Butterfly Pattern Research Articles

Do single Pollard transects represent the local butterfly community? A case study from the Spitzberg near Tübingen, Germany

Abstract Information on butterfly population size and distribution is a key tool for verifying species conservation status. Such data are rare and tend to be available on local rather than on regional scales. One important data source are monitoring schemes that have been implemented in several countries, e.g. the German butterfly monitoring scheme (TMD). The main goal of the present study was to examine how representative single transects of the TMD are able to identify local or regional diversity patterns of butterflies. Total butterfly population size, density of individuals and species diversity pattern of a 624 ha large study area (Spitzberg) were recorded by intensive surveys. Almost all butterfly species present at the Spitzberg were recorded by the TMD transects. For 22 species, local estimates based on records of the TMD transects ranged between 0 and 10% of the regional population size. Accordingly, the study demonstrates that population size changes of most of the species can potentially be identified. Increasing cover of forest has a negative effect on species diversity and butterfly density, while the cover of nutrient‐poor grassland and the diversity of biotope types has a positive impact on butterfly density and species diversity. Solar insolation has a positive effect on butterfly density. Based on the results of the species–habitat relationships, the location of TMD transects could be optimised by aiming to cover the complete butterfly diversity of the study area. Additionally, monitoring of some of the species might be improved if winter surveys of eggs and larvae were implemented in the monitoring scheme.

Recent Advances in Twisted Structures of Flatland Materials and Crafting Moiré Superlattices

AbstractThe past decade has witnessed the occurrence of novel 2D moiré patterns in nanoflatland materials. These visually beautiful moiré superlattices have become a playground on which exotic quantum phenomena can be observed. The state‐of‐the‐art experimental techniques that have been developed for crafting moiré superlattices of flatland materials are reviewed. Graphene and its heterostructure with boron nitride have now sparked new interlayer twists as a new degree of freedom for tuning several angle‐dependent physical properties, e.g., the appearance of van Hove singularities, tunable Mott insulator states, and the Hofstadter butterfly pattern. Moreover, the interplay of correlated insulating states and superconductivity is recently observed for a so‐called magic‐angle twisted bilayer graphene. Furthermore, beyond graphene, other 2D materials, such as silicene, phosphorene, and the recent black phosphorus /MoS2 heterojunctions, which are 2D allotropes of bismuth and antimony grown on highly ordered pyrolytic graphite and MoS2, are considered. Finally, the optically important exciton phenomenon, which depends on the moiré potential and has been observed for a moiré superlattice of transition metal dichalcogenides, is discussed. This overview aims to cover all the fascinating prospects that depend on the moiré superlattice, ranging from electronic structure to optical exotics among flatland materials.

Seasonality in the production of male larvae: a game model for parasitic barnacles (Cirripedia: Rhizocephala)

AbstractThe male larvae of many parasitic barnacles are planktonic and are seasonally released. To achieve reproductive success, a male must be accepted by a receptive female that has successfully infected a host. To understand the seasonality of the breeding biology of parasitic barnacles, we developed an evolutionary game theoretical model for the seasonal pattern in the production of male larvae. Assumptions are that female parasitic barnacles become receptive following a given seasonal pattern. The parental females (mothers) choose the timing of producing their own male larvae to achieve maximum reproductive success. In the evolutionarily stable seasonal pattern, the production of male larvae often shows a sharp peak on a single day, indicating strongly synchronized production of male larvae, even when the supply of receptive females is distributed over the breeding season. When the total number of male larvae is large, the evolutionarily stable male production pattern may include multiple peaks, but it never shows a continuous distribution. This is very different from the game model previously developed for the emergence pattern of butterflies, where evolutionarily stable male emergence is always continuously distributed over a fraction of the mating season. As planktonic larvae, male parasitic barnacles have a naturally limited ability to find receptive females, and females may stay receptive for many days, whereas in butterflies, newly emerged females are mated within a day of their emergence.

