Sample records for butterfly wing patterns

  1. Genetic Basis of Melanin Pigmentation in Butterfly Wings

    PubMed Central

    Zhang, Linlin; Martin, Arnaud; Perry, Michael W.; van der Burg, Karin R. L.; Matsuoka, Yuji; Monteiro, Antónia; Reed, Robert D.

    2017-01-01

    Despite the variety, prominence, and adaptive significance of butterfly wing patterns, surprisingly little is known about the genetic basis of wing color diversity. Even though there is intense interest in wing pattern evolution and development, the technical challenge of genetically manipulating butterflies has slowed efforts to functionally characterize color pattern development genes. To identify candidate wing pigmentation genes, we used RNA sequencing to characterize transcription across multiple stages of butterfly wing development, and between different color pattern elements, in the painted lady butterfly Vanessa cardui. This allowed us to pinpoint genes specifically associated with red and black pigment patterns. To test the functions of a subset of genes associated with presumptive melanin pigmentation, we used clustered regularly interspaced short palindromic repeats (CRISPR)/Cas9 genome editing in four different butterfly genera. pale, Ddc, and yellow knockouts displayed reduction of melanin pigmentation, consistent with previous findings in other insects. Interestingly, however, yellow-d, ebony, and black knockouts revealed that these genes have localized effects on tuning the color of red, brown, and ochre pattern elements. These results point to previously undescribed mechanisms for modulating the color of specific wing pattern elements in butterflies, and provide an expanded portrait of the insect melanin pathway. PMID:28193726

  2. Genetic Basis of Melanin Pigmentation in Butterfly Wings.

    PubMed

    Zhang, Linlin; Martin, Arnaud; Perry, Michael W; van der Burg, Karin R L; Matsuoka, Yuji; Monteiro, Antónia; Reed, Robert D

    2017-04-01

    Despite the variety, prominence, and adaptive significance of butterfly wing patterns, surprisingly little is known about the genetic basis of wing color diversity. Even though there is intense interest in wing pattern evolution and development, the technical challenge of genetically manipulating butterflies has slowed efforts to functionally characterize color pattern development genes. To identify candidate wing pigmentation genes, we used RNA sequencing to characterize transcription across multiple stages of butterfly wing development, and between different color pattern elements, in the painted lady butterfly Vanessa cardui This allowed us to pinpoint genes specifically associated with red and black pigment patterns. To test the functions of a subset of genes associated with presumptive melanin pigmentation, we used clustered regularly interspaced short palindromic repeats (CRISPR)/Cas9 genome editing in four different butterfly genera. pale , Ddc , and yellow knockouts displayed reduction of melanin pigmentation, consistent with previous findings in other insects. Interestingly, however, yellow-d , ebony , and black knockouts revealed that these genes have localized effects on tuning the color of red, brown, and ochre pattern elements. These results point to previously undescribed mechanisms for modulating the color of specific wing pattern elements in butterflies, and provide an expanded portrait of the insect melanin pathway. Copyright © 2017 by the Genetics Society of America.

  3. Transcriptome analysis of the painted lady butterfly, Vanessa cardui during wing color pattern development.

    PubMed

    Connahs, Heidi; Rhen, Turk; Simmons, Rebecca B

    2016-03-31

    Butterfly wing color patterns are an important model system for understanding the evolution and development of morphological diversity and animal pigmentation. Wing color patterns develop from a complex network composed of highly conserved patterning genes and pigmentation pathways. Patterning genes are involved in regulating pigment synthesis however the temporal expression dynamics of these interacting networks is poorly understood. Here, we employ next generation sequencing to examine expression patterns of the gene network underlying wing development in the nymphalid butterfly, Vanessa cardui. We identified 9, 376 differentially expressed transcripts during wing color pattern development, including genes involved in patterning, pigmentation and gene regulation. Differential expression of these genes was highest at the pre-ommochrome stage compared to early pupal and late melanin stages. Overall, an increasing number of genes were down-regulated during the progression of wing development. We observed dynamic expression patterns of a large number of pigment genes from the ommochrome, melanin and also pteridine pathways, including contrasting patterns of expression for paralogs of the yellow gene family. Surprisingly, many patterning genes previously associated with butterfly pattern elements were not significantly up-regulated at any time during pupation, although many other transcription factors were differentially expressed. Several genes involved in Notch signaling were significantly up-regulated during the pre-ommochrome stage including slow border cells, bunched and pebbles; the function of these genes in the development of butterfly wings is currently unknown. Many genes involved in ecdysone signaling were also significantly up-regulated during early pupal and late melanin stages and exhibited opposing patterns of expression relative to the ecdysone receptor. Finally, a comparison across four butterfly transcriptomes revealed 28 transcripts common to all four species that have no known homologs in other metazoans. This study provides a comprehensive list of differentially expressed transcripts during wing development, revealing potential candidate genes that may be involved in regulating butterfly wing patterns. Some differentially expressed genes have no known homologs possibly representing genes unique to butterflies. Results from this study also indicate that development of nymphalid wing patterns may arise not only from melanin and ommochrome pigments but also the pteridine pigment pathway.

  4. Single master regulatory gene coordinates the evolution and development of butterfly color and iridescence

    PubMed Central

    Zhang, Linlin

    2017-01-01

    The optix gene has been implicated in butterfly wing pattern adaptation by genetic association, mapping, and expression studies. The actual developmental function of this gene has remained unclear, however. Here we used CRISPR/Cas9 genome editing to show that optix plays a fundamental role in nymphalid butterfly wing pattern development, where it is required for determination of all chromatic coloration. optix knockouts in four species show complete replacement of color pigments with melanins, with corresponding changes in pigment-related gene expression, resulting in black and gray butterflies. We also show that optix simultaneously acts as a switch gene for blue structural iridescence in some butterflies, demonstrating simple regulatory coordination of structural and pigmentary coloration. Remarkably, these optix knockouts phenocopy the recurring “black and blue” wing pattern archetype that has arisen on many independent occasions in butterflies. Here we demonstrate a simple genetic basis for structural coloration, and show that optix plays a deeply conserved role in butterfly wing pattern development. PMID:28923944

  5. Single master regulatory gene coordinates the evolution and development of butterfly color and iridescence.

    PubMed

    Zhang, Linlin; Mazo-Vargas, Anyi; Reed, Robert D

    2017-10-03

    The optix gene has been implicated in butterfly wing pattern adaptation by genetic association, mapping, and expression studies. The actual developmental function of this gene has remained unclear, however. Here we used CRISPR/Cas9 genome editing to show that optix plays a fundamental role in nymphalid butterfly wing pattern development, where it is required for determination of all chromatic coloration. optix knockouts in four species show complete replacement of color pigments with melanins, with corresponding changes in pigment-related gene expression, resulting in black and gray butterflies. We also show that optix simultaneously acts as a switch gene for blue structural iridescence in some butterflies, demonstrating simple regulatory coordination of structural and pigmentary coloration. Remarkably, these optix knockouts phenocopy the recurring "black and blue" wing pattern archetype that has arisen on many independent occasions in butterflies. Here we demonstrate a simple genetic basis for structural coloration, and show that optix plays a deeply conserved role in butterfly wing pattern development.

  6. Waiting in the wings: what can we learn about gene co-option from the diversification of butterfly wing patterns?

    PubMed

    Jiggins, Chris D; Wallbank, Richard W R; Hanly, Joseph J

    2017-02-05

    A major challenge is to understand how conserved gene regulatory networks control the wonderful diversity of form that we see among animals and plants. Butterfly wing patterns are an excellent example of this diversity. Butterfly wings form as imaginal discs in the caterpillar and are constructed by a gene regulatory network, much of which is conserved across the holometabolous insects. Recent work in Heliconius butterflies takes advantage of genomic approaches and offers insights into how the diversification of wing patterns is overlaid onto this conserved network. WntA is a patterning morphogen that alters spatial information in the wing. Optix is a transcription factor that acts later in development to paint specific wing regions red. Both of these loci fit the paradigm of conserved protein-coding loci with diverse regulatory elements and developmental roles that have taken on novel derived functions in patterning wings. These discoveries offer insights into the 'Nymphalid Ground Plan', which offers a unifying hypothesis for pattern formation across nymphalid butterflies. These loci also represent 'hotspots' for morphological change that have been targeted repeatedly during evolution. Both convergent and divergent evolution of a great diversity of patterns is controlled by complex alleles at just a few genes. We suggest that evolutionary change has become focused on one or a few genetic loci for two reasons. First, pre-existing complex cis-regulatory loci that already interact with potentially relevant transcription factors are more likely to acquire novel functions in wing patterning. Second, the shape of wing regulatory networks may constrain evolutionary change to one or a few loci. Overall, genomic approaches that have identified wing patterning loci in these butterflies offer broad insight into how gene regulatory networks evolve to produce diversity.This article is part of the themed issue 'Evo-devo in the genomics era, and the origins of morphological diversity'. © 2016 The Author(s).

  7. Waiting in the wings: what can we learn about gene co-option from the diversification of butterfly wing patterns?

    PubMed Central

    Wallbank, Richard W. R.; Hanly, Joseph J.

    2017-01-01

    A major challenge is to understand how conserved gene regulatory networks control the wonderful diversity of form that we see among animals and plants. Butterfly wing patterns are an excellent example of this diversity. Butterfly wings form as imaginal discs in the caterpillar and are constructed by a gene regulatory network, much of which is conserved across the holometabolous insects. Recent work in Heliconius butterflies takes advantage of genomic approaches and offers insights into how the diversification of wing patterns is overlaid onto this conserved network. WntA is a patterning morphogen that alters spatial information in the wing. Optix is a transcription factor that acts later in development to paint specific wing regions red. Both of these loci fit the paradigm of conserved protein-coding loci with diverse regulatory elements and developmental roles that have taken on novel derived functions in patterning wings. These discoveries offer insights into the ‘Nymphalid Ground Plan’, which offers a unifying hypothesis for pattern formation across nymphalid butterflies. These loci also represent ‘hotspots’ for morphological change that have been targeted repeatedly during evolution. Both convergent and divergent evolution of a great diversity of patterns is controlled by complex alleles at just a few genes. We suggest that evolutionary change has become focused on one or a few genetic loci for two reasons. First, pre-existing complex cis-regulatory loci that already interact with potentially relevant transcription factors are more likely to acquire novel functions in wing patterning. Second, the shape of wing regulatory networks may constrain evolutionary change to one or a few loci. Overall, genomic approaches that have identified wing patterning loci in these butterflies offer broad insight into how gene regulatory networks evolve to produce diversity. This article is part of the themed issue ‘Evo-devo in the genomics era, and the origins of morphological diversity’. PMID:27994126

  8. Structural or pigmentary? Origin of the distinctive white stripe on the blue wing of a Morpho butterfly.

    PubMed

    Yoshioka, Shinya; Kinoshita, Shuichi

    2006-01-22

    A few species of Morpho butterflies have a distinctive white stripe pattern on their structurally coloured blue wings. Since the colour pattern of a butterfly wing is formed as a mosaic of differently coloured scales, several questions naturally arise: are the microstructures the same between the blue and white scales? How is the distinctive whiteness produced, structurally or by means of pigmentation? To answer these questions, we have performed structural and optical investigations of the stripe pattern of a butterfly, Morpho cypris. It is found that besides the dorsal and ventral scale layers, the wing substrate also has the corresponding stripe pattern. Quantitative optical measurements and analysis using a simple model for the wing structure reveal the origin of the higher reflectance which makes the white stripe brighter.

  9. Distal-less induces elemental color patterns in Junonia butterfly wings.

    PubMed

    Dhungel, Bidur; Ohno, Yoshikazu; Matayoshi, Rie; Iwasaki, Mayo; Taira, Wataru; Adhikari, Kiran; Gurung, Raj; Otaki, Joji M

    2016-01-01

    The border ocellus, or eyespot, is a conspicuous color pattern element in butterfly wings. For two decades, it has been hypothesized that transcription factors such as Distal-less (Dll) are responsible for eyespot pattern development in butterfly wings, based on their expression in the prospective eyespots. In particular, it has been suggested that Dll is a determinant for eyespot size. However, functional evidence for this hypothesis has remained incomplete, due to technical difficulties. Here, we show that ectopically expressed Dll induces ectopic elemental color patterns in the adult wings of the blue pansy butterfly, Junonia orithya (Lepidoptera, Nymphalidae). Using baculovirus-mediated gene transfer, we misexpressed Dll protein fused with green fluorescent protein (GFP) in pupal wings, resulting in ectopic color patterns, but not the formation of intact eyespots. Induced changes included clusters of black and orange scales (a basic feature of eyespot patterns), black and gray scales, and inhibition of cover scale development. In contrast, ectopic expression of GFP alone did not induce any color pattern changes using the same baculovirus-mediated gene transfer system. These results suggest that Dll plays an instructive role in the development of color pattern elements in butterfly wings, although Dll alone may not be sufficient to induce a complete eyespot. This study thus experimentally supports the hypothesis of Dll function in eyespot development.

  10. apterous A specifies dorsal wing patterns and sexual traits in butterflies

    PubMed Central

    2018-01-01

    Butterflies have evolved different colour patterns on their dorsal and ventral wing surfaces to serve different signalling functions, yet the developmental mechanisms controlling surface-specific patterning are still unknown. Here, we mutate both copies of the transcription factor apterous in Bicyclus anynana butterflies using CRISPR/Cas9 and show that apterous A, expressed dorsally, functions both as a repressor and modifier of ventral wing colour patterns, as well as a promoter of dorsal sexual ornaments in males. We propose that the surface-specific diversification of wing patterns in butterflies proceeded via the co-option of apterous A or its downstream effectors into various gene regulatory networks involved in the differentiation of discrete wing traits. Further, interactions between apterous and sex-specific factors such as doublesex may have contributed to the origin of sexually dimorphic surface-specific patterns. Finally, we discuss the evolution of eyespot number diversity in the family Nymphalidae within the context of developmental constraints due to apterous regulation. PMID:29467265

  11. Pigmentary and photonic coloration mechanisms reveal taxonomic relationships of the Cattlehearts (Lepidoptera: Papilionidae: Parides)

    PubMed Central

    2014-01-01

    Background The colorful wing patterns of butterflies, a prime example of biodiversity, can change dramatically within closely related species. Wing pattern diversity is specifically present among papilionid butterflies. Whether a correlation between color and the evolution of these butterflies exists so far remained unsolved. Results We here investigate the Cattlehearts, Parides, a small Neotropical genus of papilionid butterflies with 36 members, the wings of which are marked by distinctly colored patches. By applying various physical techniques, we investigate the coloration toolkit of the wing scales. The wing scales contain two different, wavelength-selective absorbing pigments, causing pigmentary colorations. Scale ridges with multilayered lamellae, lumen multilayers or gyroid photonic crystals in the scale lumen create structural colors that are variously combined with these pigmentary colors. Conclusions The pigmentary and structural traits strongly correlate with the taxonomical distribution of Parides species. The experimental findings add crucial insight into the evolution of butterfly wing scales and show the importance of morphological parameter mapping for butterfly phylogenetics. PMID:25064167

  12. Spectrally tuned structural and pigmentary coloration of birdwing butterfly wing scales.

    PubMed

    Wilts, Bodo D; Matsushita, Atsuko; Arikawa, Kentaro; Stavenga, Doekele G

    2015-10-06

    The colourful wing patterns of butterflies play an important role for enhancing fitness; for instance, by providing camouflage, for interspecific mate recognition, or for aposematic display. Closely related butterfly species can have dramatically different wing patterns. The phenomenon is assumed to be caused by ecological processes with changing conditions, e.g. in the environment, and also by sexual selection. Here, we investigate the birdwing butterflies, Ornithoptera, the largest butterflies of the world, together forming a small genus in the butterfly family Papilionidae. The wings of these butterflies are marked by strongly coloured patches. The colours are caused by specially structured wing scales, which act as a chirped multilayer reflector, but the scales also contain papiliochrome pigments, which act as a spectral filter. The combined structural and pigmentary effects tune the coloration of the wing scales. The tuned colours are presumably important for mate recognition and signalling. By applying electron microscopy, (micro-)spectrophotometry and scatterometry we found that the various mechanisms of scale coloration of the different birdwing species strongly correlate with the taxonomical distribution of Ornithoptera species. © 2015 The Author(s).

  13. Spectrally tuned structural and pigmentary coloration of birdwing butterfly wing scales

    PubMed Central

    Wilts, Bodo D.; Matsushita, Atsuko; Arikawa, Kentaro; Stavenga, Doekele G.

    2015-01-01

    The colourful wing patterns of butterflies play an important role for enhancing fitness; for instance, by providing camouflage, for interspecific mate recognition, or for aposematic display. Closely related butterfly species can have dramatically different wing patterns. The phenomenon is assumed to be caused by ecological processes with changing conditions, e.g. in the environment, and also by sexual selection. Here, we investigate the birdwing butterflies, Ornithoptera, the largest butterflies of the world, together forming a small genus in the butterfly family Papilionidae. The wings of these butterflies are marked by strongly coloured patches. The colours are caused by specially structured wing scales, which act as a chirped multilayer reflector, but the scales also contain papiliochrome pigments, which act as a spectral filter. The combined structural and pigmentary effects tune the coloration of the wing scales. The tuned colours are presumably important for mate recognition and signalling. By applying electron microscopy, (micro-)spectrophotometry and scatterometry we found that the various mechanisms of scale coloration of the different birdwing species strongly correlate with the taxonomical distribution of Ornithoptera species. PMID:26446560

  14. Wnt signaling underlies evolution and development of the butterfly wing pattern symmetry systems.

    PubMed

    Martin, Arnaud; Reed, Robert D

    2014-11-15

    Most butterfly wing patterns are proposed to be derived from a set of conserved pattern elements known as symmetry systems. Symmetry systems are so-named because they are often associated with parallel color stripes mirrored around linear organizing centers that run between the anterior and posterior wing margins. Even though the symmetry systems are the most prominent and diverse wing pattern elements, their study has been confounded by a lack of knowledge regarding the molecular basis of their development, as well as the difficulty of drawing pattern homologies across species with highly derived wing patterns. Here we present the first molecular characterization of symmetry system development by showing that WntA expression is consistently associated with the major basal, discal, central, and external symmetry system patterns of nymphalid butterflies. Pharmacological manipulations of signaling gradients using heparin and dextran sulfate showed that pattern organizing centers correspond precisely with WntA, wingless, Wnt6, and Wnt10 expression patterns, thus suggesting a role for Wnt signaling in color pattern induction. Importantly, this model is supported by recent genetic and population genomic work identifying WntA as the causative locus underlying wing pattern variation within several butterfly species. By comparing the expression of WntA between nymphalid butterflies representing a range of prototypical symmetry systems, slightly deviated symmetry systems, and highly derived wing patterns, we were able to infer symmetry system homologies in several challenging cases. Our work illustrates how highly divergent morphologies can be derived from modifications to a common ground plan across both micro- and macro-evolutionary time scales. Copyright © 2014 Elsevier Inc. All rights reserved.

  15. Beneficial aerodynamic effect of wing scales on the climbing flight of butterflies.

    PubMed

    Slegers, Nathan; Heilman, Michael; Cranford, Jacob; Lang, Amy; Yoder, John; Habegger, Maria Laura

    2017-01-30

    It is hypothesized that butterfly wing scale geometry and surface patterning may function to improve aerodynamic efficiency. In order to investigate this hypothesis, a method to measure butterfly flapping kinematics optically over long uninhibited flapping sequences was developed. Statistical results for the climbing flight flapping kinematics of 11 butterflies, based on a total of 236 individual flights, both with and without their wing scales, are presented. Results show, that for each of the 11 butterflies, the mean climbing efficiency decreased after scales were removed. Data was reduced to a single set of differences of climbing efficiency using are paired t-test. Results show a mean decrease in climbing efficiency of 32.2% occurred with a 95% confidence interval of 45.6%-18.8%. Similar analysis showed that the flapping amplitude decreased by 7% while the flapping frequency did not show a significant difference. Results provide strong evidence that butterfly wing scale geometry and surface patterning improve butterfly climbing efficiency. The authors hypothesize that the wing scale's effect in measured climbing efficiency may be due to an improved aerodynamic efficiency of the butterfly and could similarly be used on flapping wing micro air vehicles to potentially achieve similar gains in efficiency.

  16. Characterisation and expression of microRNAs in developing wings of the neotropical butterfly Heliconius melpomene

    PubMed Central

    2011-01-01

    Background Heliconius butterflies are an excellent system for studies of adaptive convergent and divergent phenotypic traits. Wing colour patterns are used as signals to both predators and potential mates and are inherited in a Mendelian manner. The underlying genetic mechanisms of pattern formation have been studied for many years and shed light on broad issues, such as the repeatability of evolution. In Heliconius melpomene, the yellow hindwing bar is controlled by the HmYb locus. MicroRNAs (miRNAs) are important post-transcriptional regulators of gene expression that have key roles in many biological processes, including development. miRNAs could act as regulators of genes involved in wing development, patterning and pigmentation. For this reason we characterised miRNAs in developing butterfly wings and examined differences in their expression between colour pattern races. Results We sequenced small RNA libraries from two colour pattern races and detected 142 Heliconius miRNAs with homology to others found in miRBase. Several highly abundant miRNAs were differentially represented in the libraries between colour pattern races. These candidates were tested further using Northern blots, showing that differences in expression were primarily due to developmental stage rather than colour pattern. Assembly of sequenced reads to the HmYb region identified hme-miR-193 and hme-miR-2788; located 2380 bp apart in an intergenic region. These two miRNAs are expressed in wings and show an upregulation between 24 and 72 hours post-pupation, indicating a potential role in butterfly wing development. A search for miRNAs in all available H. melpomene BAC sequences (~ 2.5 Mb) did not reveal any other miRNAs and no novel miRNAs were predicted. Conclusions Here we describe the first butterfly miRNAs and characterise their expression in developing wings. Some show differences in expression across developing pupal stages and may have important functions in butterfly wing development. Two miRNAs were located in the HmYb region and were expressed in developing pupal wings. Future work will examine the expression of these miRNAs in different colour pattern races and identify miRNA targets among wing patterning genes. PMID:21266089

  17. Characterisation and expression of microRNAs in developing wings of the neotropical butterfly Heliconius melpomene.

    PubMed

    Surridge, Alison K; Lopez-Gomollon, Sara; Moxon, Simon; Maroja, Luana S; Rathjen, Tina; Nadeau, Nicola J; Dalmay, Tamas; Jiggins, Chris D

    2011-01-26

    Heliconius butterflies are an excellent system for studies of adaptive convergent and divergent phenotypic traits. Wing colour patterns are used as signals to both predators and potential mates and are inherited in a Mendelian manner. The underlying genetic mechanisms of pattern formation have been studied for many years and shed light on broad issues, such as the repeatability of evolution. In Heliconius melpomene, the yellow hindwing bar is controlled by the HmYb locus. MicroRNAs (miRNAs) are important post-transcriptional regulators of gene expression that have key roles in many biological processes, including development. miRNAs could act as regulators of genes involved in wing development, patterning and pigmentation. For this reason we characterised miRNAs in developing butterfly wings and examined differences in their expression between colour pattern races. We sequenced small RNA libraries from two colour pattern races and detected 142 Heliconius miRNAs with homology to others found in miRBase. Several highly abundant miRNAs were differentially represented in the libraries between colour pattern races. These candidates were tested further using Northern blots, showing that differences in expression were primarily due to developmental stage rather than colour pattern. Assembly of sequenced reads to the HmYb region identified hme-miR-193 and hme-miR-2788; located 2380 bp apart in an intergenic region. These two miRNAs are expressed in wings and show an upregulation between 24 and 72 hours post-pupation, indicating a potential role in butterfly wing development. A search for miRNAs in all available H. melpomene BAC sequences (~2.5 Mb) did not reveal any other miRNAs and no novel miRNAs were predicted. Here we describe the first butterfly miRNAs and characterise their expression in developing wings. Some show differences in expression across developing pupal stages and may have important functions in butterfly wing development. Two miRNAs were located in the HmYb region and were expressed in developing pupal wings. Future work will examine the expression of these miRNAs in different colour pattern races and identify miRNA targets among wing patterning genes.

  18. Tungstate-induced color-pattern modifications of butterfly wings are independent of stress response and ecdysteroid effect.

    PubMed

    Otaki, Joji M; Ogasawara, Tsuyoshi; Yamamoto, Haruhiko

    2005-06-01

    Systemic injections of sodium tungstate, a protein-tyrosine phosphatase (PTPase) inhibitor, to pupae immediately after pupation have been shown to efficiently produce characteristic color-pattern modifications on the wings of many species of butterflies. Here we demonstrated that the tungstate-induced modification pattern was entirely different from other chemically-induced ones in a species of nymphalid butterfly Junonia (Precis) orithya. In this species, the systemic injections of tungstate produced characteristic expansion of black area and shrinkage of white area together with the move of parafocal elements toward the wing base. Overall, pattern boundaries became obscure. In contrast, an entirely different modification pattern, overall darkening of wings, was observed by the injections of stress-inducing chemicals, thapsigargin, ionomycin, or geldanamycin, to pupae under the rearing conditions for the adult summer form. On the ventral wings, this darkening was due to an increase of the proportion of peppered dark scales, which was reminiscent of the natural fall form of this species. Under the same rearing conditions, the injections of ecdysteroid, which is a well-known hormone being responsible for the seasonal polyphenism of nymphalid butterflies, yielded overall expansion of orange area especially around eyespots. Taken together, we conclude that the tungstate-induced modifications are clearly distinguishable from those of stress response and ecdysteroid effect. This conclusion then suggests that the putative PTPase signaling pathway that is sensitive to tungstate uniquely contributes to the wing-wide color-pattern development in butterflies.

