Faunal structures among European butterflies: evolutionary implications of bias for geography, endemism and taxonomic affiliation

European butterfly species have been classified, using four multivariate classification techniques, to faunal groups (a collection of species having similar distributions) each group having a unique geography (faunal element). Concordance occurs for 94% of species for at least three of the techniques. The faunal groups are brassed in geography, endemicity and taxonomic affiliation, indicating that they have historical and evolutionary significance. The inference is that a species by belonging to one faunal group, rather than to another, has a higher probability of being an endemic, and if it is not an endemic, then of evolving into one. This probability is influenced by affiliation to higher taxa, such as butterfly families, and thus by phylogenetic constraints. We argue that the fidelity of a species to a faunal element will affect its evolutionary pathway since, by belonging to a faunal element, a species is subject lo the distinctive processes linked to a unique landscape and its environmental conditions. Species have the capacity of switching between faunal groups owing to environmental changes impinging on their geographical ranges and epigenotypes. However, transfer of species across different groups are not of equal likelihood, being greatest between contiguous land elements and least between those restricted to islands. We suggest that conservation biology gives more attention to faunal structures: faunal groups have unique geography, are vulnerable to different macroevolutionary pressures and effectively underpin community assemblages within specific biotopes.     

Unique wing scale photonics of male Rajah Brooke’s birdwing butterflies

Background

Ultrastructures in butterfly wing scales can take many shapes, resulting in the often striking coloration of many butterflies due to interference of light. The plethora of coloration mechanisms is dazzling, but often only single mechanisms are described for specific animals.

Results

We have here investigated the male Rajah Brooke’s birdwing, Trogonoptera brookiana, a large butterfly from Malaysia, which is marked by striking, colorful wing patterns. The dorsal side is decorated with large, iridescent green patterning, while the ventral side of the wings is primarily brown-black with small white, blue and green patches on the hindwings. Dense arrays of red hairs, creating a distinct collar as well as contrasting areas ventrally around the thorax, enhance the butterfly’s beauty. The remarkable coloration is realized by a diverse number of intricate and complicated nanostructures in the hairs as well as the wing scales. The red collar hairs contain a broad-band absorbing pigment as well as UV-reflecting multilayers resembling the photonic structures of Morpho butterflies; the white wing patches consist of scales with prominent thin film reflectors; the blue patches have scales with ridge multilayers and these scales also have centrally concentrated melanin. The green wing areas consist of strongly curved scales, which possess a uniquely arranged photonic structure consisting of multilayers and melanin baffles that produces highly directional reflections.

Conclusion

Rajah Brooke’s birdwing employs a variety of structural and pigmentary coloration mechanisms to achieve its stunning optical appearance. The intriguing usage of order and disorder in related photonic structures in the butterfly wing scales may inspire novel optical materials as well as investigations into the development of these nanostructures in vivo.

     

Fine-Grained Distribution of a Non-Native Resource Can Alter the Population Dynamics of a Native Consumer

New interactions with non-native species can alter selection pressures on native species. Here, we examined the effect of the spatial distribution of a non-native species, a factor that determines ecological and evolutionary outcomes but that is poorly understood, particularly on a fine scale. Specifically, we explored a native butterfly population and a non-native plant on which the butterfly oviposits despite the plant’s toxicity to larvae. We developed an individual-based model to describe movement and oviposition behaviors of each butterfly, which were determined by plant distribution and the butterfly''s host preference genotype. We estimated the parameter values of the model from rich field data. We simulated various patterns of plant distributions and compared the rates of butterfly population growth and changes in the allele frequency of oviposition preference. Neither the number nor mean area of patches of non-native species affected the butterfly population, whereas plant abundance, patch shape, and distance to the nearest native and non-native patches altered both the population dynamics and genetics. Furthermore, we found a dramatic decrease in population growth rates when we reduced the distance to the nearest native patch from 147 m to 136 m. Thus changes in the non-native resource distribution that are critical to the fate of the native herbivore could only be detected at a fine-grained scale that matched the scale of a female butterfly’s movement. In addition, we found that the native butterfly population was unlikely to be rescued by the exclusion of the allele for acceptance of the non-native plant as a host. This study thus highlights the importance of including both ecological and evolutionary dynamics in analyses of the outcome of species interactions and provides insights into habitat management for non-native species.     