The Arctiid Archetype: A New Lepidopteran Groundplan

The nymphalid groundplan (NGP) has proven to be extraordinarily useful in the study of butterfly color patterns because it allows for the identification of homologous elements across species. It has long been thought that the NGP is broadly applicable to all Lepidoptera, implying that the characters which make-up butterfly color patterns are homologous to those found in the moths. However, this conjecture remains mostly untested. We analyzed the wing patterns of the hyper-diverse arctiid tiger moths, which are represented worldwide by approximately 11,000 species, and found that the color patterns of these animals can be parsed into a limited toolkit of homologous characters. Some of the pattern elements identified, such as the basal and central symmetry system, are present on the NGP, but their morphology is often quite different from what is seen in the butterflies. The border ocelli of the NGP appear to be absent altogether in the Arctiidae, and conversely, two distal pattern element we term the ‘van Bemmelen’ and ‘terminal’ bands are present in the color patterns of many arctiids, but are not represented on the NGP. In light of the observed differences, we derive a new theoretical groundplan based on the original NGP that we refer to as the ‘arctiid archetype.’ This model provides a comprehensive framework for understanding how the wing patterns of these animals develop, and yields novel insight into their evolutionary history.

Wind-driven halo in high-contrast images

Context. The wind-driven halo is a feature that is observed in images that were delivered by the latest generation of ground-based instruments that are equipped with an extreme adaptive optics system and a coronagraphic device, such as SPHERE at the Very Large Telescope (VLT). This signature appears when the atmospheric turbulence conditions vary faster than the adaptive optics loop can correct for. The wind-driven halo is observed as a radial extension of the point spread function along a distinct direction (this is sometimes referred to as the butterfly pattern). When this is present, it significantly limits the contrast capabilities of the instrument and prevents the extraction of signals at close separation or extended signals such as circumstellar disks. This limitation is consequential because it contaminates the data for a substantial fraction of the time: about 30% of the data produced by the VLT/SPHERE instrument are affected by the wind-driven halo. Aims. This paper reviews the causes of the wind-driven halo and presents a method for analyzing its contribution directly from the scientific images. Its effect on the raw contrast and on the final contrast after post-processing is demonstrated. Methods. We used simulations and on-sky SPHERE data to verify that the parameters extracted with our method can describe the wind-driven halo in the images. We studied the temporal, spatial, and spectral variation of these parameters to point out its deleterious effect on the final contrast. Results. The data-driven analysis we propose provides information to accurately describe the wind-driven halo contribution in the images. This analysis confirms that this is a fundamental limitation of the finally reached contrast performance. Conclusions. With the established procedure, we will analyze a large sample of data delivered by SPHERE in order to propose post-processing techniques that are tailored to removing the wind-driven halo.

Transcription factors underlying wing margin color patterns and pupal cuticle markings in butterflies

BackgroundThe diversity of butterfly color patterns can be attributed to a relatively small number of pattern elements that are homologous across Lepidoptera. Although genes involved in patterning some of these elements have been identified, the development of several major elements remains poorly understood. To identify genes underlying wing pupal cuticle markings and wing margin color patterns, we examined expression of the candidate transcription factors Engrailed/Invected (En/Inv), Distal-less (Dll), Cubitus interruptus (Ci), and Spalt in two nymphalids: Junonia coenia and Bicyclus anynana.ResultsWe found that En/Inv, Dll, and Ci mark domains on the J. coenia last-instar forewing disc that closely correspond to the position and shape of pupal cuticle markings. We also found that Spalt demarcates wing margin color patterns in both J. coenia and B. anynana, and that CRISPR/Cas9 deletions in the spalt gene result in reduction and loss of wing margin color patterns in J. coenia. These data demonstrate a role for spalt in promoting wing margin color patterning, in addition to its previously described role in eyespot patterning.ConclusionOur observations support the model that a core set of regulatory genes are redeployed multiple times, and in multiple roles, during butterfly wing pattern development. Of these genes, spalt is of special interest as it plays a dual role in both eyespot and margin color pattern development.

Spatial Distribution of Butterflies in Accordance with Climate Change in the Korean Peninsula

The effects of climate change are becoming apparent in the biosphere. In the 20th century, South Korea experienced a 1.5 °C temperature increase due to rapid industrialization and urbanization. If the changes continue, it is predicted that approximately 15–37% of animal and plant species will be endangered after 2050. Because butterflies act as a good indicator for changes in the temperature, the distribution of butterflies can be used to determine their adaptability to climate patterns. Local meteorological data for the period 1938–2011 were used from the National Forest Research Institute of Korea. Local temperature data were additionally considered among the basic information, and the distribution patterns of butterflies were analyzed for both the southern and northern regions. Southern butterflies (with northern limit) tend to increase in number with significant correlation between the temperature and number of habitats (p < 0.000), while northern butterflies (with southern limit) show no statistical significance between the temperature and number of habitats, indicating their sensitivity to temperature change. This finding is in accordance with the conclusion that southern butterflies are more susceptible to climate change when adapting to local environments and expanding their original temperature range for survival, which leads to an increase in the numbers of their habitats.