  19. Confocal imaging of butterfly tissue.

    PubMed

    Brunetti, Craig R

    2014-01-01

    To understand the molecular events responsible for morphological change requires the ability to examine gene expression in a wide range of organisms in addition to model systems to determine how the differences in gene expression correlate with phenotypic differences. There are approximately 12,000 species of butterflies, most, with distinct patterns on their wings. The most important tool for studying gene expression in butterflies is confocal imaging of butterfly tissue by indirect immunofluorescence using either cross-reactive antibodies from closely related species such as Drosophila or developing butterfly-specific antibodies. In this report, we describe how indirect immunofluorescence protocols can be used to visualize protein expression patterns on the butterfly wing imaginal disc and butterfly embryo.

  20. Baculovirus-mediated gene transfer in butterfly wings in vivo: an efficient expression system with an anti-gp64 antibody.

    PubMed

    Dhungel, Bidur; Ohno, Yoshikazu; Matayoshi, Rie; Otaki, Joji M

    2013-03-25

    Candidate genes for color pattern formation in butterfly wings have been known based on gene expression patterns since the 1990s, but their functions remain elusive due to a lack of a functional assay. Several methods of transferring and expressing a foreign gene in butterfly wings have been reported, but they have suffered from low success rates or low expression levels. Here, we developed a simple, practical method to efficiently deliver and express a foreign gene using baculovirus-mediated gene transfer in butterfly wings in vivo. A recombinant baculovirus containing a gene for green fluorescent protein (GFP) was injected into pupae of the blue pansy butterfly Junonia orithya (Nymphalidae). GFP fluorescence was detected in the pupal wings and other body parts of the injected individuals three to five days post-injection at various degrees of fluorescence. We obtained a high GFP expression rate at relatively high virus titers, but it was associated with pupal death before color pattern formation in wings. To reduce the high mortality rate caused by the baculovirus treatment, we administered an anti-gp64 antibody, which was raised against baculovirus coat protein gp64, to infected pupae after the baculovirus injection. This treatment greatly reduced the mortality rate of the infected pupae. GFP fluorescence was observed in pupal and adult wings and other body parts of the antibody-treated individuals at various degrees of fluorescence. Importantly, we obtained completely developed wings with a normal color pattern, in which fluorescent signals originated directly from scales or the basal membrane after the removal of scales. GFP fluorescence in wing tissues spatially coincided with anti-GFP antibody staining, confirming that the fluorescent signals originated from the expressed GFP molecules. Our baculovirus-mediated gene transfer system with an anti-gp64 antibody is reasonably efficient, and it can be an invaluable tool to transfer, express, and functionally examine foreign genes in butterfly wings and also in other non-model insect systems.

  1. Multiple scaled disorder in the photonic structure of Morpho rhetenor butterfly

    NASA Astrophysics Data System (ADS)

    Boulenguez, J.; Berthier, S.; Leroy, F.

    2012-03-01

    The iridescence of Morpho rhetenor butterfly is known to result from a photonic structure on wing scales, where multilayer interference and grating diffraction occur simultaneously. We characterize the disorder at the photonic structure length scale and at the butterfly scale. We measure the scattering pattern of the wing. Through RCWA and 1st Born approximation models, we link the different disorders to different features in the scattering patterns.

  2. A simulation study of mutations in the genetic regulatory hierarchy for butterfly eyespot focus determination.

    PubMed

    Marcus, Jeffrey M; Evans, Travis M

    2008-09-01

    The color patterns on the wings of butterflies have been an important model system in evolutionary developmental biology. A recent computational model tested genetic regulatory hierarchies hypothesized to underlie the formation of butterfly eyespot foci [Evans, T.M., Marcus, J.M., 2006. A simulation study of the genetic regulatory hierarchy for butterfly eyespot focus determination. Evol. Dev. 8, 273-283]. The computational model demonstrated that one proposed hierarchy was incapable of reproducing the known patterns of gene expression associated with eyespot focus determination in wild-type butterflies, but that two slightly modified alternative hierarchies were capable of reproducing all of the known gene expressions patterns. Here we extend the computational models previously implemented in Delphi 2.0 to two mutants derived from the squinting bush brown butterfly (Bicyclus anynana). These two mutants, comet and Cyclops, have aberrantly shaped eyespot foci that are produced by different mechanisms. The comet mutation appears to produce a modified interaction between the wing margin and the eyespot focus that results in a series of comet-shaped eyespot foci. The Cyclops mutation causes the failure of wing vein formation between two adjacent wing-cells and the fusion of two adjacent eyespot foci to form a single large elongated focus in their place. The computational approach to modeling pattern formation in these mutants allows us to make predictions about patterns of gene expression, which are largely unstudied in butterfly mutants. It also suggests a critical experiment that will allow us to distinguish between two hypothesized genetic regulatory hierarchies that may underlie all butterfly eyespot foci.

  3. A Simulation Study of Mutations in the Genetic Regulatory Hierarchy for Butterfly Eyespot Focus Determination

    PubMed Central

    Marcus, Jeffrey M.; Evans, Travis M.

    2008-01-01

    The color patterns on the wings of butterflies have been an important model system in evolutionary developmental biology. A recent computational model tested genetic regulatory hierarchies hypothesized to underlie the formation of butterfly eyespot foci (Evans and Marcus, 2006). The computational model demonstrated that one proposed hierarchy was incapable of reproducing the known patterns of gene expression associated with eyespot focus determination in wild-type butterflies, but that two slightly modified alternative hierarchies were capable of reproducing all of the known gene expressions patterns. Here we extend the computational models previously implemented in Delphi 2.0 to two mutants derived from the squinting bush brown butterfly (Bicyclus anynana). These two mutants, comet and Cyclops, have aberrantly shaped eyespot foci that are produced by different mechanisms. The comet mutation appears to produce a modified interaction between the wing margin and the eyespot focus that results in a series of comet-shaped eyespot foci. The Cyclops mutation causes the failure of wing vein formation between two adjacent wing-cells and the fusion of two adjacent eyespot foci to form a single large elongated focus in their place. The computational approach to modeling pattern formation in these mutants allows us to make predictions about patterns of gene expression, which are largely unstudied in butterfly mutants. It also suggests a critical experiment that will allow us to distinguish between two hypothesized genetic regulatory hierarchies that may underlie all butterfly eyespot foci. PMID:18586070

  4. Spontaneous long-range calcium waves in developing butterfly wings.

    PubMed

    Ohno, Yoshikazu; Otaki, Joji M

    2015-03-25

    Butterfly wing color patterns emerge as the result of a regular arrangement of scales produced by epithelial scale cells at the pupal stage. These color patterns and scale arrangements are coordinated throughout the wing. However, the mechanism by which the development of scale cells is controlled across the entire wing remains elusive. In the present study, we used pupal wings of the blue pansy butterfly, Junonia orithya, which has distinct eyespots, to examine the possible involvement of Ca(2+) waves in wing development. Here, we demonstrate that the developing pupal wing tissue of the blue pansy butterfly displayed spontaneous low-frequency Ca(2+) waves in vivo that propagated slowly over long distances. Some waves appeared to be released from the immediate peripheries of the prospective eyespot and discal spot, though it was often difficult to identify the specific origins of these waves. Physical damage, which is known to induce ectopic eyespots, led to the radiation of Ca(2+) waves from the immediate periphery of the damaged site. Thapsigargin, which is a specific inhibitor of Ca(2+)-ATPases in the endoplasmic reticulum, induced an acute increase in cytoplasmic Ca(2+) levels and halted the spontaneous Ca(2+) waves. Additionally, thapsigargin-treated wings showed incomplete scale development as well as other scale and color pattern abnormalities. We identified a novel form of Ca(2+) waves, spontaneous low-frequency slow waves, which travel over exceptionally long distances. Our results suggest that spontaneous Ca(2+) waves play a critical role in the coordinated development of scale arrangements and possibly in color pattern formation in butterflies.

  5. Genetic constraints on wing pattern variation in Lycaeides butterflies: A case study on mapping complex, multifaceted traits in structured populations.

    PubMed

    Lucas, Lauren K; Nice, Chris C; Gompert, Zachariah

    2018-03-13

    Patterns of phenotypic variation within and among species can be shaped and constrained by trait genetic architecture. This is particularly true for complex traits, such as butterfly wing patterns, that consist of multiple elements. Understanding the genetics of complex trait variation across species boundaries is difficult, as it necessitates mapping in structured populations and can involve many loci with small or variable phenotypic effects. Here, we investigate the genetic architecture of complex wing pattern variation in Lycaeides butterflies as a case study of mapping multivariate traits in wild populations that include multiple nominal species or groups. We identify conserved modules of integrated wing pattern elements within populations and species. We show that trait covariances within modules have a genetic basis and thus represent genetic constraints that can channel evolution. Consistent with this, we find evidence that evolutionary changes in wing patterns among populations and species occur in the directions of genetic covariances within these groups. Thus, we show that genetic constraints affect patterns of biological diversity (wing pattern) in Lycaeides, and we provide an analytical template for similar work in other systems. © 2018 John Wiley & Sons Ltd.

  6. Unique wing scale photonics of male Rajah Brooke's birdwing butterflies.

    PubMed

    Wilts, Bodo D; Giraldo, Marco A; Stavenga, Doekele G

    2016-01-01

    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. 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. 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.

  7. System-dependent regulations of colour-pattern development: a mutagenesis study of the pale grass blue butterfly

    PubMed Central

    Iwata, Masaki; Hiyama, Atsuki; Otaki, Joji M.

    2013-01-01

    Developmental studies on wing colour patterns have been performed in nymphalid butterflies, but efficient genetic manipulations, including mutagenesis, have not been well established. Here, we have performed mutagenesis experiments in a lycaenid butterfly, the pale grass blue Zizeeria maha, to produce colour-pattern mutants. We fed the P-generation larvae an artificial diet containing the mutagen ethyl methane sulfonate (EMS), and the F1- and F2-generation adults showed various aberrant colour patterns: dorsoventral transformation, anterioposterior background colouration gap, weak contrast, disarrangement of spots, reduction of the size of spots, loss of spots, fusion of spots, and ectopic spots. Among them, the disarrangement, reduction, and loss of spots were likely produced by the coordinated changes of many spots of a single wing around the discal spot in a system-dependent manner, demonstrating the existence of the central symmetry system. The present study revealed multiple genetic regulations for system-dependent and wing-wide colour-pattern determination in lycaenid butterflies. PMID:23917124

  8. Baculovirus-mediated gene transfer in butterfly wings in vivo: an efficient expression system with an anti-gp64 antibody

    PubMed Central

    2013-01-01

    Background Candidate genes for color pattern formation in butterfly wings have been known based on gene expression patterns since the 1990s, but their functions remain elusive due to a lack of a functional assay. Several methods of transferring and expressing a foreign gene in butterfly wings have been reported, but they have suffered from low success rates or low expression levels. Here, we developed a simple, practical method to efficiently deliver and express a foreign gene using baculovirus-mediated gene transfer in butterfly wings in vivo. Results A recombinant baculovirus containing a gene for green fluorescent protein (GFP) was injected into pupae of the blue pansy butterfly Junonia orithya (Nymphalidae). GFP fluorescence was detected in the pupal wings and other body parts of the injected individuals three to five days post-injection at various degrees of fluorescence. We obtained a high GFP expression rate at relatively high virus titers, but it was associated with pupal death before color pattern formation in wings. To reduce the high mortality rate caused by the baculovirus treatment, we administered an anti-gp64 antibody, which was raised against baculovirus coat protein gp64, to infected pupae after the baculovirus injection. This treatment greatly reduced the mortality rate of the infected pupae. GFP fluorescence was observed in pupal and adult wings and other body parts of the antibody-treated individuals at various degrees of fluorescence. Importantly, we obtained completely developed wings with a normal color pattern, in which fluorescent signals originated directly from scales or the basal membrane after the removal of scales. GFP fluorescence in wing tissues spatially coincided with anti-GFP antibody staining, confirming that the fluorescent signals originated from the expressed GFP molecules. Conclusions Our baculovirus-mediated gene transfer system with an anti-gp64 antibody is reasonably efficient, and it can be an invaluable tool to transfer, express, and functionally examine foreign genes in butterfly wings and also in other non-model insect systems. PMID:23522444

  9. Wingless is a positive regulator of eyespot color patterns in Bicyclus anynana butterflies.

    PubMed

    Özsu, Nesibe; Chan, Qian Yi; Chen, Bin; Gupta, Mainak Das; Monteiro, Antónia

    2017-09-01

    Eyespot patterns of nymphalid butterflies are an example of a novel trait yet, the developmental origin of eyespots is still not well understood. Several genes have been associated with eyespot development but few have been tested for function. One of these genes is the signaling ligand, wingless, which is expressed in the eyespot centers during early pupation and may function in eyespot signaling and color ring differentiation. Here we tested the function of wingless in wing and eyespot development by down-regulating it in transgenic Bicyclus anynana butterflies via RNAi driven by an inducible heat-shock promoter. Heat-shocks applied during larval and early pupal development led to significant decreases in wingless mRNA levels and to decreases in eyespot size and wing size in adult butterflies. We conclude that wingless is a positive regulator of eyespot and wing development in B. anynana butterflies. Copyright © 2017 Elsevier Inc. All rights reserved.

  10. Varying and unchanging whiteness on the wings of dusk-active and shade-inhabiting Carystoides escalantei butterflies.

    PubMed

    Ge, Dengteng; Wu, Gaoxiang; Yang, Lili; Kim, Hye-Na; Hallwachs, Winnie; Burns, John M; Janzen, Daniel H; Yang, Shu

    2017-07-11

    Whiteness, although frequently apparent on the wings, legs, antennae, or bodies of many species of moths and butterflies, along with other colors and shades, has often escaped our attention. Here, we investigate the nanostructure and microstructure of white spots on the wings of Carystoides escalantei , a dusk-active and shade-inhabiting Costa Rican rain forest butterfly (Hesperiidae). On both males and females, two types of whiteness occur: angle dependent (dull or bright) and angle independent, which differ in the microstructure, orientation, and associated properties of their scales. Some spots on the male wings are absent from the female wings. Whether the angle-dependent whiteness is bright or dull depends on the observation directions. The angle-dependent scales also show enhanced retro-reflection. We speculate that the biological functions and evolution of Carystoides spot patterns, scale structures, and their varying whiteness are adaptations to butterfly's low light habitat and to airflow experienced on the wing base vs. wing tip.

  11. A Single-Wing Removal Method to Assess Correspondence Between Gene Expression and Phenotype in Butterflies: The Case of Distal-less.

    PubMed

    Adhikari, Kiran; Otaki, Joji M

    2016-02-01

    It is often desirable but difficult to retrieve information on the mature phenotype of an immature tissue sample that has been subjected to gene expression analysis. This problem cannot be ignored when individual variation within a species is large. To circumvent this problem in the butterfly wing system, we developed a new surgical method for removing a single forewing from a pupa using Junonia orithya; the operated pupa was left to develop to an adult without eclosion. The removed right forewing was subjected to gene expression analysis, whereas the non-removed left forewing was examined for color patterns. As a test case, we focused on Distal-less (Dll), which likely plays an active role in inducing elemental patterns, including eyespots. The Dll expression level in forewings was paired with eyespot size data from the same individual. One third of the operated pupae survived and developed wing color patterns. Dll expression levels were significantly higher in males than in females, although male eyespots were smaller in size than female eyespots. Eyespot size data showed weak but significant correlations with the Dll expression level in females. These results demonstrate that a single-wing removal method was successfully applied to the butterfly wing system and suggest the weak and non-exclusive contribution of Dll to eyespot size determination in this butterfly. Our novel methodology for establishing correspondence between gene expression and phenotype can be applied to other candidate genes for color pattern development in butterflies. Conceptually similar methods may also be applicable in other developmental systems.

  12. The Functional Basis of Wing Patterning in Heliconius Butterflies: The Molecules Behind Mimicry

    PubMed Central

    Kronforst, Marcus R.; Papa, Riccardo

    2015-01-01

    Wing-pattern mimicry in butterflies has provided an important example of adaptation since Charles Darwin and Alfred Russell Wallace proposed evolution by natural selection >150 years ago. The neotropical butterfly genus Heliconius played a central role in the development of mimicry theory and has since been studied extensively in the context of ecology and population biology, behavior, and mimicry genetics. Heliconius species are notable for their diverse color patterns, and previous crossing experiments revealed that much of this variation is controlled by a small number of large-effect, Mendelian switch loci. Recent comparative analyses have shown that the same switch loci control wing-pattern diversity throughout the genus, and a number of these have now been positionally cloned. Using a combination of comparative genetic mapping, association tests, and gene expression analyses, variation in red wing patterning throughout Heliconius has been traced back to the action of the transcription factor optix. Similarly, the signaling ligand WntA has been shown to control variation in melanin patterning across Heliconius and other butterflies. Our understanding of the molecular basis of Heliconius mimicry is now providing important insights into a variety of additional evolutionary phenomena, including the origin of supergenes, the interplay between constraint and evolvability, the genetic basis of convergence, the potential for introgression to facilitate adaptation, the mechanisms of hybrid speciation in animals, and the process of ecological speciation. PMID:25953905

  13. Spatial patterns of correlated scale size and scale color in relation to color pattern elements in butterfly wings.

    PubMed

    Iwata, Masaki; Otaki, Joji M

    2016-02-01

    Complex butterfly wing color patterns are coordinated throughout a wing by unknown mechanisms that provide undifferentiated immature scale cells with positional information for scale color. Because there is a reasonable level of correspondence between the color pattern element and scale size at least in Junonia orithya and Junonia oenone, a single morphogenic signal may contain positional information for both color and size. However, this color-size relationship has not been demonstrated in other species of the family Nymphalidae. Here, we investigated the distribution patterns of scale size in relation to color pattern elements on the hindwings of the peacock pansy butterfly Junonia almana, together with other nymphalid butterflies, Vanessa indica and Danaus chrysippus. In these species, we observed a general decrease in scale size from the basal to the distal areas, although the size gradient was small in D. chrysippus. Scales of dark color in color pattern elements, including eyespot black rings, parafocal elements, and submarginal bands, were larger than those of their surroundings. Within an eyespot, the largest scales were found at the focal white area, although there were exceptional cases. Similarly, ectopic eyespots that were induced by physical damage on the J. almana background area had larger scales than in the surrounding area. These results are consistent with the previous finding that scale color and size coordinate to form color pattern elements. We propose a ploidy hypothesis to explain the color-size relationship in which the putative morphogenic signal induces the polyploidization (genome amplification) of immature scale cells and that the degrees of ploidy (gene dosage) determine scale color and scale size simultaneously in butterfly wings. Copyright © 2015 Elsevier Ltd. All rights reserved.

  14. Differential involvement of Hedgehog signaling in butterfly wing and eyespot development.

    PubMed

    Tong, Xiaoling; Lindemann, Anna; Monteiro, Antónia

    2012-01-01

    Butterfly eyespots may have evolved from the recruitment of pre-existent gene circuits or regulatory networks into novel locations on the wing. Gene expression data suggests one such circuit, the Hedgehog (Hh) signaling pathway and its target gene engrailed (en), was recruited from a role in patterning the anterior-posterior insect wing axis to a role patterning butterfly eyespots. However, while Junonia coenia expresses hh and en both in the posterior compartment of the wing and in eyespot centers, Bicyclus anynana lacks hh eyespot-specific expression. This suggests that Hh signaling may not be functioning in eyespot development in either species or that it functions in J. coenia but not in B. anynana. In order to test these hypotheses, we performed functional tests of Hh signaling in these species. We investigated the effects of Hh protein sequestration during the larval stage on en expression levels, and on wing size and eyespot size in adults. Hh sequestration led to significantly reduced en expression and to significantly smaller wings and eyespots in both species. But while eyespot size in B. anynana was reduced proportionately to wing size, in J. coenia, eyespots were reduced disproportionately, indicating an independent role of Hh signaling in eyespot development in J. coenia. We conclude that while Hh signaling retains a conserved role in promoting wing growth across nymphalid butterflies, it plays an additional role in eyespot development in some, but not all, lineages of nymphalid butterflies. We discuss our findings in the context of alternative evolutionary scenarios that led to the differential expression of hh and other Hh pathway signaling members across nymphalid species.

  15. Physiologically induced color-pattern changes in butterfly wings: mechanistic and evolutionary implications.

    PubMed

    Otaki, Joji M

    2008-07-01

    A mechanistic understanding of the butterfly wing color-pattern determination can be facilitated by experimental pattern changes. Here I review physiologically induced color-pattern changes in nymphalid butterflies and their mechanistic and evolutionary implications. A type of color-pattern change can be elicited by elemental changes in size and position throughout the wing, as suggested by the nymphalid groundplan. These changes of pattern elements are bi-directional and bi-sided dislocation toward or away from eyespot foci and in both proximal and distal sides of the foci. The peripheral elements are dislocated even in the eyespot-less compartments. Anterior spots are more severely modified, suggesting the existence of an anterior-posterior gradient. In one species, eyespots are transformed into white spots with remnant-like orange scales, and such patterns emerge even at the eyespot-less "imaginary" foci. A series of these color-pattern modifications probably reveal "snap-shots" of a dynamic morphogenic signal due to heterochronic uncoupling between the signaling and reception steps. The conventional gradient model can be revised to account for these observed color-pattern changes.

  16. Integration of wings and their eyespots in the speckled wood butterfly Pararge aegeria.

    PubMed

    Breuker, Casper J; Gibbs, Melanie; Van Dyck, Hans; Brakefield, Paul M; Klingenberg, Christian Peter; Van Dongen, Stefan

    2007-07-15

    We investigated both the phenotypic and developmental integration of eyespots on the fore- and hindwings of speckled wood butterflies Pararge aegeria. Eyespots develop within a framework of wing veins, which may not only separate eyespots developmentally, but may at the same time also integrate them by virtue of being both signalling sources and barriers during eyespot development. We therefore specifically investigated the interaction between wing venation patterns and eyespot integration. Phenotypic covariation among eyespots was very high, but only eyespots in neighbouring wing cells and in homologous wing cells on different wing surfaces were developmentally integrated. This can be explained by the fact that the wing cells of these eyespots share one or more wing veins. The wing venation patterns of fore- and hindwings were highly integrated, both phenotypically and developmentally. This did not affect overall developmental integration of the eyespots. The adaptive significance of integration patterns is discussed and more specifically we stress the need to conduct studies on phenotypic plasticity of integration.

  17. Molecular basis of wing coloration in a Batesian mimic butterfly, Papilio polytes

    PubMed Central

    Nishikawa, Hideki; Iga, Masatoshi; Yamaguchi, Junichi; Saito, Kazuki; Kataoka, Hiroshi; Suzuki, Yutaka; Sugano, Sumio; Fujiwara, Haruhiko

    2013-01-01

    Batesian mimicry protects animals from predators through resemblance with distasteful models in shape, color pattern, or behavior. To elucidate the wing coloration mechanisms involved in the mimicry, we investigated chemical composition and gene expression of the pale yellow and red pigments of a swallowtail butterfly, Papilio polytes, whose females mimic the unpalatable butterfly Pachliopta aristolochiae. Using LC/MS, we showed that the pale yellow wing regions in non-mimetic females consist of kynurenine and N-β-alanyldopamine (NBAD). Moreover, qRT-PCR showed that kynurenine/NBAD biosynthetic genes were upregulated in these regions in non-mimetic females. However, these pigments were absent in mimetic females. RNA-sequencing showed that kynurenine/NBAD synthesis and Toll signaling genes were upregulated in the red spots specific to mimetic female wings. These results demonstrated that drastic changes in gene networks in the red and pale yellow regions can switch wing color patterns between non-mimetic and mimetic females of P. polytes. PMID:24212474

  18. Molecular basis of wing coloration in a Batesian mimic butterfly, Papilio polytes.

    PubMed

    Nishikawa, Hideki; Iga, Masatoshi; Yamaguchi, Junichi; Saito, Kazuki; Kataoka, Hiroshi; Suzuki, Yutaka; Sugano, Sumio; Fujiwara, Haruhiko

    2013-11-11

    Batesian mimicry protects animals from predators through resemblance with distasteful models in shape, color pattern, or behavior. To elucidate the wing coloration mechanisms involved in the mimicry, we investigated chemical composition and gene expression of the pale yellow and red pigments of a swallowtail butterfly, Papilio polytes, whose females mimic the unpalatable butterfly Pachliopta aristolochiae. Using LC/MS, we showed that the pale yellow wing regions in non-mimetic females consist of kynurenine and N-β-alanyldopamine (NBAD). Moreover, qRT-PCR showed that kynurenine/NBAD biosynthetic genes were upregulated in these regions in non-mimetic females. However, these pigments were absent in mimetic females. RNA-sequencing showed that kynurenine/NBAD synthesis and Toll signaling genes were upregulated in the red spots specific to mimetic female wings. These results demonstrated that drastic changes in gene networks in the red and pale yellow regions can switch wing color patterns between non-mimetic and mimetic females of P. polytes.