Spectral organization of the eye of a butterfly,Papilio

Journal of Comparative Physiology A - This review outlines our recent studies on the spectral organization of butterfly compound eyes, with emphasis on the Japanese yellow swallowtail butterfly,...     

Structural Colour in Butterfly Apatura Ilia Scales and the Microstructure Simulation of Photonic Crystal

The butterfly Apatura ilia is a species in the Northeast of China. There are billions of tiny scales on its wings, which overlap like roof tiles and completely cover the membrane, appearing as dust to people naked eye. The scales produce brilliant structural colour through their multilayer microstructure. In this paper, the microstructure and geometrical dimension of the scales were observed using a Scanning Electron Microscope (SEM). The cross section micro-configuration of the purple scales was achieved using a Transmission Electron Microscope (TEM). The reflectivity of the wing was measured by a spectrometer. The 3D multilayer microstructure of the ridges was optimized to 1D photonic crystal structure. The spectrometer experimental graph is in accord with the 1D photonic crystal simulation curves basically. In the end, the phenomenon of the purple structural colour was explained through the Snell equation.     

Anisotropism of the Non-Smooth Surface of Butterfly Wing

Twenty-nine species of butterflies were collected for observation and determination of the wing surfaces using a ScanningElectron Microscope(SEM).Butterfly wing surface displays structural anisotropism in micro-,submicro- and nano-scales.Thescales on butterfly wing surface arrange like overlapping roof tiles.There are submicrometric vertical gibbosities,horizontallinks,and nano-protuberances on the scales.First-incline-then-drip method and first-drip-then-incline method were used tomeasure the Sliding Angle(SA)of droplet on butterfly wing surface by an optical Contact Angle(CA)measuring system.Relatively smaller sliding angles indicate that the butterfly wing surface has fine self-cleaning property.Significantly differentSAs in various directions indicate the anisotropic self-cleaning property of butterfly wing surface.The SAs on the butterfly wingsurface without scales are remarkably larger than those with scales,which proves the crucial role of scales in determining theself-cleaning property.Butterfly wing surface is a template for design and fabrication ofbiomimetic materials and self-cleaningsubstrates.This work may offer insights into how to design directional self-cleaning coatings and anisotropic wetting surface.     

Butterfly wing patterns

This paper integrates genetical studies of variation in the wing patterns of Lepidoptera with experimental investigations of developmental mechanisms. Research on the tropical butterfly,Bicyclus anynana, is described. This work includes artificial selection of lines with different patterns of wing eyespots followed by grafting experiments on the lines to examine the phenotypic and genetic differences in terms of developmental mechanisms. The results are used to show how constraints on the evolution of this wing pattern may be related to the developmental organisation. The eyespot pattrn can be envisaged as a set of developmental homologues; a common developmental mechanism is associated with a quantitative genetic system involving high genetic correlations. However, individual genes which influence only subsets of the eyespots, thus uncoupling the interdependence of the eyespots, may be important in evolutionary change. The postulated evolutionary constraints are illustrated with respect to differences in wing pattern found among other species ofBicyclus.     