Long-lived radionuclides in marine sediments from the Beibu Gulf, South China Sea: Spatial distribution, controlling factors, and proxy for transport pathway

The source and transport process of sediments are of great concern in marine sedimentology. In this study, naturally occurring radionuclides (238U, 226Ra, 228Ra, and 40K) in surface sediments and sediment cores were comprehensively analysed using high-purity germanium (HPGe) γ spectrometry in order to explore radionuclide fingerprints as novel proxies for tracking the source and transport pathway of sediments in the Beibu Gulf. We found that the radioactivity level of sediments in the Beibu Gulf was lower than that in other Chinese seas, following a comprehensive compilation and comparison of radioactivity in the region, indicating the Beibu Gulf is a high-quality marine environment. An interesting “butterfly pattern” was exhibited in the horizontal distribution of radionuclides in surface sediments from the Beibu Gulf, mainly attributable to the controlling factors of the non-linear negative regulation of sediment grain size and positive role of total organic carbon in sediment. Furthermore, the vertical distribution of radionuclides in sediment cores was affected by the redox state and a physical/biological disturbance, which were demonstrated by the simultaneous analysis of redox-sensitive element Mn and 210Pb, respectively. Additionally, the 226Ra238U endmember model and 232Th238U endmember model were proposed for quantifying local input (85%) and trans-region input (15%) of marine sediments in the Beibu Gulf in the context of the Guangdong-Qiongzhou Strait-Guangxi framework. We inferred that the Beibu Gulf was at risk of contamination by artificial radionuclides derived from nearby Guangdong Province, which has the highest number of nuclear power plants in China, through a trans-region input by ocean currents. Our study not only provided spatiotemporal distributions of radionuclides in the Beibu Gulf and their controlling factors (sediment grain size, total organic carbon, redox state, physical/biological disturbance, and ocean currents) but also indicated the radionuclide fingerprints (226Ra/238U and 232Th/238U) as potential geochemical proxies for quantifying the distinct sources of radionuclides and other associated chemicals (e.g., heavy metals, organic pollutants) in sediments.

Spectral organization of the compound eye of a migrating nymphalid, the chestnut tiger butterfly Parantica sita.

Several butterflies of family Nymphalidae perform long-distance migration. Extensive studies of migration in the iconic monarch butterfly Danaus plexippus have revealed that vision plays a crucial role in migratory orientation. Differences in the migratory patterns of butterflies suggest that not all species are exposed to the same visual conditions and yet, little is known about how the visual system varies across migratory species. Here, we used intracellular electrophysiology, dye injection and electron microscopy to assess the spectral and polarization properties of the photoreceptors of a migrating nymphalid, Parantica sita Our findings reveal three spectral classes of photoreceptors including ultraviolet, blue and green receptors. The green receptor class contains three subclasses, which are broad, narrow and double-peaking green receptors. Ultraviolet and blue receptors are sensitive to polarized light parallel to the dorso-ventral axis of the animal, while the variety of green receptors are sensitive to light polarized at 45deg, 90deg and 135deg away from the dorso-ventral axis. The polarization sensitivity ratio is constant across spectral receptor classes at around 1.8. Although P. sita has a typical nymphalid eye with three classes of spectral receptors, subtle differences exist among the eyes of migratory nymphalids, which may be genus specific.