  19. A major gene controls mimicry and crypsis in butterflies and moths

    PubMed Central

    Nadeau, Nicola J.; Pardo-Diaz, Carolina; Whibley, Annabel; Supple, Megan; Saenko, Suzanne V.; Wallbank, Richard W. R.; Wu, Grace C.; Maroja, Luana; Ferguson, Laura; Hanly, Joseph J.; Hines, Heather; Salazar, Camilo; Merrill, Richard; Dowling, Andrea; ffrench-Constant, Richard; Llaurens, Violaine; Joron, Mathieu; McMillan, W. Owen; Jiggins, Chris D.

    2016-01-01

    The wing patterns of butterflies and moths (Lepidoptera) are diverse and striking examples of evolutionary diversification by natural selection1,2. Lepidopteran wing colour patterns are a key innovation, consisting of arrays of coloured scales. We still lack a general understanding of how these patterns are controlled and if there is any commonality across the 160,000 moth and 17,000 butterfly species. Here, we identify a gene, cortex, through fine-scale mapping using population genomics and gene expression analyses, which regulates pattern switches in multiple species across the mimetic radiation in Heliconius butterflies. cortex belongs to a fast evolving subfamily of the otherwise highly conserved fizzy family of cell cycle regulators3, suggesting that it most likely regulates pigmentation patterning through regulation of scale cell development. In parallel with findings in the peppered moth (Biston betularia)4, our results suggest that this mechanism is common within Lepidoptera and that cortex has become a major target for natural selection acting on colour and pattern variation in this group of insects. PMID:27251285

  20. The gene cortex controls mimicry and crypsis in butterflies and moths.

    PubMed

    Nadeau, Nicola J; Pardo-Diaz, Carolina; Whibley, Annabel; Supple, Megan A; Saenko, Suzanne V; Wallbank, Richard W R; Wu, Grace C; Maroja, Luana; Ferguson, Laura; Hanly, Joseph J; Hines, Heather; Salazar, Camilo; Merrill, Richard M; Dowling, Andrea J; ffrench-Constant, Richard H; Llaurens, Violaine; Joron, Mathieu; McMillan, W Owen; Jiggins, Chris D

    2016-06-02

    The wing patterns of butterflies and moths (Lepidoptera) are diverse and striking examples of evolutionary diversification by natural selection. Lepidopteran wing colour patterns are a key innovation, consisting of arrays of coloured scales. We still lack a general understanding of how these patterns are controlled and whether this control shows any commonality across the 160,000 moth and 17,000 butterfly species. Here, we use fine-scale mapping with population genomics and gene expression analyses to identify a gene, cortex, that regulates pattern switches in multiple species across the mimetic radiation in Heliconius butterflies. cortex belongs to a fast-evolving subfamily of the otherwise highly conserved fizzy family of cell-cycle regulators, suggesting that it probably regulates pigmentation patterning by regulating scale cell development. In parallel with findings in the peppered moth (Biston betularia), our results suggest that this mechanism is common within Lepidoptera and that cortex has become a major target for natural selection acting on colour and pattern variation in this group of insects.

  1. Ancient Wings: animating the evolution of butterfly wing patterns.

    PubMed

    Arbesman, Samuel; Enthoven, Leo; Monteiro, Antónia

    2003-10-01

    Character optimization methods can be used to reconstruct ancestral states at the internal nodes of phylogenetic trees. However, seldom are these ancestral states visualized collectively. Ancient Wings is a computer program that provides a novel method of visualizing the evolution of several morphological traits simultaneously. It allows users to visualize how the ventral hindwing pattern of 54 butterflies in the genus Bicyclus may have changed over time. By clicking on each of the nodes within the evolutionary tree, the user can see an animation of how wing size, eyespot size, and eyespot position relative the wing margin, have putatively evolved as a collective whole. Ancient Wings may be used as a pedagogical device as well as a research tool for hypothesis-generation in the fields of evolutionary, ecological, and developmental biology.

  2. Waterproof and translucent wings at the same time: problems and solutions in butterflies.

    PubMed

    Goodwyn, Pablo Perez; Maezono, Yasunori; Hosoda, Naoe; Fujisaki, Kenji

    2009-07-01

    Although the colour of butterflies attracts the most attention, the waterproofing properties of their wings are also extremely interesting. Most butterfly wings are considered "super-hydrophobic" because the contact angle (CA) with a water drop exceeds 150 degrees. Usually, butterfly wings are covered with strongly overlapping scales; however, in the case of transparent or translucent wings, scale cover is reduced; thus, the hydrophobicity could be affected. Here, we present a comparative analysis of wing hydrophobicity and its dependence on morphology for two species with translucent wings Parantica sita (Nymphalidae) and Parnassius glacialis (Papilionidae). These species have very different life histories: P. sita lives for up to 6 months as an adult and migrates over long distance, whereas P. glacialis lives for less than 1 month and does not migrate. We measured the water CA and analysed wing morphology with scanning electron microscopy and atomic force microscopy. P. sita has super-hydrophobic wing surfaces, with CA > 160 degrees, whereas P. glacialis did not (CA = 100-135 degrees). Specialised scales were found on the translucent portions of P. sita wings. These scales were ovoid and much thinner than common scales, erect at about 30 degrees, and leaving up to 80% of the wing surface uncovered. The underlying bare wing surface had a remarkable pattern of ridges and knobs. P. glacialis also had over 80% of the wing surface uncovered, but the scales were either setae-like or spade-like. The bare surface of the wing had an irregular wavy smooth pattern. We suggest a mode of action that allows this super-hydrophobic effect with an incompletely covered wing surface. The scales bend, but do not collapse, under the pressure of a water droplet, and the elastic recovery of the structure at the borders of the droplet allows a high apparent CA. Thus, P. sita can be translucent without losing its waterproof properties. This characteristic is likely necessary for the long life and migration of this species. This is the first study of some of the effects on the hydrophobicity of translucency through scales' cover reduction in butterfly wings and on the morphology associated with improved waterproofing.

  3. Live Cell Imaging of Butterfly Pupal and Larval Wings In Vivo

    PubMed Central

    Ohno, Yoshikazu; Otaki, Joji M.

    2015-01-01

    Butterfly wing color patterns are determined during the late larval and early pupal stages. Characterization of wing epithelial cells at these stages is thus critical to understand how wing structures, including color patterns, are determined. Previously, we successfully recorded real-time in vivo images of developing butterfly wings over time at the tissue level. In this study, we employed similar in vivo fluorescent imaging techniques to visualize developing wing epithelial cells in the late larval and early pupal stages 1 hour post-pupation. Both larval and pupal epithelial cells were rich in mitochondria and intracellular networks of endoplasmic reticulum, suggesting high metabolic activities, likely in preparation for cellular division, polyploidization, and differentiation. Larval epithelial cells in the wing imaginal disk were relatively large horizontally and tightly packed, whereas pupal epithelial cells were smaller and relatively loosely packed. Furthermore, larval cells were flat, whereas pupal cells were vertically elongated as deep as 130 μm. In pupal cells, many endosome-like or autophagosome-like structures were present in the cellular periphery down to approximately 10 μm in depth, and extensive epidermal feet or filopodia-like processes were observed a few micrometers deep from the cellular surface. Cells were clustered or bundled from approximately 50 μm in depth to deeper levels. From 60 μm to 80 μm in depth, horizontal connections between these clusters were observed. The prospective eyespot and marginal focus areas were resistant to fluorescent dyes, likely because of their non-flat cone-like structures with a relatively thick cuticle. These in vivo images provide important information with which to understand processes of epithelial cell differentiation and color pattern determination in butterfly wings. PMID:26107809

  4. Live Cell Imaging of Butterfly Pupal and Larval Wings In Vivo.

    PubMed

    Ohno, Yoshikazu; Otaki, Joji M

    2015-01-01

    Butterfly wing color patterns are determined during the late larval and early pupal stages. Characterization of wing epithelial cells at these stages is thus critical to understand how wing structures, including color patterns, are determined. Previously, we successfully recorded real-time in vivo images of developing butterfly wings over time at the tissue level. In this study, we employed similar in vivo fluorescent imaging techniques to visualize developing wing epithelial cells in the late larval and early pupal stages 1 hour post-pupation. Both larval and pupal epithelial cells were rich in mitochondria and intracellular networks of endoplasmic reticulum, suggesting high metabolic activities, likely in preparation for cellular division, polyploidization, and differentiation. Larval epithelial cells in the wing imaginal disk were relatively large horizontally and tightly packed, whereas pupal epithelial cells were smaller and relatively loosely packed. Furthermore, larval cells were flat, whereas pupal cells were vertically elongated as deep as 130 μm. In pupal cells, many endosome-like or autophagosome-like structures were present in the cellular periphery down to approximately 10 μm in depth, and extensive epidermal feet or filopodia-like processes were observed a few micrometers deep from the cellular surface. Cells were clustered or bundled from approximately 50 μm in depth to deeper levels. From 60 μm to 80 μm in depth, horizontal connections between these clusters were observed. The prospective eyespot and marginal focus areas were resistant to fluorescent dyes, likely because of their non-flat cone-like structures with a relatively thick cuticle. These in vivo images provide important information with which to understand processes of epithelial cell differentiation and color pattern determination in butterfly wings.

  5. Does Skipping a Meal Matter to a Butterfly's Appearance? Effects of Larval Food Stress on Wing Morphology and Color in Monarch Butterflies

    PubMed Central

    Johnson, Haley; Solensky, Michelle J.; Satterfield, Dara A.; Davis, Andrew K.

    2014-01-01

    In animals with complex life cycles, all resources needed to form adult tissues are procured at the larval stage. For butterflies, the proper development of wings involves synthesizing tissue during metamorphosis based on the raw materials obtained by larvae. Similarly, manufacture of pigment for wing scales also requires resources acquired by larvae. We conducted an experiment to test the effects of food deprivation in the larval stage on multiple measures of adult wing morphology and coloration of monarch butterflies (Danaus plexippus), a species in which long-distance migration makes flight efficiency critical. In a captive setting, we restricted food (milkweed) from late-stage larvae for either 24 hrs or 48 hrs, then after metamorphosis we used image analysis methods to measure forewing surface area and elongation (length/width), which are both important for migration. We also measured the brightness of orange pigment and the intensity of black on the wing. There were correlations between several wing features, including an unexpected association between wing elongation and melanism, which will require further study to fully understand. The clearest effect of food restriction was a reduction in adult wing size in the high stress group (by approximately 2%). Patterns observed for other wing traits were ambiguous: monarchs in the low stress group (but not the high) had less elongated and paler orange pigmentation. There was no effect on wing melanism. Although some patterns obtained in this study were unclear, our results concerning wing size have direct bearing on the monarch migration. We show that if milkweed is limited for monarch larvae, their wings become stunted, which could ultimately result in lower migration success. PMID:24695643

  6. Does skipping a meal matter to a butterfly's appearance? Effects of larval food stress on wing morphology and color in monarch butterflies.

    PubMed

    Johnson, Haley; Solensky, Michelle J; Satterfield, Dara A; Davis, Andrew K

    2014-01-01

    In animals with complex life cycles, all resources needed to form adult tissues are procured at the larval stage. For butterflies, the proper development of wings involves synthesizing tissue during metamorphosis based on the raw materials obtained by larvae. Similarly, manufacture of pigment for wing scales also requires resources acquired by larvae. We conducted an experiment to test the effects of food deprivation in the larval stage on multiple measures of adult wing morphology and coloration of monarch butterflies (Danaus plexippus), a species in which long-distance migration makes flight efficiency critical. In a captive setting, we restricted food (milkweed) from late-stage larvae for either 24 hrs or 48 hrs, then after metamorphosis we used image analysis methods to measure forewing surface area and elongation (length/width), which are both important for migration. We also measured the brightness of orange pigment and the intensity of black on the wing. There were correlations between several wing features, including an unexpected association between wing elongation and melanism, which will require further study to fully understand. The clearest effect of food restriction was a reduction in adult wing size in the high stress group (by approximately 2%). Patterns observed for other wing traits were ambiguous: monarchs in the low stress group (but not the high) had less elongated and paler orange pigmentation. There was no effect on wing melanism. Although some patterns obtained in this study were unclear, our results concerning wing size have direct bearing on the monarch migration. We show that if milkweed is limited for monarch larvae, their wings become stunted, which could ultimately result in lower migration success.

  7. EB Ford revisited: assessing the long-term stability of wing-spot patterns and population genetic structure of the meadow brown butterfly on the Isles of Scilly

    PubMed Central

    Baxter, S W; Hoffman, J I; Tregenza, T; Wedell, N; Hosken, D J

    2017-01-01

    Understanding selection in the wild remains a major aim of evolutionary ecology and work by Ford and colleagues on the meadow brown butterfly Maniola jurtina did much to ignite this agenda. A great deal of their work was conducted during the 1950s on the Isles of Scilly. They documented island-specific wing-spot patterns that remained consistent over about a decade, but patterns on some islands changed after environmental perturbation. It was suggested that these wing-spot patterns reflected island-specific selection and that there was little migration between islands. However, genetic studies to test the underlying assumption of restricted migration are lacking and it is also unknown whether the originally described wing-spot patterns have persisted over time. We therefore collected female butterflies from five of Ford's original study locations, including three large islands (St Mary's, St Martin's and Tresco) and two small islands (Tean and St Helen's). Wing-spot patterns had not changed appreciably over time on three of the islands (two large and one small), but were significantly different on the other two. Furthermore, analysis of 176 amplified fragment length polymorphisms revealed significant genome-wide differentiation among the five islands. Our findings are consistent with Ford's conclusions that despite the close proximity of these islands, there is restricted gene flow among them. PMID:27804964

  8. In situ protocol for butterfly pupal wings using riboprobes.

    PubMed

    Ramos, Diane; Monteiro, Antonia

    2007-01-01

    Here we present, in video format, a protocol for in situ hybridizations in pupal wings of the butterfly Bicyclus anynana using riboprobes. In situ hybridizations, a mainstay of developmental biology, are useful to study the spatial and temporal patterns of gene expression in developing tissues at the level of transcription. If antibodies that target the protein products of gene transcription have not yet been developed, and/or there are multiple gene copies of a particular protein in the genome that cannot be differentiated using available antibodies, in situs can be used instead. While an in situ technique for larval wing discs has been available to the butterfly community for several years, the current protocol has been optimized for the larger and more fragile pupal wings.

  9. Positional dependence of scale size and shape in butterfly wings: wing-wide phenotypic coordination of color-pattern elements and background.

    PubMed

    Kusaba, Kiseki; Otaki, Joji M

    2009-02-01

    Butterfly wing color-patterns are a phenotypically coordinated array of scales whose color is determined as cellular interpretation outputs for morphogenic signals. Here we investigated distribution patterns of scale shape and size in relation to position and coloration on the hindwings of a nymphalid butterfly Junonia orithya. Most scales had a smooth edge but scales at and near the natural and ectopic eyespot foci and in the postbasal area were jagged. Scale size decreased regularly from the postbasal to distal areas, and eyespots occasionally had larger scales than the background. Reasonable correlations were obtained between the eyespot size and focal scale size in females. Histological and real-time individual observations of the color-pattern developmental sequence showed that the background brown and blue colors expanded from the postbasal to distal areas independently from the color-pattern elements such as eyespots. These data suggest that morphogenic signals for coloration directly or indirectly influence the scale shape and size and that the blue "background" is organized by a long-range signal from an unidentified organizing center in J. orithya.

  10. Color pattern analysis of nymphalid butterfly wings: revision of the nymphalid groundplan.

    PubMed

    Otaki, Joji M

    2012-09-01

    To better understand the developmental mechanisms of color pattern variation in butterfly wings, it is important to construct an accurate representation of pattern elements, known as the "nymphalid groundplan". However, some aspects of the current groundplan remain elusive. Here, I examined wing-wide elemental patterns of various nymphalid butterflies and confirmed that wing-wide color patterns are composed of the border, central, and basal symmetry systems. The central and basal symmetry systems can express circular patterns resembling eyespots, indicating that these systems have developmental mechanisms similar to those of the border symmetry system. The wing root band commonly occurs as a distinct symmetry system independent from the basal symmetry system. In addition, the marginal and submarginal bands are likely generated as a single system, referred to as the "marginal band system". Background spaces between two symmetry systems are sometimes light in coloration and can produce white bands, contributing significantly to color pattern diversity. When an element is enlarged with a pale central area, a visually similar (yet developmentally distinct) white band is produced. Based on the symmetric relationships of elements, I propose that both the central and border symmetry systems are comprised of "core elements" (the discal spot and the border ocelli, respectively) and a pair of "paracore elements" (the distal and proximal bands and the parafocal elements, respectively). Both core and paracore elements can be doubled, or outlined. Developmentally, this system configuration is consistent with the induction model, but not with the concentration gradient model for positional information.

  11. Comparative population genetics of a mimicry locus among hybridizing Heliconius butterfly species.

    PubMed

    Chamberlain, N L; Hill, R I; Baxter, S W; Jiggins, C D; Kronforst, M R

    2011-09-01

    The comimetic Heliconius butterfly species pair, H. erato and H. melpomene, appear to use a conserved Mendelian switch locus to generate their matching red wing patterns. Here we investigate whether H. cydno and H. pachinus, species closely related to H. melpomene, use this same switch locus to generate their highly divergent red and brown color pattern elements. Using an F2 intercross between H. cydno and H. pachinus, we first map the genomic positions of two novel red/brown wing pattern elements; the G locus, which controls the presence of red vs brown at the base of the ventral wings, and the Br locus, which controls the presence vs absence of a brown oval pattern on the ventral hind wing. The results reveal that the G locus is tightly linked to markers in the genomic interval that controls red wing pattern elements of H. erato and H. melpomene. Br is on the same linkage group but approximately 26 cM away. Next, we analyze fine-scale patterns of genetic differentiation and linkage disequilibrium throughout the G locus candidate interval in H. cydno, H. pachinus and H. melpomene, and find evidence for elevated differentiation between H. cydno and H. pachinus, but no localized signature of association. Overall, these results indicate that the G locus maps to the same interval as the locus controlling red patterning in H. melpomene and H. erato. This, in turn, suggests that the genes controlling red pattern elements may be homologous across Heliconius, supporting the hypothesis that Heliconius butterflies use a limited suite of conserved genetic switch loci to generate both convergent and divergent wing patterns.

  12. Local pharmacological effects of tungstate on the color-pattern determination of butterfly wings: a possible relationship between the eyespot and parafocal element.

    PubMed

    Dhungel, Bidur; Otaki, Joji M

    2009-11-01

    Butterfly wing color patterns can be changed by the application of a temperature shock or pharmacological agents such as tungstate, producing a distinctive type of elemental modification called the TS (temperature shock) type. Heterochronic uncoupling between the signaling and reception steps during the color-pattern determination process has been proposed as a mechanism for TS-type changes. As an extension of this hypothesis, both the parafocal element (PFE) and the eyespot in the same wing compartment are considered to be determined by morphogenic signal(s) emitted from the same eyespot focus. However, these models need to be examined with additional experimental data. Furthermore, there is controversy as to whether the action of tungstate on wing color patterns is direct or Indirect. Using a species of nymphalid butterfly (Junonia orithya), we have devised a simple method for the local application of pharmacological agents directly on developing wings of pupae. Local tungstate application resulted in reduced eyespots and circular dislocated PFEs in the eyespot-less compartments only on the treated wing, demonstrating that tungstate directly induces color-pattern changes on wings. We further examined the eyespot-PFE relationship in normal and cold-shocked Individuals, showing that an eyespot can be superimposed on a PFE and vice versa, probably depending on the timing of their fate determination. Taken together, we propose a two-morphogen model for the normal color-pattern determination, in which the morphogenic signals for the eyespot and PFE are different from each other despite their Identical origin. This two-morphogen model is compatible with the heterochronic uncoupling model for TS-type changes.

  13. Color pattern evolution in Vanessa butterflies (Nymphalidae: Nymphalini): non-eyespot characters.

    PubMed

    Abbasi, Roohollah; Marcus, Jeffrey M

    2015-01-01

    A phylogenetic approach was used to study color pattern evolution in Vanessa butterflies. Twenty-four color pattern elements from the Nymphalid ground plan were identified on the dorsal and ventral surfaces of the fore- and hind wings. Eyespot characters were excluded and will be examined elsewhere. The evolution of each character was traced over a Bayesian phylogeny of Vanessa reconstructed from 7750 DNA base pairs from 10 genes. Generally, the correspondence between character states on the same surface of the two wings is stronger on the ventral side compared to the dorsal side. The evolution of character states on both sides of a wing correspond with each other in most extant species, but the correspondence between dorsal and ventral character states is much stronger in the forewing than in the hindwing. The dorsal hindwing of many species of Vanessa is covered with an extended Basal Symmetry System and the Discalis I pattern element is highly variable between species, making this wing surface dissimilar to the other wing surfaces. The Basal Symmetry System and Discalis I may contribute to behavioral thermoregulation in Vanessa. Overall, interspecific directional character state evolution of non-eyespot color patterns is relatively rare in Vanessa, with a majority of color pattern elements showing non-variable, non-directional, or ambiguous character state evolution. The ease with which the development of color patterns can be modified, including character state reversals, has likely made important contributions to the production of color pattern diversity in Vanessa and other butterfly groups. © 2014 Wiley Periodicals, Inc.

  14. Pterin pigment granules are responsible for both broadband light scattering and wavelength selective absorption in the wing scales of pierid butterflies

    PubMed Central

    Morehouse, Nathan I; Vukusic, Peter; Rutowski, Ron

    2006-01-01

    A small but growing literature indicates that many animal colours are produced by combinations of structural and pigmentary mechanisms. We investigated one such complex colour phenotype: the highly chromatic wing colours of pierid butterflies including oranges, yellows and patterns which appear white to the human eye, but strongly absorb the ultraviolet (UV) wavelengths visible to butterflies. Pierids produce these bright colours using wing scales that contain collections of minute granules. However, to date, no work has directly characterized the molecular composition or optical properties of these granules. We present results that indicate these granules contain pterin pigments. We also find that pterin granules increase light reflection from single wing scales, such that wing scales containing denser granule arrays reflect more light than those with less dense granule collections. As male wing scales contain more pterin granules than those of females, the sexual dichromatism found in many pierid species can be explained by differences in wing scale pterin deposition. Additionally, the colour pattern elements produced by these pterins are known to be important during mating interactions in a number of pierid species. Therefore, we discuss the potential relevance of our results within the framework of sexual selection and colour signal evolution. PMID:17164199

  15. Wing shape variation associated with mimicry in butterflies.

    PubMed

    Jones, Robert T; Le Poul, Yann; Whibley, Annabel C; Mérot, Claire; ffrench-Constant, Richard H; Joron, Mathieu

    2013-08-01

    Mimetic resemblance in unpalatable butterflies has been studied by evolutionary biologists for over a century, but has largely focused on the convergence in wing color patterns. In Heliconius numata, discrete color-pattern morphs closely resemble comimics in the distantly related genus Melinaea. We examine the possibility that the shape of the butterfly wing also shows adaptive convergence. First, simple measures of forewing dimensions were taken of individuals in a cross between H. numata morphs, and showed quantitative differences between two of the segregating morphs, f. elegans and f. silvana. Second, landmark-based geometric morphometric and elliptical Fourier outline analyses were used to more fully characterize these shape differences. Extension of these techniques to specimens from natural populations suggested that, although many of the coexisting morphs could not be discriminated by shape, the differences we identified between f. elegans and f. silvana hold in the wild. Interestingly, despite extensive overlap, the shape variation between these two morphs is paralleled in their respective Melinaea comimics. Our study therefore suggests that wing-shape variation is associated with mimetic resemblance, and raises the intriguing possibility that the supergene responsible for controlling the major switch in color pattern between morphs also contributes to wing shape differences in H. numata. © 2013 The Author(s). Evolution © 2013 The Society for the Study of Evolution.

  16. Underwater flight by the planktonic sea butterfly.

    PubMed

    Murphy, David W; Adhikari, Deepak; Webster, Donald R; Yen, Jeannette

    2016-02-01

    In a remarkable example of convergent evolution, we show that the zooplanktonic sea butterfly Limacina helicina 'flies' underwater in the same way that very small insects fly in the air. Both sea butterflies and flying insects stroke their wings in a characteristic figure-of-eight pattern to produce lift, and both generate extra lift by peeling their wings apart at the beginning of the power stroke (the well-known Weis-Fogh 'clap-and-fling' mechanism). It is highly surprising to find a zooplankter 'mimicking' insect flight as almost all zooplankton swim in this intermediate Reynolds number range (Re=10-100) by using their appendages as paddles rather than wings. The sea butterfly is also unique in that it accomplishes its insect-like figure-of-eight wing stroke by extreme rotation of its body (what we call 'hyper-pitching'), a paradigm that has implications for micro aerial vehicle (MAV) design. No other animal, to our knowledge, pitches to this extent under normal locomotion. © 2016. Published by The Company of Biologists Ltd.

  17. Keeping the band together: evidence for false boundary disruptive coloration in a butterfly.

    PubMed

    Seymoure, B M; Aiello, A

    2015-09-01

    There is a recent surge of evidence supporting disruptive coloration, in which patterns break up the animal's outline through false edges or boundaries, increasing survival in animals by reducing predator detection and/or preventing recognition. Although research has demonstrated that false edges are successful for reducing predation of prey, research into the role of internal false boundaries (i.e. stripes and bands) in reducing predation remains warranted. Many animals have stripes and bands that may function disruptively. Here, we test the possible disruptive function of wing band patterning in a butterfly, Anartia fatima, using artificial paper and plasticine models in Panama. We manipulated the band so that one model type had the band shifted to the wing margin (nondisruptive treatment) and another model had a discontinuous band located on the wing margin (discontinuous edge treatment). We kept the natural wing pattern to represent the false boundary treatment. Across all treatment groups, we standardized the area of colour and used avian visual models to confirm a match between manipulated and natural wing colours. False boundary models had higher survival than either the discontinuous edge model or the nondisruptive model. There was no survival difference between the discontinuous edge model and the nondisruptive model. Our results demonstrate the importance of wing bands in reducing predation on butterflies and show that markings set in from the wing margin can reduce predation more effectively than marginal bands and discontinuous marginal patterns. This study demonstrates an adaptive benefit of having stripes and bands. © 2015 European Society For Evolutionary Biology.