Light Trapping Effect in Wing Scales of Butterfly Papilio peranthus and Its Simulations

Broadband light trapping effect and arrays of sub-wavelength textured structures based on the butterfly wing scales are applicable to solar cells and stealth technologies. In this paper, the fine optical structures in wing scales of butterfly Papilio peranthus, exhibiting efficient light trapping effect, were carefully examined. First, the reflectivity was measured by reflectance spectrum. Field Emission Scanning Electronic Microscope (FESEM) and Transmission Electron Microscope (TEM) were used to observe the coupling morphologies and structures of the scales. Then, the optimized 3D model of the coupling structure was created combining Scanning Electron Microscope (SEM) and TEM data. Afterwards, the mechanism of the light trapping effect of these structures was analyzed by simulation and theoretical calculations. A multilayer nano-structure of chitin and air was found. These structures are effective in increasing optical path, resulting in that most of the incident light can be trapped and adsorbed within the structure at last. Furthermore, the simulated optical results are consistent with the experimental and calculated ones. This result reliably confirms that these structures induce an efficient light trapping effect. This work can be used as a reference for in-depth study on the fabrication of highly efficient bionic optical devices, such as solar cells, photo detectors, high-contrast, antiglare, and so forth.     

Effects of diet and metamorphosis upon the sterol composition of the butterfly Morpho peleides

Whole body sterol metabolism in insects has seldom been studied. We were able to design an appropriate study at a butterfly farm in Belize. We collected six larvas of butterfly (Morpho peleides), their food (leaves of Pterocarpus bayessii), and their excretions. In addition, six adult butterflies were collected. The sterols of the diet, the larva, and adult butterfly were analyzed by gas-liquid chromatography. The structures of these sterols were identified by digitonin precipitation, GC-MS, and NMR. Four sterols (cholesterol, campesterol, stigmasterol, and sitosterol) and a sterol mixture were found in the food, the body, and the excreta of the larva. The tissue sterol content of the larva was 326 microg. They consumed 276 microg of sterols per day. Their excretion was 185 microg per day as sterols. The total tissue sterol contents of the larva and butterfly were similar, but they had different sterol compositions, which indicated interconversion of sterols during development. There was a progressive increase in the cholesterol content from larva to butterfly and a decrease in the content of sitosterol and other plant sterols, which were likely converted to cholesterol. Our data indicated an active sterol metabolism in butterfly larva. Diet played an important role in determining its sterol composition. During metamorphosis, there was an interconversion of sterols. This is the first paper documenting the fecal sterol excretion in insects as related to dietary intakes.     

Mitochondrial phylogenomics,the origin of swallowtail butterflies,and the impact of the number of clocks in Bayesian molecular dating

Swallowtail butterflies (Lepidoptera: Papilionidae) have been instrumental in understanding many foundational concepts in biology; despite this, a resolved and robust phylogeny of the group has been a major impediment to elucidating patterns and processes of their ecological and evolutionary history. This study presents a mitogenomic, time‐calibrated phylogeny for all swallowtail genera. A shotgun sequencing approach was performed to obtain 32 complete mitogenomes that were added to available butterfly mitogenomes, resulting in a dataset including 142 butterfly taxa (and four outgroups) representing all butterfly families. Phylogenetic analyses were carried out under maximum likelihood (ML) and Bayesian inferences (BIs) with alternative partitioning strategies and the mixture (CAT) model. To test competing hypotheses about the systematics of Papilionidae, such as the enigmatic position of Baronia brevicornis or the status of the tribe Teinopalpini, we estimated the marginal likelihood of alternative topologies and computed Bayes factors. Estimates of divergence times were assessed using a Bayesian relaxed‐clock approach calibrated with six fossils while testing for the number of clocks. The results recovered a well‐resolved and supported phylogeny confirming that Baroniinae is sister to Parnassiinae + Papilioninae, both recovered as monophyletic. It also laid the foundations for classification at tribe and genus level, suggesting that the tribe Teinopalpini only contains the genus Teinopalpus (Meandrusa being sister to Papilio ). The number of molecular clocks in dating analyses had a significant impact on divergence times. A single clock recovered an origin of butterflies in the Cretaceous (98, 66–188 Ma) and also for swallowtails (85, 55–163 Ma), while partitioning the clocks yielded an origin of Papilionoidea in the very Late Cretaceous (71, 64–86 Ma), and all butterfly families originated in the aftermath of the Cretaceous–Paleogene extinction. These results challenge previous studies suggesting that butterflies appeared in the Early Cretaceous, 110 Ma, concurrently with the rise of angiosperms.     