Disentangling the effects of host resources, local, and landscape variables on the occurrence pattern of the dusky large blue butterfly (Phengaris nausithous) in upland grasslands

Determining the effects of local and landscape drivers on endangered species and predicting potential suitable habitats for their persistence is crucial for effective conservation management. Here, we applied a multi-scale approach to disentangle the effects of host resources, local, and landscape variables on the occurrence pattern of Phengaris (= Maculinea) nausithous in semi-natural upland grasslands. Our approach comprised the assessment of host parameters (plant cover, density, height, flower heads density, ant nest density, ant colony size), local grassland management (pasture, meadow), site conditions (area, shape, terrain attributes), and landscape variables (landscape composition, connectivity). We used ensemble of small models based on bivariate generalized linear models for explaining and predicting the butterfly occurrence pattern. Bivariate models revealed that host ant nest density, plant cover and height, local grassland management type (pasture), slope and eastness, landscape forest cover and grassland connectivity had a positive effect on the occurrence of P. nausithous (average explained deviance 20.5%). Host ant density, host plant cover, and local grassland management were the most influential factors on the ensemble predictions. The presence of P. nausithous in upland grasslands is not only determined by host resources, but also by local and landscape factors. Such factors proved to be relevant for identifying and predicting suitable grassland sites for this endangered species. Consequently, we recommend that conservation actions should include a landscape perspective to promote connectivity by facilitating coherent grazing networks enabling dispersal between semi-natural upland grasslands and thus species persistence.

Features of Spatiotemporal Clustering in a Maunder Butterfly Diagram

The Maunder butterfly pattern is the most complete spatial-temporal representation of observed changes in solar activity in the 11-year cycle over a period of 12–24 cycle. The well-known empirical relation is used to transform Greenwich sunspot areas into magnetic flux, and the distances between the butterfly wings are then calculated with the Fisher–Rao metric. We found that the similarities or differences in the patterns of the individual butterfly wings in this metric are approximately the same for each hemisphere. The wings were the closest for a sequence of strong cycles, while there is a tendency for a series of weak cycles to form cycles in pairs with the implementation of an analog of the Gnevyshev–Olya rule.

Interplay between Developmental Flexibility and Determinism in the Evolution of Mimetic Heliconius Wing Patterns.

To what extent can we predict how evolution occurs? Do genetic architectures and developmental processes canalize the evolution of similar outcomes in a predictable manner? Or do historical contingencies impose alternative pathways to answer the same challenge? Examples of Müllerian mimicry between distantly related butterfly species provide natural replicates of evolution, allowing us to test whether identical wing patterns followed parallel or novel trajectories. Here, we explore the role that the signaling ligand WntA plays in generating mimetic wing patterns in Heliconius butterflies, a group with extraordinary mimicry-related wing pattern diversity. The radiation is relatively young, and numerous cases of wing pattern mimicry have evolved within the last 2.5-4.5 Ma. WntA is an important target of natural selection and is one of four major effect loci that underlie much of the pattern variation in the group. We used CRISPR/Cas9 targeted mutagenesis to generate WntA-deficient wings in 12 species and a further 10 intraspecific variants, including three co-mimetic pairs. In all tested butterflies, WntA knockouts affect pattern broadly and cause a shift among every possible scale cell type. Interestingly, the co-mimics lacking WntA were very different, suggesting that the gene networks that pattern a wing have diverged considerably among different lineages. Thus, although natural selection channeled phenotypic convergence, divergent developmental contexts between the two major Heliconius lineages opened different developmental routes to evolve resemblance. Consequently, even under very deterministic evolutionary scenarios, our results underscore a surprising unpredictability in the developmental paths underlying convergence in a recent radiation.

Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths

Butterflies and moths (Lepidoptera) are one of the major superradiations of insects, comprising nearly 160,000 described extant species. As herbivores, pollinators, and prey, Lepidoptera play a fundamental role in almost every terrestrial ecosystem. Lepidoptera are also indicators of environmental change and serve as models for research on mimicry and genetics. They have been central to the development of coevolutionary hypotheses, such as butterflies with flowering plants and moths' evolutionary arms race with echolocating bats. However, these hypotheses have not been rigorously tested, because a robust lepidopteran phylogeny and timing of evolutionary novelties are lacking. To address these issues, we inferred a comprehensive phylogeny of Lepidoptera, using the largest dataset assembled for the order (2,098 orthologous protein-coding genes from transcriptomes of 186 species, representing nearly all superfamilies), and dated it with carefully evaluated synapomorphy-based fossils. The oldest members of the Lepidoptera crown group appeared in the Late Carboniferous (∼300 Ma) and fed on nonvascular land plants. Lepidoptera evolved the tube-like proboscis in the Middle Triassic (∼241 Ma), which allowed them to acquire nectar from flowering plants. This morphological innovation, along with other traits, likely promoted the extraordinary diversification of superfamily-level lepidopteran crown groups. The ancestor of butterflies was likely nocturnal, and our results indicate that butterflies became day-flying in the Late Cretaceous (∼98 Ma). Moth hearing organs arose multiple times before the evolutionary arms race between moths and bats, perhaps initially detecting a wide range of sound frequencies before being co-opted to specifically detect bat sonar. Our study provides an essential framework for future comparative studies on butterfly and moth evolution.