  18. Convergent evolution in the genetic basis of Müllerian mimicry in heliconius butterflies.

    PubMed

    Baxter, Simon W; Papa, Riccardo; Chamberlain, Nicola; Humphray, Sean J; Joron, Mathieu; Morrison, Clay; ffrench-Constant, Richard H; McMillan, W Owen; Jiggins, Chris D

    2008-11-01

    The neotropical butterflies Heliconius melpomene and H. erato are Müllerian mimics that display the same warningly colored wing patterns in local populations, yet pattern diversity between geographic regions. Linkage mapping has previously shown convergent red wing phenotypes in these species are controlled by loci on homologous chromosomes. Here, AFLP bulk segregant analysis using H. melpomene crosses identified genetic markers tightly linked to two red wing-patterning loci. These markers were used to screen a H. melpomene BAC library and a tile path was assembled spanning one locus completely and part of the second. Concurrently, a similar strategy was used to identify a BAC clone tightly linked to the locus controlling the mimetic red wing phenotypes in H. erato. A methionine rich storage protein (MRSP) gene was identified within this BAC clone, and comparative genetic mapping shows red wing color loci are in homologous regions of the genome of H. erato and H. melpomene. Subtle differences in these convergent phenotypes imply they evolved independently using somewhat different developmental routes, but are nonetheless regulated by the same switch locus. Genetic mapping of MRSP in a third related species, the "tiger" patterned H. numata, has no association with wing patterning and shows no evidence for genomic translocation of wing-patterning loci.

  19. Convergent, modular expression of ebony and tan in the mimetic wing patterns of Heliconius butterflies.

    PubMed

    Ferguson, Laura C; Maroja, Luana; Jiggins, Chris D

    2011-12-01

    The evolution of pigmentation in vertebrates and flies has involved repeated divergence at a small number of genes related to melanin synthesis. Here, we study insect melanin synthesis genes in Heliconius butterflies, a group characterised by its diversity of wing patterns consisting of black (melanin), and yellow and red (ommochrome) pigmented scales. Consistent with their respective biochemical roles in Drosophila melanogaster, ebony is upregulated in non-melanic wing regions destined to be pigmented red whilst tan is upregulated in melanic regions. Wing regions destined to be pigmented yellow, however, are downregulated for both genes. This pattern is conserved across multiple divergent and convergent phenotypes within the Heliconii, suggesting a conserved mechanism for the development of black, red and yellow pattern elements across the genus. Linkage mapping of five melanin biosynthesis genes showed that, in contrast to other organisms, these genes do not control pattern polymorphism. Thus, the pigmentation genes themselves are not the locus of evolutionary change but lie downstream of a wing pattern regulatory factor. The results suggest a modular system in which particular combinations of genes are switched on whenever red, yellow or black pattern elements are favoured by natural selection for diverse and mimetic wing patterns. © Springer-Verlag 2011

  20. The marginal band system in nymphalid butterfly wings.

    PubMed

    Taira, Wataru; Kinjo, Seira; Otaki, Joji M

    2015-01-01

    Butterfly wing color patterns are highly complex and diverse, but they are believed to be derived from the nymphalid groundplan, which is composed of several color pattern systems. Among these pattern systems, the marginal band system, including marginal and submarginal bands, has rarely been studied. Here, we examined the color pattern diversity of the marginal band system among nymphalid butterflies. Marginal and submarginal bands are usually expressed as a pair of linear bands aligned with the wing margin. However, a submarginal band can be expressed as a broken band, an elongated oval, or a single dot. The marginal focus, usually a white dot at the middle of a wing compartment along the wing edge, corresponds to the pupal edge spot, one of the pupal cuticle spots that signify the locations of color pattern organizing centers. A marginal band can be expressed as a semicircle, an elongated oval, or a pair of eyespot-like structures, which suggest the organizing activity of the marginal focus. Physical damage at the pupal edge spot leads to distal dislocation of the submarginal band in Junonia almana and in Vanessa indica, suggesting that the marginal focus functions as an organizing center for the marginal band system. Taken together, we conclude that the marginal band system is developmentally equivalent to other symmetry systems. Additionally, the marginal band is likely a core element and the submarginal band a paracore element of the marginal band system, and both bands are primarily specified by the marginal focus organizing center.

  1. Butterflies regulate wing temperatures using radiative cooling

    NASA Astrophysics Data System (ADS)

    Tsai, Cheng-Chia; Shi, Norman Nan; Ren, Crystal; Pelaez, Julianne; Bernard, Gary D.; Yu, Nanfang; Pierce, Naomi

    2017-09-01

    Butterfly wings are live organs embedded with multiple sensory neurons and, in some species, with pheromoneproducing cells. The proper function of butterfly wings demands a suitable temperature range, but the wings can overheat quickly in the sun due to their small thermal capacity. We developed an infrared technique to map butterfly wing temperatures and discovered that despite the wings' diverse visible colors, regions of wings that contain live cells are the coolest, resulting from the thickness of the wings and scale nanostructures. We also demonstrated that butterflies use behavioral traits to prevent overheating of their wings.

  2. Gene expression underlying adaptive variation in Heliconius wing patterns: non-modular regulation of overlapping cinnabar and vermilion prepatterns.

    PubMed

    Reed, Robert D; McMillan, W Owen; Nagy, Lisa M

    2008-01-07

    Geographical variation in the mimetic wing patterns of the butterfly Heliconius erato is a textbook example of adaptive polymorphism; however, little is known about how this variation is controlled developmentally. Using microarrays and qPCR, we identified and compared expression of candidate genes potentially involved with a red/yellow forewing band polymorphism in H. erato. We found that transcripts encoding the pigment synthesis enzymes cinnabar and vermilion showed pattern- and polymorphism-related expression patterns, respectively. cinnabar expression was associated with the forewing band regardless of pigment colour, providing the first gene expression pattern known to be correlated with a major Heliconius colour pattern. In contrast, vermilion expression changed spatially over time in red-banded butterflies, but was not expressed at detectable levels in yellow-banded butterflies, suggesting that regulation of this gene may be involved with the red/yellow polymorphism. Furthermore, we found that the yellow pigment, 3-hydroxykynurenine, is incorporated into wing scales from the haemolymph rather than being synthesized in situ. We propose that some aspects of Heliconius colour patterns are determined by spatio-temporal overlap of pigment gene transcription prepatterns and speculate that evolutionary changes in vermilion regulation may in part underlie an adaptive colour pattern polymorphism.

  3. Acronictinae (Lepidoptera: Macroheterocera: Noctuidae) demonstrate the variable role of wing venation in the evolution of the nymphalid groundplan

    USDA-ARS?s Scientific Manuscript database

    The nymphalid groundplan (NGP) is an idealized system used to classify and interpret wing pattern elements of butterflies. Nearly a century ago, the principles of the NGP were applied to the wing patterns of higher moths (Macroheterocera). Recent advances in phylogeny and in the comparative morpholo...

  4. Prey from the eyes of predators: Color discriminability of aposematic and mimetic butterflies from an avian visual perspective.

    PubMed

    Su, Shiyu; Lim, Matthew; Kunte, Krushnamegh

    2015-11-01

    Predation exerts strong selection on mimetic butterfly wing color patterns, which also serve other functions such as sexual selection. Therefore, specific selection pressures may affect the sexes and signal components differentially. We tested three predictions about the evolution of mimetic resemblance by comparing wing coloration of aposematic butterflies and their Batesian mimics: (a) females gain greater mimetic advantage than males and therefore are better mimics, (b) due to intersexual genetic correlations, sexually monomorphic mimics are better mimics than female-limited mimics, and (c) mimetic resemblance is better on the dorsal wing surface that is visible to predators in flight. Using a physiological model of avian color vision, we quantified mimetic resemblance from predators' perspective, which showed that female butterflies were better mimics than males. Mimetic resemblance in female-limited mimics was comparable to that in sexually monomorphic mimics, suggesting that intersexual genetic correlations did not constrain adaptive response to selection for female-limited mimicry. Mimetic resemblance on the ventral wing surface was better than that on the dorsal wing surface, implying stronger natural and sexual selection on ventral and dorsal surfaces, respectively. These results suggest that mimetic resemblance in butterfly mimicry rings has evolved under various selective pressures acting in a sex- and wing surface-specific manner. © 2015 The Author(s). Evolution © 2015 The Society for the Study of Evolution.

  5. Why Small Is Beautiful: Wing Colour Is Free from Thermoregulatory Constraint in the Small Lycaenid Butterfly, Polyommatus icarus

    PubMed Central

    De Keyser, Rien; Breuker, Casper J.; Hails, Rosemary S.; Dennis, Roger L. H.; Shreeve, Tim G.

    2015-01-01

    We examined the roles of wing melanisation, weight, and basking posture in thermoregulation in Polyommatus Icarus, a phenotypically variable and protandrous member of the diverse Polyommatinae (Lycaenidae). Under controlled experimental conditions, approximating to marginal environmental conditions for activity in the field (= infrequent flight, long duration basking periods), warming rates are maximised with fully open wings and maximum body temperatures are dependent on weight. Variation in wing melanisation within and between sexes has no effect on warming rates; males and females which differ in melanisation had similar warming rates. Posture also affected cooling rates, consistent with cooling being dependent on convective heat loss. We hypothesise that for this small sized butterfly, melanisation has little or no effect on thermoregulation. This may be a factor contributing to the diversity of wing colours in the Polyommatinae. Because of the importance of size for thermoregulation in this small butterfly, requirements for attaining a suitable size to confer thermal stability in adults may also be a factor influencing larval feeding rates, development time and patterns of voltinism. Our findings indicate that commonly accepted views of the importance of melanisation, posture and size to thermoregulation, developed using medium and large sized butterflies, are not necessarily applicable to small sized butterflies. PMID:25923738

  6. Why Small Is Beautiful: Wing Colour Is Free from Thermoregulatory Constraint in the Small Lycaenid Butterfly, Polyommatus icarus.

    PubMed

    De Keyser, Rien; Breuker, Casper J; Hails, Rosemary S; Dennis, Roger L H; Shreeve, Tim G

    2015-01-01

    We examined the roles of wing melanisation, weight, and basking posture in thermoregulation in Polyommatus Icarus, a phenotypically variable and protandrous member of the diverse Polyommatinae (Lycaenidae). Under controlled experimental conditions, approximating to marginal environmental conditions for activity in the field (= infrequent flight, long duration basking periods), warming rates are maximised with fully open wings and maximum body temperatures are dependent on weight. Variation in wing melanisation within and between sexes has no effect on warming rates; males and females which differ in melanisation had similar warming rates. Posture also affected cooling rates, consistent with cooling being dependent on convective heat loss. We hypothesise that for this small sized butterfly, melanisation has little or no effect on thermoregulation. This may be a factor contributing to the diversity of wing colours in the Polyommatinae. Because of the importance of size for thermoregulation in this small butterfly, requirements for attaining a suitable size to confer thermal stability in adults may also be a factor influencing larval feeding rates, development time and patterns of voltinism. Our findings indicate that commonly accepted views of the importance of melanisation, posture and size to thermoregulation, developed using medium and large sized butterflies, are not necessarily applicable to small sized butterflies.

  7. A wing expressed sequence tag resource for Bicyclus anynana butterflies, an evo-devo model

    PubMed Central

    Beldade, Patrícia; Rudd, Stephen; Gruber, Jonathan D; Long, Anthony D

    2006-01-01

    Background Butterfly wing color patterns are a key model for integrating evolutionary developmental biology and the study of adaptive morphological evolution. Yet, despite the biological, economical and educational value of butterflies they are still relatively under-represented in terms of available genomic resources. Here, we describe an Expression Sequence Tag (EST) project for Bicyclus anynana that has identified the largest available collection to date of expressed genes for any butterfly. Results By targeting cDNAs from developing wings at the stages when pattern is specified, we biased gene discovery towards genes potentially involved in pattern formation. Assembly of 9,903 ESTs from a subtracted library allowed us to identify 4,251 genes of which 2,461 were annotated based on BLAST analyses against relevant gene collections. Gene prediction software identified 2,202 peptides, of which 215 longer than 100 amino acids had no homology to any known proteins and, thus, potentially represent novel or highly diverged butterfly genes. We combined gene and Single Nucleotide Polymorphism (SNP) identification by constructing cDNA libraries from pools of outbred individuals, and by sequencing clones from the 3' end to maximize alignment depth. Alignments of multi-member contigs allowed us to identify over 14,000 putative SNPs, with 316 genes having at least one high confidence double-hit SNP. We furthermore identified 320 microsatellites in transcribed genes that can potentially be used as genetic markers. Conclusion Our project was designed to combine gene and sequence polymorphism discovery and has generated the largest gene collection available for any butterfly and many potential markers in expressed genes. These resources will be invaluable for exploring the potential of B. anynana in particular, and butterflies in general, as models in ecological, evolutionary, and developmental genetics. PMID:16737530

  8. Butterfly Wings Are Three-Dimensional: Pupal Cuticle Focal Spots and Their Associated Structures in Junonia Butterflies.

    PubMed

    Taira, Wataru; Otaki, Joji M

    2016-01-01

    Butterfly wing color patterns often contain eyespots, which are developmentally determined at the late larval and early pupal stages by organizing activities of focal cells that can later form eyespot foci. In the pupal stage, the focal position of a future eyespot is often marked by a focal spot, one of the pupal cuticle spots, on the pupal surface. Here, we examined the possible relationships of the pupal focal spots with the underneath pupal wing tissues and with the adult wing eyespots using Junonia butterflies. Large pupal focal spots were found in two species with large adult eyespots, J. orithya and J. almana, whereas only small pupal focal spots were found in a species with small adult eyespots, J. hedonia. The size of five pupal focal spots on a single wing was correlated with the size of the corresponding adult eyespots in J. orithya. A pupal focal spot was a three-dimensional bulge of cuticle surface, and the underside of the major pupal focal spot exhibited a hollowed cuticle in a pupal case. Cross sections of a pupal wing revealed that the cuticle layer shows a curvature at a focal spot, and a positional correlation was observed between the cuticle layer thickness and its corresponding cell layer thickness. Adult major eyespots of J. orithya and J. almana exhibited surface elevations and depressions that approximately correspond to the coloration within an eyespot. Our results suggest that a pupal focal spot is produced by the organizing activity of focal cells underneath the focal spot. Probably because the focal cell layer immediately underneath a focal spot is thicker than that of its surrounding areas, eyespots of adult butterfly wings are three-dimensionally constructed. The color-height relationship in adult eyespots might have an implication in the developmental signaling for determining the eyespot color patterns.

  9. Butterfly Wings Are Three-Dimensional: Pupal Cuticle Focal Spots and Their Associated Structures in Junonia Butterflies

    PubMed Central

    Taira, Wataru; Otaki, Joji M.

    2016-01-01

    Butterfly wing color patterns often contain eyespots, which are developmentally determined at the late larval and early pupal stages by organizing activities of focal cells that can later form eyespot foci. In the pupal stage, the focal position of a future eyespot is often marked by a focal spot, one of the pupal cuticle spots, on the pupal surface. Here, we examined the possible relationships of the pupal focal spots with the underneath pupal wing tissues and with the adult wing eyespots using Junonia butterflies. Large pupal focal spots were found in two species with large adult eyespots, J. orithya and J. almana, whereas only small pupal focal spots were found in a species with small adult eyespots, J. hedonia. The size of five pupal focal spots on a single wing was correlated with the size of the corresponding adult eyespots in J. orithya. A pupal focal spot was a three-dimensional bulge of cuticle surface, and the underside of the major pupal focal spot exhibited a hollowed cuticle in a pupal case. Cross sections of a pupal wing revealed that the cuticle layer shows a curvature at a focal spot, and a positional correlation was observed between the cuticle layer thickness and its corresponding cell layer thickness. Adult major eyespots of J. orithya and J. almana exhibited surface elevations and depressions that approximately correspond to the coloration within an eyespot. Our results suggest that a pupal focal spot is produced by the organizing activity of focal cells underneath the focal spot. Probably because the focal cell layer immediately underneath a focal spot is thicker than that of its surrounding areas, eyespots of adult butterfly wings are three-dimensionally constructed. The color-height relationship in adult eyespots might have an implication in the developmental signaling for determining the eyespot color patterns. PMID:26731532

  10. Real-Time In Vivo Imaging of Butterfly Wing Development: Revealing the Cellular Dynamics of the Pupal Wing Tissue

    PubMed Central

    Iwata, Masaki; Ohno, Yoshikazu; Otaki, Joji M.

    2014-01-01

    Butterfly wings are covered with regularly arranged single-colored scales that are formed at the pupal stage. Understanding pupal wing development is therefore crucial to understand wing color pattern formation. Here, we successfully employed real-time in vivo imaging techniques to observe pupal hindwing development over time in the blue pansy butterfly, Junonia orithya. A transparent sheet of epithelial cells that were not yet regularly arranged was observed immediately after pupation. Bright-field imaging and autofluorescent imaging revealed free-moving hemocytes and tracheal branches of a crinoid-like structure underneath the epithelium. The wing tissue gradually became gray-white, epithelial cells were arranged regularly, and hemocytes disappeared, except in the bordering lacuna, after which scales grew. The dynamics of the epithelial cells and scale growth were also confirmed by fluorescent imaging. Fluorescent in vivo staining further revealed that these cells harbored many mitochondria at the surface of the epithelium. Organizing centers for the border symmetry system were apparent immediately after pupation, exhibiting a relatively dark optical character following treatment with fluorescent dyes, as well as in autofluorescent images. The wing tissue exhibited slow and low-frequency contraction pulses with a cycle of approximately 10 to 20 minutes, mainly occurring at 2 to 3 days postpupation. The pulses gradually became slower and weaker and eventually stopped. The wing tissue area became larger after contraction, which also coincided with an increase in the autofluorescence intensity that might have been caused by scale growth. Examination of the pattern of color development revealed that the black pigment was first deposited in patches in the central areas of an eyespot black ring and a parafocal element. These results of live in vivo imaging that covered wide wing area for a long time can serve as a foundation for studying the cellular dynamics of living wing tissues in butterflies. PMID:24586829

  11. Real-time in vivo imaging of butterfly wing development: revealing the cellular dynamics of the pupal wing tissue.

    PubMed

    Iwata, Masaki; Ohno, Yoshikazu; Otaki, Joji M

    2014-01-01

    Butterfly wings are covered with regularly arranged single-colored scales that are formed at the pupal stage. Understanding pupal wing development is therefore crucial to understand wing color pattern formation. Here, we successfully employed real-time in vivo imaging techniques to observe pupal hindwing development over time in the blue pansy butterfly, Junonia orithya. A transparent sheet of epithelial cells that were not yet regularly arranged was observed immediately after pupation. Bright-field imaging and autofluorescent imaging revealed free-moving hemocytes and tracheal branches of a crinoid-like structure underneath the epithelium. The wing tissue gradually became gray-white, epithelial cells were arranged regularly, and hemocytes disappeared, except in the bordering lacuna, after which scales grew. The dynamics of the epithelial cells and scale growth were also confirmed by fluorescent imaging. Fluorescent in vivo staining further revealed that these cells harbored many mitochondria at the surface of the epithelium. Organizing centers for the border symmetry system were apparent immediately after pupation, exhibiting a relatively dark optical character following treatment with fluorescent dyes, as well as in autofluorescent images. The wing tissue exhibited slow and low-frequency contraction pulses with a cycle of approximately 10 to 20 minutes, mainly occurring at 2 to 3 days postpupation. The pulses gradually became slower and weaker and eventually stopped. The wing tissue area became larger after contraction, which also coincided with an increase in the autofluorescence intensity that might have been caused by scale growth. Examination of the pattern of color development revealed that the black pigment was first deposited in patches in the central areas of an eyespot black ring and a parafocal element. These results of live in vivo imaging that covered wide wing area for a long time can serve as a foundation for studying the cellular dynamics of living wing tissues in butterflies.

  12. The generation and diversification of butterfly eyespot color patterns.

    PubMed

    Brunetti, C R; Selegue, J E; Monteiro, A; French, V; Brakefield, P M; Carroll, S B

    2001-10-16

    A fundamental challenge of evolutionary and developmental biology is understanding how new characters arise and change. The recently derived eyespots on butterfly wings vary extensively in number and pattern between species and play important roles in predator avoidance. Eyespots form through the activity of inductive organizers (foci) at the center of developing eyespot fields. Foci are the proposed source of a morphogen, the levels of which determine the color of surrounding wing scale cells. However, it is unknown how reception of the focal signal translates into rings of different-colored scales, nor how different color schemes arise in different species. We have identified several transcription factors, including butterfly homologs of the Drosophila Engrailed/Invected and Spalt proteins, that are deployed in concentric territories corresponding to the future rings of pigmented scales that compose the adult eyespot. We have isolated a new Bicyclus anynana wing pattern mutant, Goldeneye, in which the scales of one inner color ring become the color of a different ring. These changes correlate with shifts in transcription factor expression, suggesting that Goldeneye affects an early regulatory step in eyespot color patterning. In different butterfly species, the same transcription factors are expressed in eyespot fields, but in different relative spatial domains that correlate with divergent eyespot color schemes. Our results suggest that signaling from the focus induces nested rings of regulatory gene expression that subsequently control the final color pattern. Furthermore, the remarkably plastic regulatory interactions downstream of focal signaling have facilitated the evolution of eyespot diversity.

  13. Spatial reflection patterns of iridescent wings of male pierid butterflies: curved scales reflect at a wider angle than flat scales.

    PubMed

    Pirih, Primož; Wilts, Bodo D; Stavenga, Doekele G

    2011-10-01

    The males of many pierid butterflies have iridescent wings, which presumably function in intraspecific communication. The iridescence is due to nanostructured ridges of the cover scales. We have studied the iridescence in the males of a few members of Coliadinae, Gonepteryx aspasia, G. cleopatra, G. rhamni, and Colias croceus, and in two members of the Colotis group, Hebomoia glaucippe and Colotis regina. Imaging scatterometry demonstrated that the pigmentary colouration is diffuse whereas the structural colouration creates a directional, line-shaped far-field radiation pattern. Angle-dependent reflectance measurements demonstrated that the directional iridescence distinctly varies among closely related species. The species-dependent scale curvature determines the spatial properties of the wing iridescence. Narrow beam illumination of flat scales results in a narrow far-field iridescence pattern, but curved scales produce broadened patterns. The restricted spatial visibility of iridescence presumably plays a role in intraspecific signalling.

  14. Forward flight of swallowtail butterfly with simple flapping motion.

    PubMed

    Tanaka, Hiroto; Shimoyama, Isao

    2010-06-01

    Unlike other flying insects, the wing motion of swallowtail butterflies is basically limited to flapping because their fore wings partly overlap their hind wings, structurally restricting the feathering needed for active control of aerodynamic force. Hence, it can be hypothesized that the flight of swallowtail butterflies is realized with simple flapping, requiring little feedback control of the feathering angle. To verify this hypothesis, we fabricated an artificial butterfly mimicking the wing motion and wing shape of a swallowtail butterfly and analyzed its flights using images taken with a high-speed video camera. The results demonstrated that stable forward flight could be realized without active feathering or feedback control of the wing motion. During the flights, the artificial butterfly's body moved up and down passively in synchronization with the flapping, and the artificial butterfly followed an undulating flight trajectory like an actual swallowtail butterfly. Without feedback control of the wing motion, the body movement is directly affected by change of aerodynamic force due to the wing deformation; the degree of deformation was determined by the wing venation. Unlike a veinless wing, a mimic wing with veins generated a much higher lift coefficient during the flapping flight than in a steady flow due to the large body motion.

  15. Replication of butterfly wing and natural lotus leaf structures by nanoimprint on silica sol-gel films.

    PubMed

    Saison, Tamar; Peroz, Christophe; Chauveau, Vanessa; Berthier, Serge; Sondergard, Elin; Arribart, Hervé

    2008-12-01

    An original and low cost method for the fabrication of patterned surfaces bioinspired from butterfly wings and lotus leaves is presented. Silica-based sol-gel films are thermally imprinted from elastomeric molds to produce stable structures with superhydrophobicity values as high as 160 degrees water contact angle. The biomimetic surfaces are demonstrated to be tuned from superhydrophobic to superhydrophilic by annealing between 200 degrees C and 500 degrees C.