Butterfly morphology in a molecular age – Does it still matter in butterfly systematics?

We review morphological characters considered important for understanding butterfly phylogeny and evolution in the light of recent large-scale molecular phylogenies of the group. A number of the most important morphological works from the past half century are reviewed and morphological character evolution is reassessed based on the most recent phylogenetic results. In particular, higher level butterfly morphology is evaluated based on a very recent study combining an elaborate morphological dataset with a similar molecular one. Special attention is also given to the families Papilionidae, Nymphalidae and Hesperiidae which have all seen morphological and molecular efforts come together in large, combined works in recent years. In all of the examined cases the synergistic effect of combining elaborate morphological datasets with ditto molecular clearly outweigh the merits of either data type analysed on its own (even for ‘genome size’ molecular datasets). It is evident that morphology, far from being obsolete or arcane, still has an immensely important role to play in butterfly (and insect) phylogenetics. Not least because understanding morphology is essential for understanding and evaluating the evolutionary scenarios phylogenetic trees are supposed to illustrate.     

Fission in Convolutriloba longifissura: asexual reproduction in acoelous turbellarians revisited

Studies of the asexual reproduction of Convolutriloba longifissura (Acoela, Acoelomorpha) revealed that there is no longitudinal fission of the whole animal as has been described by Bartolomaeus and Balzer (1997) . Instead, the first step is a transverse fission. This results in the detachment of the caudal fourth of the mother animal. The detached part forms what we call the butterfly stage, which initially has no mouth and no eye fields. This stage gives rise to two new individuals by a longitudinal fission. Within 2–3 days the eye fields and a mouth develop in each of the two progenies formed in this way. In the meantime the mother individual grows and develops the three typical caudal lobes. The mother animal can repeat this process resulting in three individuals every fourth day. The finding of this new pattern of reproduction in the Acoela has prompted us to review the various ways by which asexual reproduction occurs in the group. The peculiar combination of few cases but high diversity of asexual reproduction in the Acoela is discussed from an evolutionary point of view.     

Signaling,solidarity, and the sacred: The evolution of religious behavior

Anthropologists have repeatedly noted that there has been little theoretical progress in the anthropology of religion over the past fifty years.^1–7 By the 1960s, Geertz² had pronounced the field dead. Recently, however, evolutionary researchers have turned their attention toward understanding the selective pressures that have shaped the human capacity for religious thoughts and behaviors, and appear to be resurrecting this long‐dormant but important area of research.^8–19 This work, which focuses on ultimate evolutionary explanations, is being complemented by advances in neuropsychology and a growing interest among neuroscientists in how ritual, trance, meditation, and other altered states affect brain functioning and development.^20–26 This latter research is providing critical insights into the evolution of the proximate mechanisms responsible for religious behavior. Here we review these literatures and examine both the proximate mechanisms and ultimate evolutionary processes essential for developing a comprehensive evolutionary explanation of religion.     

The Functional Role of the Hollow Region of the Butterfly Pyrameis atalanta （L.） Scale

Questions concerning the functional role of the hollow region of the butterfly Pyrameis atalanta （L.） scale are experimentally investigated. Attention was initially directed to this problem by observation of the complex microstrucmre of the butterfly scale as well as other studies indicating higher lift on butterfly wings covered with scale. The aerodynamic forces were measured for two oscillating scale models. Results indicated that the air cavity of an oscillating model of the Pyrameis atalanta （L.） scale increased the lift by a factor of 1.15 and reduced the damping coefficients by a factor of 1.38. The modification of the aerodynamic effects on the model of butterfly scale was due to an increase of the virtual air mass, which influenced the body. The hollow region of the scale increased the virtual air mass by a factor of 1.2. The virtual mass of the butterfly scale with the hollow region was represented as the sum of air mass of two imaginary geometrical figures： a circular cylinder around the scale and a right-angled parallelepiped within the hollow region. The interaction mechanism of the butterfly Pyrameis atalanta （L.） scale with a flow was described. This novel interaction mechanism explained most geometrical features of the airpermeable butterfly scale （inverted V-profile of the ridges, nozzle of the tip edge, hollow region, and openings of the upper lamina） and their arrangement.     