A review of red-eye pigmentation and diel activity patterns in skippers (Lepidoptera, Papilionoidea, Hesperiidae)

ABSTRACTThe visual systems and diel activity patterns of butterflies and moths have been studied for decades, yet understanding the underlying mechanisms that are associated with the evolution of these remains a major challenge. The order Lepidoptera is principally composed of nocturnal lineages with extreme morphological and behavioural adaptations to this lifestyle. Although most butterflies are diurnal, some clades are predominantly nocturnal, for instance, the Hedylidae. Skippers are another family that comprises multiple crepuscular and/or nocturnal lineages. Interestingly, many such lineages have red-eye pigmentation, and this morphological feature has been hypothesised to be an adaptation to the night-time lifestyle. Here, we review the occurrence and frequency of red-eye pigmentation as well as diel activity patterns in Hesperiidae. We place these data within a phylogenomic framework and provide hypotheses and predictions relative to the evolution of red-eye pigmentation in skippers with respect to diel activity. We suggest that several types of red-eye pigmentation exist in skippers, with possible structural red-eye pigmentation in some lineages of diurnal skippers. The colouration of eyes in Hesperiidae, the evolutionary mechanisms that drive these traits and the physiological underlying processes require further attention and could bring new insights into our understanding of butterfly vision systems.

Evolution of aposematism and mimicry in butterflies: Causes, consequences and paradoxes

Evolution of aposematism and mimicry in butterflies: Causes, consequences and paradoxes

Gene Flow in the Müllerian Mimicry Ring of a Poisonous Papuan Songbird Clade (Pitohui; Aves).

Müllerian mimicry rings are remarkable symbiotic species assemblages in which multiple members share a similar phenotype. However, their evolutionary origin remains poorly understood. Although gene flow among species has been shown to generate mimetic patterns in some Heliconius butterflies, mimicry is believed to be due to true convergence without gene flow in many other cases. We investigated the evolutionary history of multiple members of a passerine mimicry ring in the poisonous Papuan pitohuis. Previous phylogenetic evidence indicates that the aposematic coloration shared by many, but not all, members of this genus is ancestral and has only been retained by members of the mimicry ring. Using a newly assembled genome and thousands of genomic DNA markers, we demonstrate gene flow from the hooded pitohui (Pitohui dichrous) into the southern variable pitohui (Pitohui uropygialis), consistent with shared patterns of aposematic coloration. The vicinity of putatively introgressed loci is significantly enriched for genes that are important in melanin pigment expression and toxin resistance, suggesting that gene flow may have been instrumental in the sharing of plumage patterns and toxicity. These results indicate that interspecies gene flow may be a more general mechanism in generating mimicry rings than hitherto appreciated.

The mimetic wing pattern of Papilio polytes butterflies is regulated by a doublesex-orchestrated gene network

The swallowtail butterfly Papilio polytes is sexually dimorphic and exhibits female-limited Batesian mimicry. This species also has two female forms, a non-mimetic form with male-like wing patterns, and a mimetic form resembling an unpalatable model, Pachliopta aristolochiae. The mimicry locus H constitutes a dimorphic Mendelian ‘supergene’, including a transcription factor gene doublesex (dsx). However, how the mimetic-type dsx (dsx-H) orchestrates the downstream gene network and causes the mimetic traits remains unclear. Here we performed RNA-seq-based gene screening and found that Wnt1 and Wnt6 are up-regulated by dsx-H during the early pupal stage and are involved in the red/white pigmentation and patterning of mimetic female wings. In contrast, a homeobox gene abdominal-A is repressed by dsx-H and involved in the non-mimetic colouration pattern. These findings suggest that dual regulation by dsx-H, induction of mimetic gene networks and repression of non-mimetic gene networks, is essential for the switch from non-mimetic to mimetic pattern in mimetic female wings.