  16. The Lycaenid Central Symmetry System: Color Pattern Analysis of the Pale Grass Blue Butterfly Zizeeria maha.

    PubMed

    Iwata, Masaki; Taira, Wataru; Hiyama, Atsuki; Otaki, Joji M

    2015-06-01

    The nymphalid groundplan has been proposed to explain diverse butterfly wing color patterns. In this model, each symmetry system is composed of a core element and a pair of paracore elements. The development of this elemental configuration has been explained by the induction model for positional information. However, the diversity of color patterns in other butterfly families in relation to the nymphalid groundplan has not been thoroughly examined. Here, we examined aberrant color pattern phenotypes of a lycaenid butterfly, Zizeeria maha, from mutagenesis and plasticity studies as well as from field surveys. In several mutants, the third and fourth spot arrays were coordinately positioned much closer to the discal spot in comparison to the normal phenotype. In temperature-shock types, the third and fourth array spots were elongated inwardly or outwardly from their normal positions. In field-caught spontaneous mutants, small black spots were located adjacent to normal black spots. Analysis of these aberrant phenotypes indicated that the spots belonging to the third and fourth arrays are synchronously changeable in position and shape around the discal spot. Thus, these arrays constitute paracore elements of the central symmetry system of the lycaenid butterflies, and the discal spot comprises the core element. These aberrant phenotypes can be explained by the black-inducing signals that propagate from the prospective discal spot, as predicted by the induction model. These results suggest the existence of long-range developmental signals that cover a large area of a wing not only in nymphalid butterflies, but also in lycaenid butterflies.

  17. Morphological outcomes of gynandromorphism in Lycaeides butterflies (Lepidoptera: Lycaenidae).

    PubMed

    Jahner, Joshua P; Lucas, Lauren K; Wilson, Joseph S; Forister, Matthew L

    2015-01-01

    The genitalia of male insects have been widely used in taxonomic identification and systematics and are potentially involved in maintaining reproductive isolation between species. Although sexual selection has been invoked to explain patterns of morphological variation in genitalia among populations and species, developmental plasticity in genitalia likely contributes to observed variation but has been rarely examined, particularly in wild populations. Bilateral gynandromorphs are individuals that are genetically male on one side of the midline and genetically female on the other, while mosaic gynandromorphs have only a portion of their body developing as the opposite sex. Gynandromorphs might offer unique insights into developmental plasticity because individuals experience abnormal cellular interactions at the genitalic midline. In this study, we compare the genitalia and wing patterns of gynandromorphic Anna and Melissa blue butterflies, Lycaeides anna (Edwards) (formerly L. idas anna) and L. melissa (Edwards) (Lepidoptera: Lycaenidae), to the morphology of normal individuals from the same populations. Gynandromorph wing markings all fell within the range of variation of normal butterflies; however, a number of genitalic measurements were outliers when compared with normal individuals. From these results, we conclude that the gynandromorphs' genitalia, but not wing patterns, can be abnormal when compared with normal individuals and that the gynandromorphic genitalia do not deviate developmentally in a consistent pattern across individuals. Finally, genetic mechanisms are considered for the development of gynandromorphism in Lycaeides butterflies. © The Author 2015. Published by Oxford University Press on behalf of the Entomological Society of America.

  18. Subtractive Structural Modification of Morpho Butterfly Wings.

    PubMed

    Shen, Qingchen; He, Jiaqing; Ni, Mengtian; Song, Chengyi; Zhou, Lingye; Hu, Hang; Zhang, Ruoxi; Luo, Zhen; Wang, Ge; Tao, Peng; Deng, Tao; Shang, Wen

    2015-11-11

    Different from studies of butterfly wings through additive modification, this work for the first time studies the property change of butterfly wings through subtractive modification using oxygen plasma etching. The controlled modification of butterfly wings through such subtractive process results in gradual change of the optical properties, and helps the further understanding of structural optimization through natural evolution. The brilliant color of Morpho butterfly wings is originated from the hierarchical nanostructure on the wing scales. Such nanoarchitecture has attracted a lot of research effort, including the study of its optical properties, its potential use in sensing and infrared imaging, and also the use of such structure as template for the fabrication of high-performance photocatalytic materials. The controlled subtractive processes provide a new path to modify such nanoarchitecture and its optical property. Distinct from previous studies on the optical property of the Morpho wing structure, this study provides additional experimental evidence for the origination of the optical property of the natural butterfly wing scales. The study also offers a facile approach to generate new 3D nanostructures using butterfly wings as the templates and may lead to simpler structure models for large-scale man-made structures than those offered by original butterfly wings. © 2015 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim.

  19. Focusing on butterfly eyespot focus: uncoupling of white spots from eyespot bodies in nymphalid butterflies.

    PubMed

    Iwata, Masaki; Otaki, Joji M

    2016-01-01

    Developmental studies on butterfly wing color patterns often focus on eyespots. A typical eyespot (such as that of Bicyclus anynana) has a few concentric rings of dark and light colors and a white spot (called a focus) at the center. The prospective eyespot center during the early pupal stage is known to act as an organizing center. It has often been assumed, according to gradient models for positional information, that a white spot in adult wings corresponds to an organizing center and that the size of the white spot indicates how active that organizing center was. However, there is no supporting evidence for these assumptions. To evaluate the feasibility of these assumptions in nymphalid butterflies, we studied the unique color patterns of Calisto tasajera (Nymphalidae, Satyrinae), which have not been analyzed before in the literature. In the anterior forewing, one white spot was located at the center of an eyespot, but another white spot associated with either no or only a small eyespot was present in the adjacent compartment. The anterior hindwing contained two adjacent white spots not associated with eyespots, one of which showed a sparse pattern. The posterior hindwing contained two adjacent pear-shaped eyespots, and the white spots were located at the proximal side or even outside the eyespot bodies. The successive white spots within a single compartment along the midline in the posterior hindwing showed a possible trajectory of a positional determination process for the white spots. Several cases of focus-less eyespots in other nymphalid butterflies were also presented. These results argue for the uncoupling of white spots from eyespot bodies, suggesting that an eyespot organizing center does not necessarily differentiate into a white spot and that a prospective white spot does not necessarily signify organizing activity for an eyespot. Incorporation of these results in future models for butterfly wing color pattern formation is encouraged.

  20. What shapes the continuum of reproductive isolation? Lessons from Heliconius butterflies.

    PubMed

    Mérot, C; Salazar, C; Merrill, R M; Jiggins, C D; Joron, M

    2017-06-14

    The process by which species evolve can be illuminated by investigating barriers that limit gene flow between taxa. Recent radiations, such as Heliconius butterflies, offer the opportunity to compare isolation between pairs of taxa at different stages of ecological, geographical, and phylogenetic divergence. Here, we report a comparative analysis of existing and novel data in order to quantify the strength and direction of isolating barriers within a well-studied clade of Heliconius Our results highlight that increased divergence is associated with the accumulation of stronger and more numerous barriers to gene flow. Wing pattern is both under natural selection for Müllerian mimicry and involved in mate choice, and therefore underlies several isolating barriers. However, pairs which share a similar wing pattern also display strong reproductive isolation mediated by traits other than wing pattern. This suggests that, while wing pattern is a key factor for early stages of divergence, it may become facultative at later stages of divergence. Additional factors including habitat partitioning, hybrid sterility, and chemically mediated mate choice are associated with complete speciation. Therefore, although most previous work has emphasized the role of wing pattern, our comparative results highlight that speciation is a multi-dimensional process, whose completion is stabilized by many factors. © 2017 The Author(s).

  1. Longwing (Heliconius) butterflies combine a restricted set of pigmentary and structural coloration mechanisms.

    PubMed

    Wilts, Bodo D; Vey, Aidan J M; Briscoe, Adriana D; Stavenga, Doekele G

    2017-11-21

    Longwing butterflies, Heliconius sp., also called heliconians, are striking examples of diversity and mimicry in butterflies. Heliconians feature strongly colored patterns on their wings, arising from wing scales colored by pigments and/or nanostructures, which serve as an aposematic signal. Here, we investigate the coloration mechanisms among several species of Heliconius by applying scanning electron microscopy, (micro)spectrophotometry, and imaging scatterometry. We identify seven kinds of colored scales within Heliconius whose coloration is derived from pigments, nanostructures or both. In yellow-, orange- and red-colored wing patches, both cover and ground scales contain wavelength-selective absorbing pigments, 3-OH-kynurenine, xanthommatin and/or dihydroxanthommatin. In blue wing patches, the cover scales are blue either due to interference of light in the thin-film lower lamina (e.g., H. doris) or in the multilayered lamellae in the scale ridges (so-called ridge reflectors, e.g., H. sara and H. erato); the underlying ground scales are black. In the white wing patches, both cover and ground scales are blue due to their thin-film lower lamina, but because they are stacked upon each other and at the wing substrate, a faint bluish to white color results. Lastly, green wing patches (H. doris) have cover scales with blue-reflecting thin films and short-wavelength absorbing 3-OH-kynurenine, together causing a green color. The pigmentary and structural traits are discussed in relation to their phylogenetic distribution and the evolution of vision in this highly interesting clade of butterflies.

  2. Gradual and contingent evolutionary emergence of leaf mimicry in butterfly wing patterns.

    PubMed

    Suzuki, Takao K; Tomita, Shuichiro; Sezutsu, Hideki

    2014-11-25

    Special resemblance of animals to natural objects such as leaves provides a representative example of evolutionary adaptation. The existence of such sophisticated features challenges our understanding of how complex adaptive phenotypes evolved. Leaf mimicry typically consists of several pattern elements, the spatial arrangement of which generates the leaf venation-like appearance. However, the process by which leaf patterns evolved remains unclear. In this study we show the evolutionary origin and process for the leaf pattern in Kallima (Nymphalidae) butterflies. Using comparative morphological analyses, we reveal that the wing patterns of Kallima and 45 closely related species share the same ground plan, suggesting that the pattern elements of leaf mimicry have been inherited across species with lineage-specific changes of their character states. On the basis of these analyses, phylogenetic comparative methods estimated past states of the pattern elements and enabled reconstruction of the wing patterns of the most recent common ancestor. This analysis shows that the leaf pattern has evolved through several intermediate patterns. Further, we use Bayesian statistical methods to estimate the temporal order of character-state changes in the pattern elements by which leaf mimesis evolved, and show that the pattern elements changed their spatial arrangement (e.g., from a curved line to a straight line) in a stepwise manner and finally establish a close resemblance to a leaf venation-like appearance. Our study provides the first evidence for stepwise and contingent evolution of leaf mimicry.  Leaf mimicry patterns evolved in a gradual, rather than a sudden, manner from a non-mimetic ancestor. Through a lineage of Kallima butterflies, the leaf patterns evolutionarily originated through temporal accumulation of orchestrated changes in multiple pattern elements.

  3. 76 FR 49541 - Endangered and Threatened Wildlife and Plants; Emergency Listing of the Miami Blue Butterfly as...

    Federal Register 2010, 2011, 2012, 2013, 2014

    2011-08-10

    ... 1993, p. 134). Wings of males are blue above (dorsally), with a narrow black outer border and white... Krizek 1984, p. 112). Seasonal wing pattern variation may be caused by changes in humidity, temperature... the wings (Minno and Emmel 1994, p. 647). The Miami blue can be distinguished from the ceraunus blue...

  4. 77 FR 20947 - Endangered and Threatened Wildlife and Plants; Listing of the Miami Blue Butterfly as Endangered...

    Federal Register 2010, 2011, 2012, 2013, 2014

    2012-04-06

    .... 134). Wings of males are blue above (dorsally), with a narrow black outer border and white fringes... Krizek 1984, p. 112). Seasonal wing pattern variation may be caused by changes in humidity, temperature... on the undersides of the wings (Minno and Emmel 1994, p. 647). The Miami blue can be distinguished...

  5. Colour pattern homology and evolution in Vanessa butterflies (Nymphalidae: Nymphalini): eyespot characters.

    PubMed

    Abbasi, R; Marcus, J M

    2015-11-01

    Ocelli are serially repeated colour patterns on the wings of many butterflies. Eyespots are elaborate ocelli that function in predator avoidance and deterrence as well as in mate choice. A phylogenetic approach was used to study ocelli and eyespot evolution in Vanessa butterflies, a genus exhibiting diverse phenotypes among these serial homologs. Forty-four morphological characters based on eyespot number, arrangement, shape and the number of elements in each eyespot were defined and scored. Ocelli from eight wing cells on the dorsal and ventral surfaces of the forewing and hindwing were evaluated. The evolution of these characters was traced over a phylogeny of Vanessa based on 7750 DNA base pairs from 10 genes. Our reconstruction predicts that the ancestral Vanessa had 5 serially arranged ocelli on all four wing surfaces. The ancestral state on the dorsal forewing and ventral hindwing was ocelli arranged in two heterogeneous groups. On the dorsal hindwing, the ancestral state was either homogenous or ocelli arranged in two heterogeneous groups. On the ventral forewing, we determined that the ancestral state was organized into three heterogeneous groups. In Vanessa, almost all ocelli are individuated and capable of independent evolution relative to other colour patterns except for the ocelli in cells -1 and 0 on the dorsal and ventral forewings, which appear to be constrained to evolve in parallel. The genus Vanessa is a good model system for the study of serial homology and the interaction of selective forces with developmental architecture to produce diversity in butterfly colour patterns. © 2015 European Society For Evolutionary Biology. Journal of Evolutionary Biology © 2015 European Society For Evolutionary Biology.

  6. Distal-less regulates eyespot patterns and melanization in Bicyclus butterflies.

    PubMed

    Monteiro, Antónia; Chen, Bin; Ramos, Diane M; Oliver, Jeffrey C; Tong, Xiaoling; Guo, Min; Wang, Wen-Kai; Fazzino, Lisa; Kamal, Firdous

    2013-07-01

    Butterfly eyespots represent novel complex traits that display substantial diversity in number and size within and across species. Correlative gene expression studies have implicated a large suite of transcription factors, including Distal-less (Dll), Engrailed (En), and Spalt (Sal), in eyespot development in butterflies, but direct evidence testing the function of any of these proteins is still missing. Here we show that the characteristic two-eyespot pattern of wildtype Bicyclus anynana forewings is correlated with dynamic progression of Dll, En, and Sal expression in larval wings from four spots to two spots, whereas no such decline in gene expression ensues in a four-eyespot mutant. We then conduct transgenic experiments testing whether over-expression of any of these genes in a wild-type genetic background is sufficient to induce eyespot differentiation in these pre-patterned wing compartments. We also produce a Dll-RNAi transgenic line to test how Dll down-regulation affects eyespot development. Finally we test how ectopic expression of these genes during the pupal stages of development alters adults color patters. We show that over-expressing Dll in larvae is sufficient to induce the differentiation of additional eyespots and increase the size of eyespots, whereas down-regulating Dll leads to a decrease in eyespot size. Furthermore, ectopic expression of Dll in the early pupal wing led to the appearance of ectopic patches of black scales. We conclude that Dll is a positive regulator of focal differentiation and eyespot signaling and that this gene is also a possible selector gene for scale melanization in butterflies. Copyright © 2013 Wiley Periodicals, Inc.

  7. Morpho morphometrics: Shared ancestry and selection drive the evolution of wing size and shape in Morpho butterflies.

    PubMed

    Chazot, Nicolas; Panara, Stephen; Zilbermann, Nicolas; Blandin, Patrick; Le Poul, Yann; Cornette, Raphaël; Elias, Marianne; Debat, Vincent

    2016-01-01

    Butterfly wings harbor highly diverse phenotypes and are involved in many functions. Wing size and shape result from interactions between adaptive processes, phylogenetic history, and developmental constraints, which are complex to disentangle. Here, we focus on the genus Morpho (Nymphalidae: Satyrinae, 30 species), which presents a high diversity of sizes, shapes, and color patterns. First, we generate a comprehensive molecular phylogeny of these 30 species. Next, using 911 collection specimens, we quantify the variation of wing size and shape across species, to assess the importance of shared ancestry, microhabitat use, and sexual selection in the evolution of the wings. While accounting for phylogenetic and allometric effects, we detect a significant difference in wing shape but not size among microhabitats. Fore and hindwings covary at the individual and species levels, and the covariation differs among microhabitats. However, the microhabitat structure in covariation disappears when phylogenetic relationships are taken into account. Our results demonstrate that microhabitat has driven wing shape evolution, although it has not strongly affected forewing and hindwing integration. We also found that sexual dimorphism of forewing shape and color pattern are coupled, suggesting a common selective force. © 2015 The Author(s). Evolution © 2015 The Society for the Study of Evolution.

  8. Butterfly wing coloration studied with a novel imaging scatterometer

    NASA Astrophysics Data System (ADS)

    Stavenga, Doekele

    2010-03-01

    Animal coloration functions for display or camouflage. Notably insects provide numerous examples of a rich variety of the applied optical mechanisms. For instance, many butterflies feature a distinct dichromatism, that is, the wing coloration of the male and the female differ substantially. The male Brimstone, Gonepteryx rhamni, has yellow wings that are strongly UV iridescent, but the female has white wings with low reflectance in the UV and a high reflectance in the visible wavelength range. In the Small White cabbage butterfly, Pieris rapae crucivora, the wing reflectance of the male is low in the UV and high at visible wavelengths, whereas the wing reflectance of the female is higher in the UV and lower in the visible. Pierid butterflies apply nanosized, strongly scattering beads to achieve their bright coloration. The male Pipevine Swallowtail butterfly, Battus philenor, has dorsal wings with scales functioning as thin film gratings that exhibit polarized iridescence; the dorsal wings of the female are matte black. The polarized iridescence probably functions in intraspecific, sexual signaling, as has been demonstrated in Heliconius butterflies. An example of camouflage is the Green Hairstreak butterfly, Callophrys rubi, where photonic crystal domains exist in the ventral wing scales, resulting in a matte green color that well matches the color of plant leaves. The spectral reflection and polarization characteristics of biological tissues can be rapidly and with unprecedented detail assessed with a novel imaging scatterometer-spectrophotometer, built around an elliptical mirror [1]. Examples of butterfly and damselfly wings, bird feathers, and beetle cuticle will be presented. [4pt] [1] D.G. Stavenga, H.L. Leertouwer, P. Pirih, M.F. Wehling, Optics Express 17, 193-202 (2009)

  9. Effects of structural flexibility of wings in flapping flight of butterfly.

    PubMed

    Senda, Kei; Obara, Takuya; Kitamura, Masahiko; Yokoyama, Naoto; Hirai, Norio; Iima, Makoto

    2012-06-01

    The objective of this paper is to clarify the effects of structural flexibility of wings of a butterfly in flapping flight. For this purpose, a dynamics model of a butterfly is derived by Lagrange's method, where the butterfly is considered as a rigid multi-body system. The panel method is employed to simulate the flow field and the aerodynamic forces acting on the wings. The mathematical model is validated by the agreement of the numerical result with the experimentally measured data. Then, periodic orbits of flapping-of-wings flights are parametrically searched in order to fly the butterfly models. Almost periodic orbits are found, but they are unstable. Deformation of the wings is modeled in two ways. One is bending and its effect on the aerodynamic forces is discussed. The other is passive wing torsion caused by structural flexibility. Numerical simulations demonstrate that flexible torsion reduces the flight instability.

  10. Structural analysis of eyespots: dynamics of morphogenic signals that govern elemental positions in butterfly wings

    PubMed Central

    2012-01-01

    Background To explain eyespot colour-pattern determination in butterfly wings, the induction model has been discussed based on colour-pattern analyses of various butterfly eyespots. However, a detailed structural analysis of eyespots that can serve as a foundation for future studies is still lacking. In this study, fundamental structural rules related to butterfly eyespots are proposed, and the induction model is elaborated in terms of the possible dynamics of morphogenic signals involved in the development of eyespots and parafocal elements (PFEs) based on colour-pattern analysis of the nymphalid butterfly Junonia almana. Results In a well-developed eyespot, the inner black core ring is much wider than the outer black ring; this is termed the inside-wide rule. It appears that signals are wider near the focus of the eyespot and become narrower as they expand. Although fundamental signal dynamics are likely to be based on a reaction-diffusion mechanism, they were described well mathematically as a type of simple uniformly decelerated motion in which signals associated with the outer and inner black rings of eyespots and PFEs are released at different time points, durations, intervals, and initial velocities into a two-dimensional field of fundamentally uniform or graded resistance; this produces eyespots and PFEs that are diverse in size and structure. The inside-wide rule, eyespot distortion, structural differences between small and large eyespots, and structural changes in eyespots and PFEs in response to physiological treatments were explained well using mathematical simulations. Natural colour patterns and previous experimental findings that are not easily explained by the conventional gradient model were also explained reasonably well by the formal mathematical simulations performed in this study. Conclusions In a mode free from speculative molecular interactions, the present study clarifies fundamental structural rules related to butterfly eyespots, delineates a theoretical basis for the induction model, and proposes a mathematically simple mode of long-range signalling that may reflect developmental mechanisms associated with butterfly eyespots. PMID:22409965

  11. Structural analysis of eyespots: dynamics of morphogenic signals that govern elemental positions in butterfly wings.

    PubMed

    Otaki, Joji M

    2012-03-13

    To explain eyespot colour-pattern determination in butterfly wings, the induction model has been discussed based on colour-pattern analyses of various butterfly eyespots. However, a detailed structural analysis of eyespots that can serve as a foundation for future studies is still lacking. In this study, fundamental structural rules related to butterfly eyespots are proposed, and the induction model is elaborated in terms of the possible dynamics of morphogenic signals involved in the development of eyespots and parafocal elements (PFEs) based on colour-pattern analysis of the nymphalid butterfly Junonia almana. In a well-developed eyespot, the inner black core ring is much wider than the outer black ring; this is termed the inside-wide rule. It appears that signals are wider near the focus of the eyespot and become narrower as they expand. Although fundamental signal dynamics are likely to be based on a reaction-diffusion mechanism, they were described well mathematically as a type of simple uniformly decelerated motion in which signals associated with the outer and inner black rings of eyespots and PFEs are released at different time points, durations, intervals, and initial velocities into a two-dimensional field of fundamentally uniform or graded resistance; this produces eyespots and PFEs that are diverse in size and structure. The inside-wide rule, eyespot distortion, structural differences between small and large eyespots, and structural changes in eyespots and PFEs in response to physiological treatments were explained well using mathematical simulations. Natural colour patterns and previous experimental findings that are not easily explained by the conventional gradient model were also explained reasonably well by the formal mathematical simulations performed in this study. In a mode free from speculative molecular interactions, the present study clarifies fundamental structural rules related to butterfly eyespots, delineates a theoretical basis for the induction model, and proposes a mathematically simple mode of long-range signalling that may reflect developmental mechanisms associated with butterfly eyespots.

  12. Wing patterning genes and coevolution of Müllerian mimicry in Heliconius butterflies: Support from phylogeography, cophylogeny, and divergence times.

    PubMed

    Hoyal Cuthill, Jennifer F; Charleston, Michael

    2015-12-01

    Examples of long-term coevolution are rare among free-living organisms. Müllerian mimicry in Heliconius butterflies had been suggested as a key example of coevolution by early genetic studies. However, research over the last two decades has been dominated by the idea that the best-studied comimics, H. erato and H. melpomene, did not coevolve at all. Recently sequenced genes associated with wing color pattern phenotype offer a new opportunity to resolve this controversy. Here, we test the hypothesis of coevolution between H. erato and H. melpomene using Bayesian multilocus analysis of five color pattern genes and five neutral genetic markers. We first explore the extent of phylogenetic agreement versus conflict between the different genes. Coevolution is then tested against three aspects of the mimicry diversifications: phylogenetic branching patterns, divergence times, and, for the first time, phylogeographic histories. We show that all three lines of evidence are compatible with strict coevolution of the diverse mimicry wing patterns, contrary to some recent suggestions. Instead, these findings tally with a coevolutionary diversification driven primarily by the ecological force of Müllerian mimicry. © 2015 The Author(s). Evolution © 2015 The Society for the Study of Evolution.

  13. Genome-wide introgression among distantly related Heliconius butterfly species.

    PubMed

    Zhang, Wei; Dasmahapatra, Kanchon K; Mallet, James; Moreira, Gilson R P; Kronforst, Marcus R

    2016-02-27

    Although hybridization is thought to be relatively rare in animals, the raw genetic material introduced via introgression may play an important role in fueling adaptation and adaptive radiation. The butterfly genus Heliconius is an excellent system to study hybridization and introgression but most studies have focused on closely related species such as H. cydno and H. melpomene. Here we characterize genome-wide patterns of introgression between H. besckei, the only species with a red and yellow banded 'postman' wing pattern in the tiger-striped silvaniform clade, and co-mimetic H. melpomene nanna. We find a pronounced signature of putative introgression from H. melpomene into H. besckei in the genomic region upstream of the gene optix, known to control red wing patterning, suggesting adaptive introgression of wing pattern mimicry between these two distantly related species. At least 39 additional genomic regions show signals of introgression as strong or stronger than this mimicry locus. Gene flow has been on-going, with evidence of gene exchange at multiple time points, and bidirectional, moving from the melpomene to the silvaniform clade and vice versa. The history of gene exchange has also been complex, with contributions from multiple silvaniform species in addition to H. besckei. We also detect a signature of ancient introgression of the entire Z chromosome between the silvaniform and melpomene/cydno clades. Our study provides a genome-wide portrait of introgression between distantly related butterfly species. We further propose a comprehensive and efficient workflow for gene flow identification in genomic data sets.

