of the loops in BX-C affected the phenotypes regulated by Antp in ANT-C, resulting in legless larva. Our results reveal evolutionary patterns of insect 3D chromatin structures and provide evidence that 3D chromatin structure changes can play important roles in the evolution of traits.

Optix and cortex/ivory/mir-193 again: the repeated use of two mimicry hotspot loci.

The extent to which evolution is repeatable has been a debated topic among evolutionary biologists. Although rewinding the tape of life perhaps would not lead to the same outcome every time, repeated evolution of analogous genes for similar functions has been extensively reported. Wing phenotypes of butterflies and moths have provided a wealth of examples of gene re-use, with certain 'hotspot loci' controlling wing patterns across diverse taxa. Here, we present an example of convergent evolution in the molecular genetic basis of Batesian wing mimicry in two Hypolimnas butterfly species. We show that mimicry is controlled by variation near cortex/ivory/mir-193, a known butterfly hotspot locus. By dissecting the genetic architecture of mimicry in Hypolimnas misippus and Hypolimnas bolina, we present evidence that distinct non-coding regions control the development of white pattern elements in the forewing and hindwing of the two species, suggesting independent evolution, and that no structural variation is found at the locus. Finally, we also show that orange coloration in H. bolina is associated with optix, a well-known patterning gene. Overall, our study once again implicates variation near the hotspot loci cortex/ivory/mir-193 and optix in butterfly wing mimicry and thereby highlights the repeatability of adaptive evolution.

New data on distribution of Phoenicurusia transcaucasicus (Miller, 1923) (Lepidoptera, Lycaenidae) with description of a new subspecies from Iran.

Distribution of Phoenicurusia transcaucasicus (Miller, 1923) in Iran and neighbouring territories is clarified based on analysis of DNA barcodes, the male genitalia and wing pattern of adults. Our study revealed the widespread distribution of Ph. transcaucasicus throughout northern, northeastern and central Iran. Based on integrative analysis, a new subspecies, Ph. transcaucasicus flamma ssp. n., is described from the Zagros Mountains, the Central Iranian Range and the Alborz Mountains. Compared to the nominotypical subspecies, the new subspecies differs in well-developed orange pattern both of dorsal and ventral sides of the wings, shape of the valva in the male genitalia and distinct COI haplotypes. Additionally, distinct phylogenetic lineage of Ph. transcaucasicus is reported from Kerman Province. Diagnostic characters of the species of Ph. phoenicurus (Lederer, 1870) group of Iran are illustrated.

Mating preferences act independently on different elements of visual signals in Heliconius butterflies

Abstract Mating cues are often comprised of several elements, which can act independently, or in concert to attract a suitable partner. Individual elements may also function in other contexts, such as anti-predator defense or camouflage. In Heliconius butterflies, wing patterns comprise several individual color pattern elements, which advertise the butterflies’ toxicity to predators. These wing patterns are also mating cues and males predominantly court females that possess the same wing pattern as their own. However, it is not known whether male preference is based on the full wing pattern or only individual pattern elements. We compared preferences of male H. erato lativitta between female models with the full wing pattern and those with some pattern elements removed. We found no differences in preference between the full wing pattern model and a model with pattern elements removed, indicating that the complete composition of all elements is not essential to the mating signal. Wing pattern preferences also contribute to pre-mating isolation between two other Heliconius taxa, H. erato cyrbia, and H. himera; therefore, we next compared preferences for the same models in these species. H. erato cyrbia and H. himera strongly differed in preferences for the models, potentially providing a mechanism for how pre-mating isolation acts between these species. These findings suggest that contrasting levels of selective constraint act on elements across the wing pattern.

Schistophleps kendricki sp. nov. (Lepidoptera: Erebidae: Arctiinae) from Bangladesh with an overview of the genus.

Schistophleps Hampson, 1891 is one of the most frequently occurring genera of tiger moths, widely distributed in the Oriental and Australian regions. However, a lion's share of the species belonging to this genus are poorly known except the original descriptions. In this article, we provide a brief overview of all known species currently assigning to this genus. We divide the species into albida, bicolora, bipuncta, flavia, fulvia, hyalina, and subtilis species groups on the basis of wing pattern elements along with geographic distributions, and four highly obscure species are discussed as a miscellaneous group. Among the species, Schistophleps bipuncta Hampson, 1891 is the most frequently occurring taxon having a distribution stretching all over the South East Asia. However, the variable wing maculations and other features, reflected through the observations on various citizen science platforms from different regions of the Indian subcontinent indicate to a sibling species yet to discover. Herein, we describe the species as Schistophleps kendricki Rayhan, Bucsek, & Jahan sp. nov., from Chittagong, Bangladesh that is similar to the Schistophleps bipuncta species group in wing maculation, but markedly differs from all of its closely allied congeners in the genitalia configurations in having characteristic modification of uncus. Besides, a comparison of observations on different citizen science platforms from the Indian subcontinent is provided that suggests the possible distribution of this new species throughout the Bengal including West Bengal, India and Bangladesh.