Generating phenotypic variation: prospects from "evo-devo" research on Bicyclus anynana wing patterns

Understanding the generation of phenotypic variation is an important challenge for modern evolutionary biology, and butterfly wing patterns are an exciting system that can shed some light on this issue. Here, we report on recent advances in the genetics of Bicyclus anynana butterflies. This system provides the potential for a fully integrated study of the evolutionary and developmental processes underlying diversity in morphology.     

Automatic recognition and measurement of butterfly eyespot patterns

A favorite wing pattern element in butterflies that has been the focus of intense study in evolutionary and developmental biology, as well as in behavioral ecology, is the eyespot. Because the pace of research on these bull's eye patterns is accelerating we sought to develop a tool to automatically detect and measure butterfly eyespot patterns in digital images of the wings. We used a machine learning algorithm with features based on circularity and symmetry to detect eyespots on the images. The algorithm is first trained with examples from a database of images with two different labels (eyespot and non-eyespot), and subsequently is able to provide classification for a new image. After an eyespot is detected the radius measurements of its color rings are performed by a 1D Hough Transform which corresponds to histogramming. We trained software to recognize eyespot patterns of the nymphalid butterfly Bicyclus anynana but eyespots of other butterfly species were also successfully detected by the software.     

Kowalevsky, comparative evolutionary embryology, and the intellectual lineage of evo-devo

Alexander Kowalevsky was one of the most significant 19th century biologists working at the intersection of evolution and embryology. The reinstatement of the Alexander Kowalevsky Medal by the St. Petersburg Society of Naturalists for outstanding contributions to understanding evolutionary relationships in the animal kingdom, evolutionary developmental biology, and comparative zoology is timely now that Evo-devo has emerged as a major research discipline in contemporary biology. Consideration of the intellectual lineage of comparative evolutionary embryology explicitly forces a reconsideration of some current conceptions of the modern emergence of Evo-devo, which has tended to exist in the shadow of experimental embryology throughout the 20th century, especially with respect to the recent success of developmental biology and developmental genetics. In particular we advocate a sharper distinction between the heritage of problems and the heritage of tools for contemporary Evo-devo. We provide brief overviews of the work of N. J. Berrill and D. T. Anderson to illustrate comparative evolutionary embryology in the 20th century, which provides an appropriate contextualization for a conceptual review of our research on the sea urchin genus Heliocidaris over the past two decades. We conclude that keeping research questions rather than experimental capabilities at the forefront of Evo-devo may be an antidote to any repeat of the stagnation experienced by the first group of evolutionary developmental biologists over one hundred years ago and acknowledges Kowalevsky's legacy in evolutionary embryology.     

Culture, embodiment and genes: unravelling the triple helix

Much recent work stresses the role of embodiment and action in thought and reason, and celebrates the power of transmitted cultural and environmental structures to transform the problem-solving activity required of individual brains. By apparent contrast, much work in evolutionary psychology has stressed the selective fit of the biological brain to an ancestral environment of evolutionary adaptedness, with an attendant stress upon the limitations and cognitive biases that result. On the face of it, this suggests either a tension or, at least, a mismatch, with the symbiotic dyad of cultural evolution and embodied cognition. In what follows, we explore this mismatch by focusing on three key ideas: cognitive niche construction; cognitive modularity; and the existence (or otherwise) of an evolved universal human nature. An appreciation of the power and scope of the first, combined with consequently more nuanced visions of the latter two, allow us to begin to glimpse a much richer vision of the combined interactive potency of biological and cultural evolution for active, embodied agents.