  14. Extensive transcriptional response associated with seasonal plasticity of butterfly wing patterns.

    PubMed

    Daniels, Emily V; Murad, Rabi; Mortazavi, Ali; Reed, Robert D

    2014-12-01

    In the eastern United States, the buckeye butterfly, Junonia coenia, shows seasonal wing colour plasticity where adults emerging in the spring are tan, while those emerging in the autumn are dark red. This variation can be artificially induced in laboratory colonies, thus making J. coenia a useful model system to examine the mechanistic basis of plasticity. To better understand the developmental basis of seasonal plasticity, we used RNA-seq to quantify transcription profiles associated with development of alternative seasonal wing morphs. Depending on the developmental stage, between 547 and 1420 transfrags were significantly differentially expressed between morphs. These extensive differences in gene expression stand in contrast to the much smaller numbers of differentially expressed transcripts identified in previous studies of genetic wing pattern variation in other species and suggest that environmentally induced phenotypic shifts arise from very broad systemic processes. Analyses of candidate endocrine and pigmentation transcripts revealed notable genes upregulated in the red morph, including several ecdysone-associated genes, and cinnabar, an ommochrome pigmentation gene implicated in colour pattern variation in other butterflies. We also found multiple melanin-related transcripts strongly upregulated in the red morph, including tan and yellow-family genes, leading us to speculate that dark red pigmentation in autumn J. coenia may involve nonommochrome pigments. While we identified several endocrine and pigmentation genes as obvious candidates for seasonal colour morph differentiation, we speculate that the majority of observed expression differences were due to thermal stress response. The buckeye transcriptome provides a basis for further developmental studies of phenotypic plasticity. © 2014 John Wiley & Sons Ltd.

  15. Dynamics of F-actin prefigure the structure of butterfly wing scales.

    PubMed

    Dinwiddie, April; Null, Ryan; Pizzano, Maria; Chuong, Lisa; Leigh Krup, Alexis; Ee Tan, Hwei; Patel, Nipam H

    2014-08-15

    The wings of butterflies and moths consist of dorsal and ventral epidermal surfaces that give rise to overlapping layers of scales and hairs (Lepidoptera, "scale wing"). Wing scales (average length ~200 µm) are homologous to insect bristles (macrochaetes), and their colors create the patterns that characterize lepidopteran wings. The topology and surface sculpture of wing scales vary widely, and this architectural complexity arises from variations in the developmental program of the individual scale cells of the wing epithelium. One of the more striking features of lepidopteran wing scales are the longitudinal ridges that run the length of the mature (dead) cell, gathering the cuticularized scale cell surface into pleats on the sides of each scale. While also present around the periphery of other insect bristles and hairs, longitudinal ridges in lepidopteran wing scales gain new significance for their creation of iridescent color through microribs and lamellae. Here we show the dynamics of the highly organized F-actin filaments during scale cell development, and present experimental manipulations of actin polymerization that reveal the essential role of this cytoskeletal component in wing scale elongation and the positioning of longitudinal ribs. Copyright © 2014 Elsevier Inc. All rights reserved.

  16. Refractive index dependence of Papilio Ulysses butterfly wings reflectance spectra

    NASA Astrophysics Data System (ADS)

    Isnaeni, Muslimin, Ahmad Novi; Birowosuto, Muhammad Danang

    2016-02-01

    We have observed and utilized butterfly wings of Papilio Ulysses for refractive index sensor. We noticed this butterfly wings have photonic crystal structure, which causes blue color appearance on the wings. The photonic crystal structure, which consists of cuticle and air void, is approximated as one dimensional photonic crystal structure. This photonic crystal structure opens potential to several optical devices application, such as refractive index sensor. We have utilized small piece of Papilio Ulysses butterfly wings to characterize refractive index of several liquid base on reflectance spectrum of butterfly wings in the presence of sample liquid. For comparison, we simulated reflectance spectrum of one dimensional photonic crystal structure having material parameter based on real structure of butterfly wings. We found that reflectance spectrum peaks shifted as refractive index of sample changes. Although there is a slight difference in reflectance spectrum peaks between measured spectrum and calculated spectrum, the trend of reflectance spectrum peaks as function of sample's refractive index is the similar. We assume that during the measurement, the air void that filled by sample liquid is expanded due to liquid pressure. This change of void shape causes non-similarity between measured spectrum and calculated spectrum.

  17. Color-pattern analysis of eyespots in butterfly wings: a critical examination of morphogen gradient models.

    PubMed

    Otaki, Joji M

    2011-06-01

    Butterfly wing color patterns consist of many color-pattern elements such as eyespots. It is believed that eyespot patterns are determined by a concentration gradient of a single morphogen species released by diffusion from the prospective eyespot focus in conjunction with multiple thresholds in signal-receiving cells. As alternatives to this single-morphogen model, more flexible multiple-morphogen model and induction model can be proposed. However, the relevance of these conceptual models to actual eyespots has not been examined systematically. Here, representative eyespots from nymphalid butterflies were analyzed morphologically to determine if they are consistent with these models. Measurement of ring widths of serial eyespots from a single wing surface showed that the proportion of each ring in an eyespot is quite different among homologous rings of serial eyespots of different sizes. In asymmetric eyespots, each ring is distorted to varying degrees. In extreme cases, only a portion of rings is expressed remotely from the focus. Similarly, there are many eyespots where only certain rings are deleted, added, or expanded. In an unusual case, the central area of an eyespot is composed of multiple "miniature eyespots," but the overall macroscopic eyespot structure is maintained. These results indicate that each eyespot ring has independence and flexibility to a certain degree, which is less consistent with the single-morphogen model. Considering a "periodic eyespot", which has repeats of a set of rings, damage-induced eyespots in mutants, and a scale-size distribution pattern in an eyespot, the induction model is the least incompatible with the actual eyespot diversity.

  18. Cruising the rain forest floor: butterfly wing shape evolution and gliding in ground effect.

    PubMed

    Cespedes, Ann; Penz, Carla M; DeVries, Philip J

    2015-05-01

    Flight is a key innovation in the evolutionary success of insects and essential to dispersal, territoriality, courtship and oviposition. Wing shape influences flight performance and selection likely acts to maximize performance for conducting essential behaviours that in turn results in the evolution of wing shape. As wing shape also contributes to fitness, optimal shapes for particular flight behaviours can be assessed with aerodynamic predictions and placed in an ecomorphological context. Butterflies in the tribe Haeterini (Nymphalidae) are conspicuous members of understorey faunas in lowland Neotropical forests. Field observations indicate that the five genera in this clade differ in flight height and behaviour: four use gliding flight at the forest floor level, and one utilizes flapping flight above the forest floor. Nonetheless, the association of ground level gliding flight behaviour and wing shape has never been investigated in this or any other butterfly group. We used landmark-based geometric morphometrics to test whether wing shapes in Haeterini and their close relatives reflected observed flight behaviours. Four genera of Haeterini and some distantly related Satyrinae showed significant correspondence between wing shape and theoretical expectations in performance trade-offs that we attribute to selection for gliding in ground effect. Forewing shape differed between sexes for all taxa, and male wing shapes were aerodynamically more efficient for gliding flight than corresponding females. This suggests selection acts differentially on male and female wing shapes, reinforcing the idea that sex-specific flight behaviours contribute to the evolution of sexual dimorphism. Our study indicates that wing shapes in Haeterini butterflies evolved in response to habitat-specific flight behaviours, namely gliding in ground effect along the forest floor, resulting in ecomorphological partitions of taxa in morphospace. The convergent flight behaviour and wing morphology between tribes of Satyrinae suggest that the flight environment may offset phylogenetic constraints. Overall, this study provides a basis for exploring similar patterns of wing shape evolution in other taxa that glide in ground effect. © 2014 The Authors. Journal of Animal Ecology © 2014 British Ecological Society.

  19. Study of nano-architecture of the wings of Paris Peacock butterfly

    NASA Astrophysics Data System (ADS)

    Ghate, Ekata; Bhoraskar, S. V.; Kulkarni, G. R.

    Butterflies are one of the most colorful creatures in animal Kingdom. Wings of the male butterfly are brilliantly colored to attract females. Color of the wings plays an important role in camouflage. Study of structural colors in case of insects and butterflies are important for their biomimic and biophotonic applications. Structural color is the color which is produced by physical structures and their interaction with light. Paris Peacock or Papilio paris butterfly belongs to the family Papilionidae. The basis of structural color of this butterfly is investigated in the present study. The upper surface of the wings in this butterfly is covered with blue, green and brown colored scales. Nano-architecture of these scales was investigated with scanning electron microscope (SEM) and environmental scanning electron microscope (ESEM). Photomicrographs were analyzed using image analysis software. Goniometric color or iridescence in blue and green colored scales of this butterfly was observed and studied with the help of gonio spectrophotometer in the visible range. No iridescence was observed in brown colored scales of the butterfly. Hues of the blue and green color were measured with spectrophotometer and were correlated with nano-architecture of the wing. Results of electron microscopy and reflection spectroscopy are used to explain the iridescent nature of blue and green scales. Sinusoidal grating like structures of these scales were prominently seen in the blue scales. It is possible that the structure of these wings can act as a template for the fabrication of sinusoidal gratings using nano-imprint technology.

  20. Conservatism and novelty in the genetic architecture of adaptation in Heliconius butterflies.

    PubMed

    Huber, B; Whibley, A; Poul, Y L; Navarro, N; Martin, A; Baxter, S; Shah, A; Gilles, B; Wirth, T; McMillan, W O; Joron, M

    2015-05-01

    Understanding the genetic architecture of adaptive traits has been at the centre of modern evolutionary biology since Fisher; however, evaluating how the genetic architecture of ecologically important traits influences their diversification has been hampered by the scarcity of empirical data. Now, high-throughput genomics facilitates the detailed exploration of variation in the genome-to-phenotype map among closely related taxa. Here, we investigate the evolution of wing pattern diversity in Heliconius, a clade of neotropical butterflies that have undergone an adaptive radiation for wing-pattern mimicry and are influenced by distinct selection regimes. Using crosses between natural wing-pattern variants, we used genome-wide restriction site-associated DNA (RAD) genotyping, traditional linkage mapping and multivariate image analysis to study the evolution of the architecture of adaptive variation in two closely related species: Heliconius hecale and H. ismenius. We implemented a new morphometric procedure for the analysis of whole-wing pattern variation, which allows visualising spatial heatmaps of genotype-to-phenotype association for each quantitative trait locus separately. We used the H. melpomene reference genome to fine-map variation for each major wing-patterning region uncovered, evaluated the role of candidate genes and compared genetic architectures across the genus. Our results show that, although the loci responding to mimicry selection are highly conserved between species, their effect size and phenotypic action vary throughout the clade. Multilocus architecture is ancestral and maintained across species under directional selection, whereas the single-locus (supergene) inheritance controlling polymorphism in H. numata appears to have evolved only once. Nevertheless, the conservatism in the wing-patterning toolkit found throughout the genus does not appear to constrain phenotypic evolution towards local adaptive optima.

  1. Time-varying wing-twist improves aerodynamic efficiency of forward flight in butterflies.

    PubMed

    Zheng, Lingxiao; Hedrick, Tyson L; Mittal, Rajat

    2013-01-01

    Insect wings can undergo significant chordwise (camber) as well as spanwise (twist) deformation during flapping flight but the effect of these deformations is not well understood. The shape and size of butterfly wings leads to particularly large wing deformations, making them an ideal test case for investigation of these effects. Here we use computational models derived from experiments on free-flying butterflies to understand the effect of time-varying twist and camber on the aerodynamic performance of these insects. High-speed videogrammetry is used to capture the wing kinematics, including deformation, of a Painted Lady butterfly (Vanessa cardui) in untethered, forward flight. These experimental results are then analyzed computationally using a high-fidelity, three-dimensional, unsteady Navier-Stokes flow solver. For comparison to this case, a set of non-deforming, flat-plate wing (FPW) models of wing motion are synthesized and subjected to the same analysis along with a wing model that matches the time-varying wing-twist observed for the butterfly, but has no deformation in camber. The simulations show that the observed butterfly wing (OBW) outperforms all the flat-plate wings in terms of usable force production as well as the ratio of lift to power by at least 29% and 46%, respectively. This increase in efficiency of lift production is at least three-fold greater than reported for other insects. Interestingly, we also find that the twist-only-wing (TOW) model recovers much of the performance of the OBW, demonstrating that wing-twist, and not camber is key to forward flight in these insects. The implications of this on the design of flapping wing micro-aerial vehicles are discussed.

  2. Time-Varying Wing-Twist Improves Aerodynamic Efficiency of Forward Flight in Butterflies

    PubMed Central

    Zheng, Lingxiao; Hedrick, Tyson L.; Mittal, Rajat

    2013-01-01

    Insect wings can undergo significant chordwise (camber) as well as spanwise (twist) deformation during flapping flight but the effect of these deformations is not well understood. The shape and size of butterfly wings leads to particularly large wing deformations, making them an ideal test case for investigation of these effects. Here we use computational models derived from experiments on free-flying butterflies to understand the effect of time-varying twist and camber on the aerodynamic performance of these insects. High-speed videogrammetry is used to capture the wing kinematics, including deformation, of a Painted Lady butterfly (Vanessa cardui) in untethered, forward flight. These experimental results are then analyzed computationally using a high-fidelity, three-dimensional, unsteady Navier-Stokes flow solver. For comparison to this case, a set of non-deforming, flat-plate wing (FPW) models of wing motion are synthesized and subjected to the same analysis along with a wing model that matches the time-varying wing-twist observed for the butterfly, but has no deformation in camber. The simulations show that the observed butterfly wing (OBW) outperforms all the flat-plate wings in terms of usable force production as well as the ratio of lift to power by at least 29% and 46%, respectively. This increase in efficiency of lift production is at least three-fold greater than reported for other insects. Interestingly, we also find that the twist-only-wing (TOW) model recovers much of the performance of the OBW, demonstrating that wing-twist, and not camber is key to forward flight in these insects. The implications of this on the design of flapping wing micro-aerial vehicles are discussed. PMID:23341923

  3. Complex dynamics underlie the evolution of imperfect wing pattern convergence in butterflies.

    PubMed

    Finkbeiner, Susan D; Briscoe, Adriana D; Mullen, Sean P

    2017-04-01

    Adaptive radiation is characterized by rapid diversification that is strongly associated with ecological specialization. However, understanding the evolutionary mechanisms fueling adaptive diversification requires a detailed knowledge of how natural selection acts at multiple life-history stages. Butterflies within the genus Adelpha represent one of the largest and most diverse butterfly lineages in the Neotropics. Although Adelpha species feed on an extraordinary diversity of larval hosts, convergent evolution is widespread in this group, suggesting that selection for mimicry may contribute to adaptive divergence among species. To investigate this hypothesis, we conducted predation studies in Costa Rica using artificial butterfly facsimiles. Specifically, we predicted that nontoxic, palatable Adelpha species that do not feed on host plants in the family Rubiaceae would benefit from sharing a locally convergent wing pattern with the presumably toxic Rubiaceae-feeding species via reduced predation. Contrary to expectations, we found that the presumed mimic was attacked significantly more than its locally convergent model at a frequency paralleling attack rates on both novel and palatable prey. Although these data reveal the first evidence for protection from avian predators by the supposed toxic, Rubiaceae-feeding Adelpha species, we conclude that imprecise mimetic patterns have high costs for Batesian mimics in the tropics. © 2017 The Author(s). Evolution © 2017 The Society for the Study of Evolution.

  4. Coloration principles of nymphaline butterflies - thin films, melanin, ommochromes and wing scale stacking.

    PubMed

    Stavenga, Doekele G; Leertouwer, Hein L; Wilts, Bodo D

    2014-06-15

    The coloration of the common butterflies Aglais urticae (small tortoiseshell), Aglais io (peacock) and Vanessa atalanta (red admiral), belonging to the butterfly subfamily Nymphalinae, is due to the species-specific patterning of differently coloured scales on their wings. We investigated the scales' structural and pigmentary properties by applying scanning electron microscopy, (micro)spectrophotometry and imaging scatterometry. The anatomy of the wing scales appears to be basically identical, with an approximately flat lower lamina connected by trabeculae to a highly structured upper lamina, which consists of an array of longitudinal, parallel ridges and transversal crossribs. Isolated scales observed at the abwing (upper) side are blue, yellow, orange, red, brown or black, depending on their pigmentation. The yellow, orange and red scales contain various amounts of 3-OH-kynurenine and ommochrome pigment, black scales contain a high density of melanin, and blue scales have a minor amount of melanin pigment. Observing the scales from their adwing (lower) side always revealed a structural colour, which is blue in the case of blue, red and black scales, but orange for orange scales. The structural colours are created by the lower lamina, which acts as an optical thin film. Its reflectance spectrum, crucially determined by the lamina thickness, appears to be well tuned to the scales' pigmentary spectrum. The colours observed locally on the wing are also due to the degree of scale stacking. Thin films, tuned pigments and combinations of stacked scales together determine the wing coloration of nymphaline butterflies. © 2014. Published by The Company of Biologists Ltd.

  5. Flutter-by Interactive Butterfly Using interactivity to excite and educate children about butterflies and the National Museum of Play at The Strong's Dancing Wings Butterfly Garden

    NASA Astrophysics Data System (ADS)

    Powers, Lydia

    The National Museum of Play at The Strong's Dancing Wings Butterfly Garden is a tropical rainforest that allows visitors to step into the world of butterflies, but lacks a more comprehensive educational element to teach visitors additional information about butterflies. Flutter-by Interactive Butterfly is a thesis project designed to enhance younger visitors' experience of the Dancing Wings Butterfly Garden with an interactive educational application that aligns with The Strong's mission of encouraging learning, creativity, and discovery. This was accomplished through a series of fun and educational games and animations, designed for use as a kiosk outside the garden and as a part of The Strong's website. Content, planning, and organization of this project has been completed through research and observation of the garden in the following areas: its visitors, butterflies, best usability practices for children, and game elements that educate and engage children. Flutter-by Interactive Butterfly teaches users about the butterfly's life cycle, anatomy, and characteristics as well as their life in the Dancing Wings Butterfly Garden. Through the use of the design programs Adobe Illustrator, Flash, and After Effects; the programming language ActionScript3.0; a child-friendly user interface and design; audio elements and user takeaways, Flutter-by Interactive Butterfly appeals to children of all ages, interests, and learning styles. The project can be viewed at lydiapowers.com/Thesis/FlutterByButterfly.html

  6. White butterflies as solar photovoltaic concentrators.

    PubMed

    Shanks, Katie; Senthilarasu, S; Ffrench-Constant, Richard H; Mallick, Tapas K

    2015-07-31

    Man's harvesting of photovoltaic energy requires the deployment of extensive arrays of solar panels. To improve both the gathering of thermal and photovoltaic energy from the sun we have examined the concept of biomimicry in white butterflies of the family Pieridae. We tested the hypothesis that the V-shaped posture of basking white butterflies mimics the V-trough concentrator which is designed to increase solar input to photovoltaic cells. These solar concentrators improve harvesting efficiency but are both heavy and bulky, severely limiting their deployment. Here, we show that the attachment of butterfly wings to a solar cell increases its output power by 42.3%, proving that the wings are indeed highly reflective. Importantly, and relative to current concentrators, the wings improve the power to weight ratio of the overall structure 17-fold, vastly expanding their potential application. Moreover, a single mono-layer of scale cells removed from the butterflies' wings maintained this high reflectivity showing that a single layer of scale cell-like structures can also form a useful coating. As predicted, the wings increased the temperature of the butterflies' thorax dramatically, showing that the V-shaped basking posture of white butterflies has indeed evolved to increase the temperature of their flight muscles prior to take-off.

  7. Butterfly eyespot serial homology: enter the Hox genes

    PubMed Central

    2011-01-01

    Hox genes modify serial homology patterns in many organisms, exemplified in vertebrates by modification of the axial skeleton and in arthropods by diversification of the body segments. Butterfly wing eyespots also appear in a serial homologous pattern that, in certain species, is subject to local modification. A paper in EvoDevo reports the Hox gene Antp is the earliest known gene to have eyespot-specific expression; however, not all Lepidoptera express Antp in eyespots, suggesting some developmental flexibility. See research article: http://www.evodevojournal.com/content/2/1/9 PMID:21527048

  8. Adaptive introgression across species boundaries in Heliconius butterflies.

    PubMed

    Pardo-Diaz, Carolina; Salazar, Camilo; Baxter, Simon W; Merot, Claire; Figueiredo-Ready, Wilsea; Joron, Mathieu; McMillan, W Owen; Jiggins, Chris D

    2012-01-01

    It is widely documented that hybridisation occurs between many closely related species, but the importance of introgression in adaptive evolution remains unclear, especially in animals. Here, we have examined the role of introgressive hybridisation in transferring adaptations between mimetic Heliconius butterflies, taking advantage of the recent identification of a gene regulating red wing patterns in this genus. By sequencing regions both linked and unlinked to the red colour locus, we found a region that displays an almost perfect genotype by phenotype association across four species, H. melpomene, H. cydno, H. timareta, and H. heurippa. This particular segment is located 70 kb downstream of the red colour specification gene optix, and coalescent analysis indicates repeated introgression of adaptive alleles from H. melpomene into the H. cydno species clade. Our analytical methods complement recent genome scale data for the same region and suggest adaptive introgression has a crucial role in generating adaptive wing colour diversity in this group of butterflies.

  9. Hormonal control of GTP cyclohydrolase I gene expression and enzyme activity during color pattern development in wings of Precis coenia.

    PubMed

    Sawada, H; Nakagoshi, M; Reinhardt, R K; Ziegler, I; Koch, P B

    2002-06-01

    Color patterns of butterfly wings are composed of single color points represented by each scale. In the case of Precis coenia, at the end of pupal development, different types of pigments are synthesized sequentially in the differently colored scales beginning with white (pterins) followed by red (ommatins) and then black (melanin). In order to explain how formation of these different colors is regulated, we examined the expression of an mRNA-encoding guanosine triphosphate-cyclohydrolase I (GTP-CH I; EC 3.5.4.16), the first key enzyme in the biosynthesis of pteridines, during pigment formation in the wings of P. coenia. The strongest positive signal was recognized around pigment formation one day before butterfly emergence. This GTP-CH I gene expression is paralleled by GTP-CH I enzyme activity measured in wing extracts. We also investigated the effect of 20-hydroxyecdysone on the expression of GTP-CH I mRNA and the enzyme activity during color formation. The results strongly suggest that the onset and duration of the expression of a GTP-CH I mRNA is triggered by a declining ecdysteroid hormone titer during late pupal development.

  10. Three-Dimensional CdS/Au Butterfly Wing Scales with Hierarchical Rib Structures for Plasmon-Enhanced Photocatalytic Hydrogen Production.

    PubMed

    Fang, Jing; Gu, Jiajun; Liu, Qinglei; Zhang, Wang; Su, Huilan; Zhang, Di

    2018-06-13

    Localized surface plasmon resonance (LSPR) of plasmonic metals (e.g., Au) can help semiconductors improve their photocatalytic hydrogen (H 2 ) production performance. However, an artificial synthesis of hierarchical plasmonic structures down to nanoscales is usually difficult. Here, we adopt the butterfly wing scales from Morpho didius to fabricate three-dimensional (3D) CdS/Au butterfly wing scales for plasmonic photocatalysis. The as-prepared materials well-inherit the pristine hierarchical biostructures. The 3D CdS/Au butterfly wing scales exhibit a high H 2 production rate (221.8 μmol·h -1 within 420-780 nm), showing a 241-fold increase over the CdS butterfly wing scales. This is attributed to the effective potentiation effect of LSPR introduced by multilayer metallic rib structures and a good interface bonding state between Au and CdS nanoparticles. Thus, our study provides a relatively simple method to learn from nature and inspiration for preparing highly efficient plasmonic photocatalysts.

  11. Theoretical and experimental analysis of the structural pattern responsible for the iridescence of Morpho butterflies.

    PubMed

    Siddique, Radwanul Hasan; Diewald, Silvia; Leuthold, Juerg; Hölscher, Hendrik

    2013-06-17

    Morpho butterflies are well-known for their iridescence originating from nanostructures in the scales of their wings. These optical active structures integrate three design principles leading to the wide angle reflection: alternating lamellae layers, "Christmas tree" like shape, and offsets between neighboring ridges. We study their individual effects rigorously by 2D FEM simulations of the nanostructures of the Morpho sulkowskyi butterfly and show how the reflection spectrum can be controlled by the design of the nanostructures. The width of the spectrum is broad (≈ 90 nm) for alternating lamellae layers (or "brunches") of the structure while the "Christmas tree" pattern together with a height offset between neighboring ridges reduces the directionality of the reflectance. Furthermore, we fabricated the simulated structures by e-beam lithography. The resulting samples mimicked all important optical features of the original Morpho butterfly scales and feature the intense blue iridescence with a wide angular range of reflection.

  12. The effect of butterfly scales on flight efficiency and leading edge vortex formation

    NASA Astrophysics Data System (ADS)

    Lang, Amy; Wilroy, Jacob; Wahidi, Redha; Slegers, Nathan; Heilman, Micahel; Cranford, Jacob

    2016-11-01

    It is hypothesized that the scales on a butterfly wing lead to increased flight efficiency. Recent testing of live butterflies tracked their motion over 246 flights for 11 different specimens. Results show a 37.8 percent mean decrease in flight efficiency and a flapping amplitude reduction of 6.7 percent once the scales were removed. This change could be largely a result of how the leading edge vortex (LEV) interacts with the wing. To simplify this complex flow problem, an experiment was designed to focus on the alteration of 2-D vortex development with a variation in surface patterning. Specifically, the secondary vorticity generated by the LEV interacting at the patterned surface was studied, as well as the subsequent effect on the LEV's growth rate and peak circulation. For this experiment butterfly inspired grooves were created using additive manufacturing and were attached to a flat plate with a chordwise orientation, thus increasing plate surface area. The vortex generated by the grooved plate was then compared to a smooth case as the plate translated vertically through a tow tank at Re = 1500, 3000, and 6000. Using DPIV, the vortex formation was documented and a maximum vortex formation time of 4.22 was found based on the flat plate travel distance and chord length. Results indicate that the patterned surface slows down the growth of the vortex which corroborates the flight test results. Funding from NSF CBET Fluid Dynamcis is gratefully acknowledged.