Radio telemetry helps record the dispersal patterns of birdwing butterflies in mountainous habitats: Golden Birdwing (Troides aeacus) as an example

Birdwing butterflies are a monophyletic group of swallowtail butterflies (Papilionidae) protected by national and international laws and often serve as flagships of insect conservation. Selecting the Golden Birdwing (Troides aeacus) as an example, we demonstrate an effective way to simultaneously record the activity of multiple birdwing butterflies using radio telemetry in hard-to-access mountainous terrain. During the summer flight period of T. aeacus (June and July), a single researcher was able to obtain 30-min records of movement patterns for ten individuals for an average of 4 days by fastening 0.22 g transmitters onto the butterflies, in the mountainous valleys of Mt. Gongga, China. The maximum distance the butterflies traveled over the 4 day period was 4314 m away from the starting location. During this time, the average dispersal rate was 38.07 m/h (n = 9, sd = 85.11); average movement speed was 293.48 m/h (n = 9, sd = 121.45). Flight patterns of butterflies collected from low and high elevation habitats showed no significant differences. Activity levels of individuals from both low and high elevation habitats track diurnal fluctuation in temperature. Flight activity is positively correlated with temperature and negatively correlated with humidity. Our data provide basic parameters of real-time flight activity and dispersal ability for a species of conservation importance. The methodology is highly suitable for monitoring endangered lepidopteran species in otherwise difficult-to-access terrain.

Genomes reveal drastic and recurrent phenotypic divergence in firetip skipper butterflies (Hesperiidae: Pyrrhopyginae)

Biologists marvel at the powers of adaptive convergence, when distantly related animals look alike. While mimetic wing patterns of butterflies have fooled predators for millennia, entomologists inferred that mimics were distant relatives despite similar appearance. However, the obverse question has not been frequently asked. Who are the close relatives of mimetic butterflies and what are their features? As opposed to close convergence, divergence from a non-mimetic relative would also be extreme. When closely related animals look unalike, it is challenging to pair them. Genomic analysis promises to elucidate evolutionary relationships and shed light on molecular mechanisms of divergence. We chose the firetip skipper butterfly as a model due to its phenotypic diversity and abundance of mimicry. We sequenced and analysed whole genomes of nearly 120 representative species. Genomes partitioned this subfamily Pyrrhopyginae into five tribes (1 new), 23 genera and, additionally, 22 subgenera (10 new). The largest tribe Pyrrhopygini is divided into four subtribes (three new). Surprisingly, we found five cases where a uniquely patterned butterfly was formerly placed in a genus of its own and separately from its close relatives. In several cases, extreme and rapid phenotypic divergence involved not only wing patterns but also the structure of the male genitalia. The visually striking wing pattern difference between close relatives frequently involves disappearance or suffusion of spots and colour exchange between orange and blue. These differences (in particular, a transition between unspotted black and striped wings) happen recurrently on a short evolutionary time scale, and are therefore probably achieved by a small number of mutations.

Modular cis-regulatory logic of yellow gene expression in silkmoth larvae.

Colour patterns in butterflies and moths are crucial traits for adaptation. Previous investigations have highlighted genes responsible for pigmentation (ie yellow and ebony). However, the mechanisms by which these genes are regulated in lepidopteran insects remain poorly understood. To elucidate this, molecular studies involving dipterans have largely analysed the cis‐regulatory regions of pigmentation genes and have revealed cis‐regulatory modularity. Here, we used well‐developed transgenic techniques in Bombyx mori and demonstrated that cis‐regulatory modularity controls tissue‐specific expression of the yellow gene. We first identified which body parts are regulated by the yellow gene via black pigmentation. We then isolated three discrete regulatory elements driving tissue‐specific gene expression in three regions of B. mori larvae. Finally, we found that there is no apparent sequence conservation of cis‐regulatory regions between B. mori and Drosophila melanogaster, and no expression driven by the regulatory regions of one species when introduced into the other species. Therefore, the trans‐regulatory landscapes of the yellow gene differ significantly between the two taxa. The results of this study confirm that lepidopteran species use cis‐regulatory modules to control gene expression related to pigmentation, and represent a powerful cadre of transgenic tools for studying evolutionary developmental mechanisms.