  13. Magnetron sputtering in the creation of photonic nanostructures derived from Sasakia Charonda Formosana-butterfly wings for applied in dye-sensitized solar cells

    NASA Astrophysics Data System (ADS)

    Niu, Haihong; Zhou, Ru; Cheng, Cong; Zhang, Gonghai; Hu, Yu; Huang, Bin; Zhang, Shouwei; Shang, Xin; Xia, Mei; Xu, Jinzhang

    2016-09-01

    Creating new functional materials derived from the structures seen on butterfly wings has achieved interest in a variety of research topics. However, there need a concision approach could result in a high-quality, precise, and convenient process for the fabrication of complex nanostructures replication with unique functionalities based on the butterfly wings. Here we developed a pithy approach based on a magnetron sputtering metal Ti process for biotemplating used to refine hierarchically porous titanium dioxide photonic crystal nanostructures (TiO2sbnd PCN), themselves derived from nanostructures present on the wings of Sasakia Charonda Formosana (S. Charonda) butterflies. For the first time, the TiO2sbnd PCN were deposited on the top of the P25 active layer and were used to fabricate DSSCs as the light-scattering layers of photoanodes with power conversion efficiencies of up to 8.7%. Remarkably, a much enhanced photocurrent density and a prominent photoelectrochemical conversion capability have been achieved, which are exceeding most of the previously reported photoanodes as well as a similar butterflies replication-based device structure. Our study suggests many exciting opportunities of developing artificially engineered butterfly wing-based solar-to-fuel conversion.

  14. Butterfly wing color: A photonic crystal demonstration

    NASA Astrophysics Data System (ADS)

    Proietti Zaccaria, Remo

    2016-01-01

    We have theoretically modeled the optical behavior of a natural occurring photonic crystal, as defined by the geometrical characteristics of the Teinopalpus Imperialis butterfly. In particular, following a genetic algorithm approach, we demonstrate how its wings follow a triclinic crystal geometry with a tetrahedron unit base. By performing both photonic band analysis and transmission/reflection simulations, we are able to explain the characteristic colors emerging by the butterfly wings, thus confirming their crystal form.

  15. Germline transformation of the butterfly Bicyclus anynana.

    PubMed

    Marcus, Jeffrey M; Ramos, Diane M; Monteiro, Antónia

    2004-08-07

    Ecological and evolutionary theory has frequently been inspired by the diversity of colour patterns on the wings of butterflies. More recently, these varied patterns have also become model systems for studying the evolution of developmental mechanisms. A technique that will facilitate our understanding of butterfly colour-pattern development is germline transformation. Germline transformation permits functional tests of candidate gene products and of cis-regulatory regions, and provides a means of generating new colour-pattern mutants by insertional mutagenesis. We report the successful transformation of the African satyrid butterfly Bicyclus anynana with two different transposable element vectors, Hermes and piggyBac, each carrying EGFP coding sequences driven by the 3XP3 synthetic enhancer that drives gene expression in the eyes. Candidate lines identified by screening for EGFP in adult eyes were later confirmed by PCR amplification of a fragment of the EGFP coding sequence from genomic DNA. Flanking DNA surrounding the insertions was amplified by inverse PCR and sequenced. Transformation rates were 5% for piggyBac and 10.2% for Hermes. Ultimately, the new data generated by these techniques may permit an integrated understanding of the developmental genetics of colour-pattern formation and of the ecological and evolutionary processes in which these patterns play a role.

  16. Insect Wing Displacement Measurement Using Digital Holography

    DOE Office of Scientific and Technical Information (OSTI.GOV)

    Aguayo, Daniel D.; Mendoza Santoyo, Fernando; Torre I, Manuel H. de la

    2008-04-15

    Insects in flight have been studied with optical non destructive techniques with the purpose of using meaningful results in aerodynamics. With the availability of high resolution and large dynamic range CCD sensors the so called interferometric digital holographic technique was used to measure the surface displacement of in flight insect wings, such as butterflies. The wings were illuminated with a continuous wave Verdi laser at 532 nm, and observed with a CCD Pixelfly camera that acquire images at a rate of 11.5 frames per second at a resolution of 1392x1024 pixels and 12 Bit dynamic range. At this frame ratemore » digital holograms of the wings were captured and processed in the usual manner, namely, each individual hologram is Fourier processed in order to find the amplitude and phase corresponding to the digital hologram. The wings displacement is obtained when subtraction between two digital holograms is performed for two different wings position, a feature applied to all consecutive frames recorded. The result of subtracting is seen as a wrapped phase fringe pattern directly related to the wing displacement. The experimental data for different butterfly flying conditions and exposure times are shown as wire mesh plots in a movie of the wings displacement.« less

  17. Geographic variation in ultraviolet reflectance of the wings of the female cabbage butterfly, Pieris rapae.

    PubMed

    Obara, Yoshiaki; Ozawa, Gaku; Fukano, Yuya

    2008-11-01

    The British and Japanese subspecies of the cabbage butterfly, Pieris rapae , differ in terms of the UV reflectance of their wings ( Obara and Majerus, 2000 ). We studied the biogeographical distribution of the female cabbage butterfly having wings with UV reflectance around the Eurasian continent, and between Britain and Japan. For the study, we collected specimens from various locations. A gradient in the UV reflectance of the wings appears to exist along the west-east axis; reflectance was higher toward the east and reached a peak in butterflies in Japan. The UV-reflecting Japanese subspecies Pieris rapae crucivora was found exclusively along the east coast of the Eurasian continent. This suggests that the Japanese subspecies has evolved from a continental ancestor, with females having UV-absorbing wings. We discuss the results of our study with regard to the evolution and adaptive significance of UV coloration in the Japanese subspecies.

  18. Investigation of Surface Enhanced Coherent Raman Scattering on Nano-patterned Insect Wings

    NASA Astrophysics Data System (ADS)

    Ujj, Laszlo; Lawhead, Carlos

    2015-03-01

    Many insect wings (cicadas, butterflies, mosquitos) poses nano-patterned surface structure. Characterization of surface morphology and chemical composition of insect wings is important to understand the extreme mechanical properties and the biophysical functionalities of the wings. We have measured the image of the membrane of a cicada's wing with the help of Scanning Electron Microscopy (SEM). The results confirm the existing periodic structure of the wing measured previously. In order to identify the chemical composition of the wing, we have deposited silver nanoparticles on it and applied Coherent anti-Stokes Raman Spectroscopy to measure the vibrational spectra of the molecules comprising the wing for the first time. The measured spectra are consistent with the original assumption that the wing membrane is composed of protein, wax, and chitin. The results of these studies can be used to measure other nano-patterned surfaces and to make artificial materials in the future. Authors grateful for financial support from the Department of Physics of the College of Sciences Engineering and Health of UWF and the Pall Corporation for SEM imaging.

  19. Asymmetric ratchet effect for directional transport of fog drops on static and dynamic butterfly wings.

    PubMed

    Liu, Chengcheng; Ju, Jie; Zheng, Yongmei; Jiang, Lei

    2014-02-25

    Inspired by novel creatures, researchers have developed varieties of fog drop transport systems and made significant contributions to the fields of heat transferring, water collecting, antifogging, and so on. Up to now, most of the efforts in directional fog drop transport have been focused on static surfaces. Considering it is not practical to keep surfaces still all the time in reality, conducting investigations on surfaces that can transport fog drops in both static and dynamic states has become more and more important. Here we report the wings of Morpho deidamia butterflies can directionally transport fog drops in both static and dynamic states. This directional drop transport ability results from the micro/nano ratchet-like structure of butterfly wings: the surface of butterfly wings is composed of overlapped scales, and the scales are covered with porous asymmetric ridges. Influenced by this special structure, fog drops on static wings are transported directionally as a result of the fog drops' asymmetric growth and coalescence. Fog drops on vibrating wings are propelled directionally due to the fog drops' asymmetric dewetting from the wings.

  20. Neighboring genes shaping a single adaptive mimetic trait.

    PubMed

    Pardo-Diaz, Carolina; Jiggins, Chris D

    2014-01-01

    The colorful wing patterns of Heliconius butterflies represent an excellent system in which to study the genetic and developmental control of adaptation and convergence. Using qRT-PCR and in situ hybridization on developing wings of the co-mimic species Heliconius melpomene and Heliconius erato, we have profiled the expression of three candidate genes located in the genomic locus controlling red color pattern variation. We found convergent domains of gene expression in H. melpomene and H. erato associated with red wing elements in the two genes optix and kinesin. During early pupal development of both species, the expression of optix perfectly associated with all red pattern elements whereas that of kinesin was specifically correlated with the presence of the red forewing band. These results provide evidence for the use of these two tightly linked patterning genes, acting together to create convergent wing phenotypes in Heliconius and constituting a hotspot of adaptation. © 2013 Wiley Periodicals, Inc.

  1. Aerodynamic evaluation of wing shape and wing orientation in four butterfly species using numerical simulations and a low-speed wind tunnel, and its implications for the design of flying micro-robots.

    PubMed

    Ortega Ancel, Alejandro; Eastwood, Rodney; Vogt, Daniel; Ithier, Carter; Smith, Michael; Wood, Rob; Kovač, Mirko

    2017-02-06

    Many insects are well adapted to long-distance migration despite the larger energetic costs of flight for small body sizes. To optimize wing design for next-generation flying micro-robots, we analyse butterfly wing shapes and wing orientations at full scale using numerical simulations and in a low-speed wind tunnel at 2, 3.5 and 5 m s -1 . The results indicate that wing orientations which maximize wing span lead to the highest glide performance, with lift to drag ratios up to 6.28, while spreading the fore-wings forward can increase the maximum lift produced and thus improve versatility. We discuss the implications for flying micro-robots and how the results assist in understanding the behaviour of the butterfly species tested.

  2. Aerodynamic evaluation of wing shape and wing orientation in four butterfly species using numerical simulations and a low-speed wind tunnel, and its implications for the design of flying micro-robots

    PubMed Central

    Eastwood, Rodney; Vogt, Daniel; Ithier, Carter; Smith, Michael; Wood, Rob; Kovač, Mirko

    2017-01-01

    Many insects are well adapted to long-distance migration despite the larger energetic costs of flight for small body sizes. To optimize wing design for next-generation flying micro-robots, we analyse butterfly wing shapes and wing orientations at full scale using numerical simulations and in a low-speed wind tunnel at 2, 3.5 and 5 m s−1. The results indicate that wing orientations which maximize wing span lead to the highest glide performance, with lift to drag ratios up to 6.28, while spreading the fore-wings forward can increase the maximum lift produced and thus improve versatility. We discuss the implications for flying micro-robots and how the results assist in understanding the behaviour of the butterfly species tested. PMID:28163879

  3. Preserving and vouchering butterflies and moths for large-scale museum-based molecular research

    PubMed Central

    Epstein, Samantha W.; Mitter, Kim; Hamilton, Chris A.; Plotkin, David; Mitter, Charles

    2016-01-01

    Butterflies and moths (Lepidoptera) comprise significant portions of the world’s natural history collections, but a standardized tissue preservation protocol for molecular research is largely lacking. Lepidoptera have traditionally been spread on mounting boards to display wing patterns and colors, which are often important for species identification. Many molecular phylogenetic studies have used legs from pinned specimens as the primary source for DNA in order to preserve a morphological voucher, but the amount of available tissue is often limited. Preserving an entire specimen in a cryogenic freezer is ideal for DNA preservation, but without an easily accessible voucher it can make specimen identification, verification, and morphological work difficult. Here we present a procedure that creates accessible and easily visualized “wing vouchers” of individual Lepidoptera specimens, and preserves the remainder of the insect in a cryogenic freezer for molecular research. Wings are preserved in protective holders so that both dorsal and ventral patterns and colors can be easily viewed without further damage. Our wing vouchering system has been implemented at the University of Maryland (AToL Lep Collection) and the University of Florida (Florida Museum of Natural History, McGuire Center of Lepidoptera and Biodiversity), which are among two of the largest Lepidoptera molecular collections in the world. PMID:27366654

  4. Adaptive Introgression across Species Boundaries in Heliconius Butterflies

    PubMed Central

    Pardo-Diaz, Carolina; Salazar, Camilo; Baxter, Simon W.; Merot, Claire; Figueiredo-Ready, Wilsea; Joron, Mathieu; McMillan, W. Owen; Jiggins, Chris D.

    2012-01-01

    It is widely documented that hybridisation occurs between many closely related species, but the importance of introgression in adaptive evolution remains unclear, especially in animals. Here, we have examined the role of introgressive hybridisation in transferring adaptations between mimetic Heliconius butterflies, taking advantage of the recent identification of a gene regulating red wing patterns in this genus. By sequencing regions both linked and unlinked to the red colour locus, we found a region that displays an almost perfect genotype by phenotype association across four species, H. melpomene, H. cydno, H. timareta, and H. heurippa. This particular segment is located 70 kb downstream of the red colour specification gene optix, and coalescent analysis indicates repeated introgression of adaptive alleles from H. melpomene into the H. cydno species clade. Our analytical methods complement recent genome scale data for the same region and suggest adaptive introgression has a crucial role in generating adaptive wing colour diversity in this group of butterflies. PMID:22737081

  5. UV photoreceptors and UV-yellow wing pigments in Heliconius butterflies allow a color signal to serve both mimicry and intraspecific communication.

    PubMed

    Bybee, Seth M; Yuan, Furong; Ramstetter, Monica D; Llorente-Bousquets, Jorge; Reed, Robert D; Osorio, Daniel; Briscoe, Adriana D

    2012-01-01

    Mimetic wing coloration evolves in butterflies in the context of predator confusion. Unless butterfly eyes have adaptations for discriminating mimetic color variation, mimicry also carries a risk of confusion for the butterflies themselves. Heliconius butterfly eyes, which express recently duplicated ultraviolet (UV) opsins, have such an adaptation. To examine bird and butterfly color vision as sources of selection on butterfly coloration, we studied yellow wing pigmentation in the tribe Heliconiini. We confirmed, using reflectance and mass spectrometry, that only Heliconius use 3-hydroxy-DL-kynurenine (3-OHK), which looks yellow to humans but reflects both UV- and long-wavelength light, whereas butterflies in related genera have chemically unknown yellow pigments mostly lacking UV reflectance. Modeling of these color signals reveals that the two UV photoreceptors of Heliconius are better suited to separating 3-OHK from non-3-OHK spectra compared with the photoreceptors of related genera or birds. The co-occurrence of potentially enhanced UV vision and a UV-reflecting yellow wing pigment could allow unpalatable Heliconius private intraspecific communication in the presence of mimics. Our results are the best available evidence for the correlated evolution of a color signal and color vision. They also suggest that predator visual systems are error prone in the context of mimicry. © 2011 by The University of Chicago.

  6. Localization of Defensive Chemicals in Two Congeneric Butterflies (Euphydryas, Nymphalidae).

    PubMed

    Mason, Peri A; Deane Bowers, M

    2017-05-01

    Many insect species sequester compounds acquired from their host plants for defense against natural enemies. The distribution of these compounds is likely to affect both their efficacy as defenses, and their costs. In this study we examined the distribution of sequestered iridoid glycosides (IGs) in two congeneric species of nymphalid butterfly, Euphydryas anicia and E. phaeton, and found that the pattern of localization of IGs differed between the two species. Although IG concentrations were quite high in the heads of both species, the relative concentrations in wings and abdomens differed substantially. Euphydryas anicia had relatively high IG concentrations in their abdomens and low IG concentrations in their wings, whereas the reverse was true in E. phaeton. We interpret these results in light of two current hypotheses regarding where sequestered chemicals should be localized: that they should be found in wings, which would allow non-lethal sampling by predators; and that their distribution is constrained by the distribution of tissue types to which sequestered compounds bind. We also offer the third hypothesis, that costs of storage may differ among body parts, and that the localization of compounds may reflect a cost-reduction strategy. Results from E. phaeton were consistent with all three of these non-mutually exclusive hypotheses, whereas results from E. anicia were only consistent with the notion that tissue bias among body parts plays a role in IG distribution. The finding that these two congeneric butterflies exhibit different patterns of IG localization suggests that they have been shaped by different selection regimes.

  7. Coupling effects in 3D plasmonic structures templated by Morpho butterfly wings.

    PubMed

    He, Jiaqing; Shen, Qingchen; Yang, Shuai; He, Gufeng; Tao, Peng; Song, Chengyi; Wu, Jianbo; Deng, Tao; Shang, Wen

    2018-01-03

    This paper presents the study of the coupling effects of three dimensional (3D) plasmonic nanostructures templated by Morpho butterfly wings. Different from the random deposition of metallic nanoparticles (NPs) or conformal coating of metallic layers on butterfly wings reported previously, the 3D plasmonic nanostructures studied in this work consist of gold (Au) nanostrips quasi-periodically arranged in 3D, which allows us to investigate the plasmonic coupling effects. Through refractive index (RI) matching, the plasmonic coupling can be differentiated from the optical contribution of butterfly wings. By tuning the deposition thickness of Au from 30 to 90 nm, the plasmonic coupling effects between the 3D Au nanostrips are gradually enhanced. In particular, the near-field coupling results in two resonant modes and enhances the surface-enhanced Raman scattering (SERS) signals.

  8. An Ingenious Super Light Trapping Surface Templated from Butterfly Wing Scales

    NASA Astrophysics Data System (ADS)

    Han, Zhiwu; Li, Bo; Mu, Zhengzhi; Yang, Meng; Niu, Shichao; Zhang, Junqiu; Ren, Luquan

    2015-08-01

    Based on the super light trapping property of butterfly Trogonoptera brookiana wings, the SiO2 replica of this bionic functional surface was successfully synthesized using a simple and highly effective synthesis method combining a sol-gel process and subsequent selective etching. Firstly, the reflectivity of butterfly wing scales was carefully examined. It was found that the whole reflectance spectroscopy of the butterfly wings showed a lower level (less than 10 %) in the visible spectrum. Thus, it was confirmed that the butterfly wings possessed a super light trapping effect. Afterwards, the morphologies and detailed architectures of the butterfly wing scales were carefully investigated using the ultra-depth three-dimensional (3D) microscope and field emission scanning electronic microscopy (FESEM). It was composed by the parallel ridges and quasi-honeycomb-like structure between them. Based on the biological properties and function above, an exact SiO2 negative replica was fabricated through a synthesis method combining a sol-gel process and subsequent selective etching. At last, the comparative analysis of morphology feature size and the reflectance spectroscopy between the SiO2 negative replica and the flat plate was conducted. It could be concluded that the SiO2 negative replica inherited not only the original super light trapping architectures, but also the super light trapping characteristics of bio-template. This work may open up an avenue for the design and fabrication of super light trapping materials and encourage people to look for more super light trapping architectures in nature.

  9. Characterization of Structural and Pigmentary Colors in Common Emigrant (Catopsilia Pomona) Butterfly

    DOE Office of Scientific and Technical Information (OSTI.GOV)

    Ghate, Ekata; Kulkarni, G. R.; Bhoraskar, S. V.

    2011-10-20

    Study of structural colors in case of insects and butterflies is important for their biomimic and biophotonics applications. Structural color is the color which is produced by physical structures and their interaction with light while pigmentary color is produced by absorption of light by pigments. Common Emigrant butterfly is widely distributed in India. It is of moderate size with wing span of about 60-80 mm. The wings are broadly white with yellow or sulphur yellow coloration at places as well as few dark black patches. It belongs to family Pieridae. A study of structural color in case of Common Emigrantmore » butterfly has been carried out in the present work. The characterization of wing color was performed using absorption spectroscopy. Scanning electron microscopic study of the wings of Common Emigrant butterfly showed that three different types of scales are present on the wing surface dorsally. Diffracting structures are present in certain parts of the surfaces of the various scales. Bead like structures are embedded in the intricate structures of the scales. Absorption spectra revealed that a strong absorption peak is seen in the UV-range. Crystalline structure of beads was confirmed by the X-ray diffraction analysis.« less

  10. White butterflies as solar photovoltaic concentrators

    NASA Astrophysics Data System (ADS)

    Shanks, Katie; Senthilarasu, S.; Ffrench-Constant, Richard H.; Mallick, Tapas K.

    2015-07-01

    Man’s harvesting of photovoltaic energy requires the deployment of extensive arrays of solar panels. To improve both the gathering of thermal and photovoltaic energy from the sun we have examined the concept of biomimicry in white butterflies of the family Pieridae. We tested the hypothesis that the V-shaped posture of basking white butterflies mimics the V-trough concentrator which is designed to increase solar input to photovoltaic cells. These solar concentrators improve harvesting efficiency but are both heavy and bulky, severely limiting their deployment. Here, we show that the attachment of butterfly wings to a solar cell increases its output power by 42.3%, proving that the wings are indeed highly reflective. Importantly, and relative to current concentrators, the wings improve the power to weight ratio of the overall structure 17-fold, vastly expanding their potential application. Moreover, a single mono-layer of scale cells removed from the butterflies’ wings maintained this high reflectivity showing that a single layer of scale cell-like structures can also form a useful coating. As predicted, the wings increased the temperature of the butterflies’ thorax dramatically, showing that the V-shaped basking posture of white butterflies has indeed evolved to increase the temperature of their flight muscles prior to take-off.

  11. Nanofabrication and coloration study of artificial Morpho butterfly wings with aligned lamellae layers

    PubMed Central

    Zhang, Sichao; Chen, Yifang

    2015-01-01

    The bright and iridescent blue color from Morpho butterfly wings has attracted worldwide attentions to explore its mysterious nature for long time. Although the physics of structural color by the nanophotonic structures built on the wing scales has been well established, replications of the wing structure by standard top-down lithography still remains a challenge. This paper reports a technical breakthrough to mimic the blue color of Morpho butterfly wings, by developing a novel nanofabrication process, based on electron beam lithography combined with alternate PMMA/LOR development/dissolution, for photonic structures with aligned lamellae multilayers in colorless polymers. The relationship between the coloration and geometric dimensions as well as shapes is systematically analyzed by solving Maxwell’s Equations with a finite domain time difference simulator. Careful characterization of the mimicked blue by spectral measurements under both normal and oblique angles are carried out. Structural color in blue reflected by the fabricated wing scales, is demonstrated and further extended to green as an application exercise of the new technique. The effects of the regularity in the replicas on coloration are analyzed. In principle, this approach establishes a starting point for mimicking structural colors beyond the blue in Morpho butterfly wings. PMID:26577813

  12. Nanofabrication and coloration study of artificial Morpho butterfly wings with aligned lamellae layers.

    PubMed

    Zhang, Sichao; Chen, Yifang

    2015-11-18

    The bright and iridescent blue color from Morpho butterfly wings has attracted worldwide attentions to explore its mysterious nature for long time. Although the physics of structural color by the nanophotonic structures built on the wing scales has been well established, replications of the wing structure by standard top-down lithography still remains a challenge. This paper reports a technical breakthrough to mimic the blue color of Morpho butterfly wings, by developing a novel nanofabrication process, based on electron beam lithography combined with alternate PMMA/LOR development/dissolution, for photonic structures with aligned lamellae multilayers in colorless polymers. The relationship between the coloration and geometric dimensions as well as shapes is systematically analyzed by solving Maxwell's Equations with a finite domain time difference simulator. Careful characterization of the mimicked blue by spectral measurements under both normal and oblique angles are carried out. Structural color in blue reflected by the fabricated wing scales, is demonstrated and further extended to green as an application exercise of the new technique. The effects of the regularity in the replicas on coloration are analyzed. In principle, this approach establishes a starting point for mimicking structural colors beyond the blue in Morpho butterfly wings.

  13. Nanofabrication and coloration study of artificial Morpho butterfly wings with aligned lamellae layers

    NASA Astrophysics Data System (ADS)

    Zhang, Sichao; Chen, Yifang

    2015-11-01

    The bright and iridescent blue color from Morpho butterfly wings has attracted worldwide attentions to explore its mysterious nature for long time. Although the physics of structural color by the nanophotonic structures built on the wing scales has been well established, replications of the wing structure by standard top-down lithography still remains a challenge. This paper reports a technical breakthrough to mimic the blue color of Morpho butterfly wings, by developing a novel nanofabrication process, based on electron beam lithography combined with alternate PMMA/LOR development/dissolution, for photonic structures with aligned lamellae multilayers in colorless polymers. The relationship between the coloration and geometric dimensions as well as shapes is systematically analyzed by solving Maxwell’s Equations with a finite domain time difference simulator. Careful characterization of the mimicked blue by spectral measurements under both normal and oblique angles are carried out. Structural color in blue reflected by the fabricated wing scales, is demonstrated and further extended to green as an application exercise of the new technique. The effects of the regularity in the replicas on coloration are analyzed. In principle, this approach establishes a starting point for mimicking structural colors beyond the blue in Morpho butterfly wings.

  14. Active dendrites: colorful wings of the mysterious butterflies.

    PubMed

    Johnston, Daniel; Narayanan, Rishikesh

    2008-06-01

    Santiago Ramón y Cajal had referred to neurons as the 'mysterious butterflies of the soul.' Wings of these butterflies--their dendrites--were traditionally considered as passive integrators of synaptic information. Owing to a growing body of experimental evidence, it is now widely accepted that these wings are colorful, endowed with a plethora of active conductances, with each family of these butterflies made of distinct hues and shades. Furthermore, rapidly evolving recent literature also provides direct and indirect demonstrations for activity-dependent plasticity of these active conductances, pointing toward chameleonic adaptability in these hues. These experimental findings firmly establish the immense computational power of a single neuron, and thus constitute a turning point toward the understanding of various aspects of neuronal information processing. In this brief historical perspective, we track important milestones in the chameleonic transmogrification of these mysterious butterflies.

  15. Disruptive ecological selection on a mating cue.

    PubMed

    Merrill, Richard M; Wallbank, Richard W R; Bull, Vanessa; Salazar, Patricio C A; Mallet, James; Stevens, Martin; Jiggins, Chris D

    2012-12-22

    Adaptation to divergent ecological niches can result in speciation. Traits subject to disruptive selection that also contribute to non-random mating will facilitate speciation with gene flow. Such 'magic' or 'multiple-effect' traits may be widespread and important for generating biodiversity, but strong empirical evidence is still lacking. Although there is evidence that putative ecological traits are indeed involved in assortative mating, evidence that these same traits are under divergent selection is considerably weaker. Heliconius butterfly wing patterns are subject to positive frequency-dependent selection by predators, owing to aposematism and Müllerian mimicry, and divergent colour patterns are used by closely related species to recognize potential mates. The amenability of colour patterns to experimental manipulation, independent of other traits, presents an excellent opportunity to test their role during speciation. We conducted field experiments with artificial butterflies, designed to match natural butterflies with respect to avian vision. These were complemented with enclosure trials with live birds and real butterflies. Our experiments showed that hybrid colour-pattern phenotypes are attacked more frequently than parental forms. For the first time, we demonstrate disruptive ecological selection on a trait that also acts as a mating cue.

  16. Inferring the Provenance of an Alien Species with DNA Barcodes: The Neotropical Butterfly Dryas iulia in Thailand

    PubMed Central

    Burg, Noah A.; Pradhan, Ashman; Gonzalez, Rebecca M.; Morban, Emely Z.; Zhen, Erica W.; Sakchoowong, Watana; Lohman, David J.

    2014-01-01

    The Neotropical butterfly Dryas iulia has been collected from several locations in Thailand and Malaysia since 2007, and has been observed breeding in the wild, using introduced Passiflora foetida as a larval host plant. The butterfly is bred by a butterfly house in Phuket, Thailand, for release at weddings and Buddhist ceremonies, and we hypothesized that this butterfly house was the source of wild, Thai individuals. We compared wing patterns and COI barcodes from two, wild Thai populations with individuals obtained from this butterfly house. All Thai individuals resemble the subspecies D. iulia modesta, and barcodes from wild and captive Thai specimens were identical. This unique, Thai barcode was not found in any of the 30 specimens sampled from the wild in the species' native range, but is most similar to specimens from Costa Rica, where many exporting butterfly farms are located. These data implicate the butterfly house as the source of Thailand's wild D. iulia populations, which are currently so widespread that eradication efforts are unlikely to be successful. PMID:25119899

  17. Inferring the provenance of an alien species with DNA barcodes: the neotropical butterfly Dryas iulia in Thailand.

    PubMed

    Burg, Noah A; Pradhan, Ashman; Gonzalez, Rebecca M; Morban, Emely Z; Zhen, Erica W; Sakchoowong, Watana; Lohman, David J

    2014-01-01

    The Neotropical butterfly Dryas iulia has been collected from several locations in Thailand and Malaysia since 2007, and has been observed breeding in the wild, using introduced Passiflora foetida as a larval host plant. The butterfly is bred by a butterfly house in Phuket, Thailand, for release at weddings and Buddhist ceremonies, and we hypothesized that this butterfly house was the source of wild, Thai individuals. We compared wing patterns and COI barcodes from two, wild Thai populations with individuals obtained from this butterfly house. All Thai individuals resemble the subspecies D. iulia modesta, and barcodes from wild and captive Thai specimens were identical. This unique, Thai barcode was not found in any of the 30 specimens sampled from the wild in the species' native range, but is most similar to specimens from Costa Rica, where many exporting butterfly farms are located. These data implicate the butterfly house as the source of Thailand's wild D. iulia populations, which are currently so widespread that eradication efforts are unlikely to be successful.

  18. Genomic architecture of adaptive color pattern divergence and convergence in Heliconius butterflies

    PubMed Central

    Supple, Megan A.; Hines, Heather M.; Dasmahapatra, Kanchon K.; Lewis, James J.; Nielsen, Dahlia M.; Lavoie, Christine; Ray, David A.; Salazar, Camilo; McMillan, W. Owen; Counterman, Brian A.

    2013-01-01

    Identifying the genetic changes driving adaptive variation in natural populations is key to understanding the origins of biodiversity. The mosaic of mimetic wing patterns in Heliconius butterflies makes an excellent system for exploring adaptive variation using next-generation sequencing. In this study, we use a combination of techniques to annotate the genomic interval modulating red color pattern variation, identify a narrow region responsible for adaptive divergence and convergence in Heliconius wing color patterns, and explore the evolutionary history of these adaptive alleles. We use whole genome resequencing from four hybrid zones between divergent color pattern races of Heliconius erato and two hybrid zones of the co-mimic Heliconius melpomene to examine genetic variation across 2.2 Mb of a partial reference sequence. In the intergenic region near optix, the gene previously shown to be responsible for the complex red pattern variation in Heliconius, population genetic analyses identify a shared 65-kb region of divergence that includes several sites perfectly associated with phenotype within each species. This region likely contains multiple cis-regulatory elements that control discrete expression domains of optix. The parallel signatures of genetic differentiation in H. erato and H. melpomene support a shared genetic architecture between the two distantly related co-mimics; however, phylogenetic analysis suggests mimetic patterns in each species evolved independently. Using a combination of next-generation sequencing analyses, we have refined our understanding of the genetic architecture of wing pattern variation in Heliconius and gained important insights into the evolution of novel adaptive phenotypes in natural populations. PMID:23674305

  19. Argonne News Brief: Secret of Butterfly Wings Revealed With X-Rays

    DOE Office of Scientific and Technical Information (OSTI.GOV)

    None

    Scientists used powerful X-rays at the Advanced Photon Source at Argonne to get a unique look at the structure of the tiny crystals that make up the wings of butterflies. The results show us how the wings get their iridescent, brilliant colors—and could give us ideas for technology of our own.

  20. Prey survival by predator intimidation: an experimental study of peacock butterfly defence against blue tits

    PubMed Central

    Vallin, Adrian; Jakobsson, Sven; Lind, Johan; Wiklund, Christer

    2005-01-01

    Long-lived butterflies that hibernate as adults are expected to have well-developed antipredation devices as a result of their long exposure to natural enemies. The peacock butterfly, Inachis io, for instance, is a cryptic leaf mimic when resting, but shifts to active defence when disturbed, performing a repeated sequence of movements exposing major eyespots on the wings accompanied by a hissing noise. We studied the effect of visual and auditory defence by staging experiments in which wild-caught blue tits, Parus caeruleus, were presented with one of six kinds of experimentally manipulated living peacock butterflies as follows: butterflies with eyespots painted over and their controls (painted on another part of the wing), butterflies with their sound production aborted (small part of wings removed) and their controls, and butterflies with eyespots painted over and sound production aborted and their controls. The results showed that eyespots alone, or in combination with sound, constituted an effective defence; only 1 out of 34 butterflies with intact eyespots was killed, whereas 13 out of 20 butterflies without eyespots were killed. The killed peacocks were eaten, indicating that they are not distasteful. Hence, intimidation by bluffing can be an efficient means of defence for an edible prey. PMID:16024383

  1. Study of structural colour of Hebomoia glaucippe butterfly wing scales

    NASA Astrophysics Data System (ADS)

    Shur, V. Ya; Kuznetsov, D. K.; Pryakhina, V. I.; Kosobokov, M. S.; Zubarev, I. V.; Boymuradova, S. K.; Volchetskaya, K. V.

    2017-10-01

    Structural colours of Hebomoia glaucippe butterfly wing scales have been studied experimentally using high resolution scanning electron microscopy. Visualization of scales structures and computer simulation allowed distinguishing correlation between nanostructures on the scales and their colour.

  2. Detailed electromagnetic simulation for the structural color of butterfly wings.

    PubMed

    Lee, R Todd; Smith, Glenn S

    2009-07-20

    Many species of butterflies exhibit interesting optical phenomena due to structural color. The physical reason for this color is subwavelength features on the surface of a single scale. The exposed surface of a scale is covered with a ridge structure. The fully three-dimensional, periodic, finite-difference time-domain method is used to create a detailed electromagnetic model of a generic ridge. A novel method for presenting the three-dimensional observed color pattern is developed. Using these tools, the change in color that is a result of varying individual features of the scale is explored. Computational models are developed that are similar to three butterflies: Morpho rhetenor, Troides magellanus, and Ancyluris meliboeus.

  3. Wound healing, calcium signaling, and other novel pathways are associated with the formation of butterfly eyespots.

    PubMed

    Özsu, Nesibe; Monteiro, Antónia

    2017-10-16

    One hypothesis surrounding the origin of novel traits is that they originate from the co-option of pre-existing genes or larger gene regulatory networks into novel developmental contexts. Insights into a trait's evolutionary origins can, thus, be gained via identification of the genes underlying trait development, and exploring whether those genes also function in other developmental contexts. Here we investigate the set of genes associated with the development of eyespot color patterns, a trait that originated once within the Nymphalid family of butterflies. Although several genes associated with eyespot development have been identified, the eyespot gene regulatory network remains largely unknown. In this study, next-generation sequencing and transcriptome analyses were used to identify a large set of genes associated with eyespot development of Bicyclus anynana butterflies, at 3-6 h after pupation, prior to the differentiation of the color rings. Eyespot-associated genes were identified by comparing the transcriptomes of homologous micro-dissected wing tissues that either develop or do not develop eyespots in wild-type and a mutant line of butterflies, Spotty, with extra eyespots. Overall, 186 genes were significantly up and down-regulated in wing tissues that develop eyespots compared to wing tissues that do not. Many of the differentially expressed genes have yet to be annotated. New signaling pathways, including the Toll, Fibroblast Growth Factor (FGF), extracellular signal-regulated kinase (ERK) and/or Jun N-terminal kinase (JNK) signaling pathways are associated for the first time with eyespot development. In addition, several genes involved in wound healing and calcium signaling were also found to be associated with eyespots. Overall, this study provides the identity of many new genes and signaling pathways associated with eyespots, and suggests that the ancient wound healing gene regulatory network may have been co-opted to cells at the center of the pattern to aid in eyespot origins. New transcription factors that may be providing different identities to distinct wing sectors, and genes with sexually dimorphic expression in the eyespots were also identified.

  4. The development of wing shape in Lepidoptera: mitotic density, not orientation, is the primary determinant of shape.

    PubMed

    Nijhout, H Frederik; Cinderella, Margaret; Grunert, Laura W

    2014-03-01

    The wings of butterflies and moths develop from imaginal disks whose structure is always congruent with the final adult wing. It is therefore possible to map every point on the imaginal disk to a location on the adult wing throughout ontogeny. We studied the growth patterns of the wings of two distantly related species with very different adult wing shapes, Junonia coenia and Manduca sexta. The shape of the wing disks change throughout their growth phase in a species-specific pattern. We measured mitotic densities and mitotic orientation in successive stages of wing development approximately one cell division apart. Cell proliferation was spatially patterned, and the density of mitoses was highly correlated with local growth. Unlike other systems in which the direction of mitoses has been viewed as the primary determinant of directional growth, we found that in these two species the direction of growth was only weakly correlated with the orientation of mitoses. Directional growth appears to be imposed by a constantly changing spatial pattern of cell division coupled with a weak bias in the orientation of cell division. Because growth and cell division in imaginal disk require ecdysone and insulin signaling, the changing spatial pattern of cell division may due to a changing pattern of expression of receptors or downstream elements in the signaling pathways for one or both of these hormones. Evolution of wing shape comes about by changes in the progression of spatial patterns of cell division. © 2014 Wiley Periodicals, Inc.

  5. Nano

    Science.gov Websites

    inspire light-manipulating surface for medical implants Butterfly wings inspire light-manipulating surface for medical implants Inspired by tiny nanostructures on transparent butterfly wings, engineers at -manipulating surface for medical implants Our phones could soon be sending data by light Scalable manufacturing

  6. Colour formation on the wings of the butterfly Hypolimnas salmacis by scale stacking

    NASA Astrophysics Data System (ADS)

    Siddique, Radwanul Hasan; Vignolini, Silvia; Bartels, Carolin; Wacker, Irene; Hölscher, Hendrik

    2016-11-01

    The butterfly genus Hypolimnas features iridescent blue colouration in some areas of its dorsal wings. Here, we analyse the mechanisms responsible for such colouration on the dorsal wings of Hypolimnas salmacis and experimentally demonstrate that the lower thin lamina in the white cover scales causes the blue iridescence. This outcome contradicts other studies reporting that the radiant blue in Hypolimnas butterflies is caused by complex ridge-lamellar architectures in the upper lamina of the cover scales. Our comprehensive optical study supported by numerical calculation however shows that scale stacking primarily induces the observed colour appearance of Hypolimnas salmacis.

  7. Colour formation on the wings of the butterfly Hypolimnas salmacis by scale stacking.

    PubMed

    Siddique, Radwanul Hasan; Vignolini, Silvia; Bartels, Carolin; Wacker, Irene; Hölscher, Hendrik

    2016-11-02

    The butterfly genus Hypolimnas features iridescent blue colouration in some areas of its dorsal wings. Here, we analyse the mechanisms responsible for such colouration on the dorsal wings of Hypolimnas salmacis and experimentally demonstrate that the lower thin lamina in the white cover scales causes the blue iridescence. This outcome contradicts other studies reporting that the radiant blue in Hypolimnas butterflies is caused by complex ridge-lamellar architectures in the upper lamina of the cover scales. Our comprehensive optical study supported by numerical calculation however shows that scale stacking primarily induces the observed colour appearance of Hypolimnas salmacis.

  8. White butterflies as solar photovoltaic concentrators

    PubMed Central

    Shanks, Katie; Senthilarasu, S.; ffrench-Constant, Richard H.; Mallick, Tapas K.

    2015-01-01

    Man’s harvesting of photovoltaic energy requires the deployment of extensive arrays of solar panels. To improve both the gathering of thermal and photovoltaic energy from the sun we have examined the concept of biomimicry in white butterflies of the family Pieridae. We tested the hypothesis that the V-shaped posture of basking white butterflies mimics the V-trough concentrator which is designed to increase solar input to photovoltaic cells. These solar concentrators improve harvesting efficiency but are both heavy and bulky, severely limiting their deployment. Here, we show that the attachment of butterfly wings to a solar cell increases its output power by 42.3%, proving that the wings are indeed highly reflective. Importantly, and relative to current concentrators, the wings improve the power to weight ratio of the overall structure 17-fold, vastly expanding their potential application. Moreover, a single mono-layer of scale cells removed from the butterflies’ wings maintained this high reflectivity showing that a single layer of scale cell-like structures can also form a useful coating. As predicted, the wings increased the temperature of the butterflies’ thorax dramatically, showing that the V-shaped basking posture of white butterflies has indeed evolved to increase the temperature of their flight muscles prior to take-off. PMID:26227341

  9. Infrared detection based on localized modification of Morpho butterfly wings.

    PubMed

    Zhang, Fangyu; Shen, Qingchen; Shi, Xindong; Li, Shipu; Wang, Wanlin; Luo, Zhen; He, Gufeng; Zhang, Peng; Tao, Peng; Song, Chengyi; Zhang, Wang; Zhang, Di; Deng, Tao; Shang, Wen

    2015-02-01

    Inspired by butterflies an advanced detection and sensing system is developed. The hierarchical nanoarchitecture of Morpho butterfly wings is shown to facilitate the selective modification of such a structure, which results in a sensitive infrared response. These findings offer a new path both for detecting infrared photons and for generating nanostructured bimaterial systems for high-performance sensing platforms. © 2014 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim.

  10. X-ray Tomography and Chemical Imaging within Butterfly Wing Scales

    DOE Office of Scientific and Technical Information (OSTI.GOV)

    Chen Jianhua; Lee Yaochang; Tang, M.-T.

    2007-01-19

    The rainbow like color of butterfly wings is associated with the internal and surface structures of the wing scales. While the photonic structure of the scales is believed to diffract specific lights at different angle, there is no adequate probe directly answering the 3-D structures with sufficient spatial resolution. The NSRRC nano-transmission x-ray microscope (nTXM) with tens nanometers spatial resolution is able to image biological specimens without artifacts usually introduced in sophisticated sample staining processes. With the intrinsic deep penetration of x-rays, the nTXM is capable of nondestructively investigating the internal structures of fragile and soft samples. In this study,more » we imaged the structure of butterfly wing scales in 3-D view with 60 nm spatial resolution. In addition, synchrotron-radiation-based Fourier transform Infrared (FT-IR) microspectroscopy was employed to analyze the chemical components with spatial information of the butterfly wing scales. Based on the infrared spectral images, we suggest that the major components of scale structure were rich in protein and polysaccharide.« less

  11. Species richness and trait composition of butterfly assemblages change along an altitudinal gradient.

    PubMed

    Leingärtner, Annette; Krauss, Jochen; Steffan-Dewenter, Ingolf

    2014-06-01

    Species richness patterns along altitudinal gradients are well-documented ecological phenomena, yet very little data are available on how environmental filtering processes influence the composition and traits of butterfly assemblages at high altitudes. We have studied the diversity patterns of butterfly species at 34 sites along an altitudinal gradient ranging from 600 to 2,000 m a.s.l. in the National Park Berchtesgaden (Germany) and analysed traits of butterfly assemblages associated with dispersal capacity, reproductive strategies and developmental time from lowlands to highlands, including phylogenetic analyses. We found a linear decline in butterfly species richness along the altitudinal gradient, but the phylogenetic relatedness of the butterfly assemblages did not increase with altitude. Compared to butterfly assemblages at lower altitudes, those at higher altitudes were composed of species with larger wings (on average 9%) which laid an average of 68% more eggs. In contrast, egg maturation time in butterfly assemblages decreased by about 22% along the altitudinal gradient. Further, butterfly assemblages at higher altitudes were increasingly dominated by less widespread species. Based on our abundance data, but not on data in the literature, population density increased with altitude, suggesting a reversed density-distribution relationship, with higher population densities of habitat specialists in harsh environments. In conclusion, our data provide evidence for significant shifts in the composition of butterfly assemblages and for the dominance of different traits along the altitudinal gradient. In our study, these changes were mainly driven by environmental factors, whereas phylogenetic filtering played a minor role along the studied altitudinal range.

  12. Physiological Perturbation Reveals Modularity of Eyespot Development in the Painted Lady Butterfly, Vanessa cardui

    PubMed Central

    Rhen, Turk; Simmons, Rebecca B.

    2016-01-01

    Butterfly eyespots are complex morphological traits that can vary in size, shape and color composition even on the same wing surface. Homology among eyespots suggests they share a common developmental basis and function as an integrated unit in response to selection. Despite strong evidence of genetic integration, eyespots can also exhibit modularity or plasticity, indicating an underlying flexibility in pattern development. The extent to which particular eyespots or eyespot color elements exhibit modularity or integration is poorly understood, particularly following exposure to novel conditions. We used perturbation experiments to explore phenotypic correlations among different eyespots and their color elements on the ventral hindwing of V. cardui. Specifically, we identified which eyespots and eyespot features are most sensitive to perturbation by heat shock and injection of heparin—a cold shock mimic. For both treatments, the two central eyespots (3 + 4) were most affected by the experimental perturbations, whereas the outer eyespot border was more resistant to modification than the interior color elements. Overall, the individual color elements displayed a similar response to heat shock across all eyespots, but varied in their response to each other. Graphical modeling also revealed that although eyespots differ morphologically, regulation of eyespot size and colored elements appear to be largely integrated across the wing. Patterns of integration, however, were disrupted following heat shock, revealing that the strength of integration varies across the wing and is strongest between the two central eyespots. These findings support previous observations that document coupling between eyespots 3 + 4 in other nymphalid butterflies. PMID:27560365

  13. Physiological Perturbation Reveals Modularity of Eyespot Development in the Painted Lady Butterfly, Vanessa cardui.

    PubMed

    Connahs, Heidi; Rhen, Turk; Simmons, Rebecca B

    2016-01-01

    Butterfly eyespots are complex morphological traits that can vary in size, shape and color composition even on the same wing surface. Homology among eyespots suggests they share a common developmental basis and function as an integrated unit in response to selection. Despite strong evidence of genetic integration, eyespots can also exhibit modularity or plasticity, indicating an underlying flexibility in pattern development. The extent to which particular eyespots or eyespot color elements exhibit modularity or integration is poorly understood, particularly following exposure to novel conditions. We used perturbation experiments to explore phenotypic correlations among different eyespots and their color elements on the ventral hindwing of V. cardui. Specifically, we identified which eyespots and eyespot features are most sensitive to perturbation by heat shock and injection of heparin-a cold shock mimic. For both treatments, the two central eyespots (3 + 4) were most affected by the experimental perturbations, whereas the outer eyespot border was more resistant to modification than the interior color elements. Overall, the individual color elements displayed a similar response to heat shock across all eyespots, but varied in their response to each other. Graphical modeling also revealed that although eyespots differ morphologically, regulation of eyespot size and colored elements appear to be largely integrated across the wing. Patterns of integration, however, were disrupted following heat shock, revealing that the strength of integration varies across the wing and is strongest between the two central eyespots. These findings support previous observations that document coupling between eyespots 3 + 4 in other nymphalid butterflies.

  14. Spectral reflectance properties of iridescent pierid butterfly wings.

    PubMed

    Wilts, Bodo D; Pirih, Primož; Stavenga, Doekele G

    2011-06-01

    The wings of most pierid butterflies exhibit a main, pigmentary colouration: white, yellow or orange. The males of many species have in restricted areas of the wing upper sides a distinct structural colouration, which is created by stacks of lamellae in the ridges of the wing scales, resulting in iridescence. The amplitude of the reflectance is proportional to the number of lamellae in the ridge stacks. The angle-dependent peak wavelength of the observed iridescence is in agreement with classical multilayer theory. The iridescence is virtually always in the ultraviolet wavelength range, but some species have a blue-peaking iridescence. The spectral properties of the pigmentary and structural colourations are presumably tuned to the spectral sensitivities of the butterflies' photoreceptors.

  15. Photonic Crystals on the Wing

    DTIC Science & Technology

    2011-11-01

    Giraldo MA Stavenga DG (2007) Sexual dichroism and pigment localization in the wing scales of Pieris rapae butterflies. Proc R Soc B 274:97-102 4...723 41. Stavenga DG, Arikawa K (2011) Photoreceptor spectral sensitivities of the Small White butterfly Pieris rapae crucivora interpreted with

  16. The Effects of Scales on Autorotation of Monarch Butterfly Forewings

    NASA Astrophysics Data System (ADS)

    Dechello, Nicole; Lang, Amy

    2014-11-01

    The wings of Monarch butterflies (Danus plexippus) have scales of approximately 100 micrometers that cover their wings in a roof-shingle pattern, and these scales are hypothesized to help improve flight efficiency for their long migration. The aerodynamic effects of the scales, particularly involving the leading edge vortex formation and resulting lift, were investigated by observing the natural autorotation of forewing specimen when dropped in quiescent air. A high-speed camera recorded drop tests of 32 forewings both with scales and after removal of the scales. It was found that the scales, on average, comprised 17% of the forewing mass. Tracking software was used to analyze the videos for several parameters, including descent speed and radius of rotation. NSF ECE Grant #1358991 supported the first author as an research experience for undergraduate (REU) student.

  17. Structural colours of nickel bioreplicas of butterfly wings

    NASA Astrophysics Data System (ADS)

    Tolenis, Tomas; Swiontek, Stephen E.; Lakhtakia, Akhlesh

    2017-04-01

    The two-angle conformally evaporated-film-by-rotation technique (TA-CEFR) was devised to coat the wings of the monarch butterfly with nickel in order to form a 500-nm thick bioreplica thereof. The bioreplica exhibits structural colours that are completely obscured in actual wings by pigmental colours. Thus, the TA-CEFR technique provides a way to replicate, study and exploit hidden morphologies of biological surfaces.

  18. Butterfly inclusions in Van Schrieck masterpieces. Techniques and optical properties

    NASA Astrophysics Data System (ADS)

    Berthier, S.; Boulenguez, J.; Menu, M.; Mottin, B.

    2008-07-01

    Dutch painter Otto Marseus Van Schrieck (1619 1678) is famous for his invention of “sottobosco”. These specific still-life paintings are characterized by the presence of various living organisms (mainly insects and plants) directly on the canvas. We will focus our attention on the painting kept in the museum of Grenoble, France, where a real butterfly is pasted on the canvas. The actual butterfly is a common Nymphalidae, Inachis io, presented in a static position on the dorsal side, without any perspective, compared to the neighboring butterflies. The colors of this butterfly are mainly due to pigments, melanin (black to brown) and ommochromes (yellow, orange, red) often in granules configuration that introduce scattering of light superimposed to the classical selective absorption, except in the ocelli of the hind wings where the blue coloration is due to interferential effects. The nearly perfect refraction index equality between the varnish and the chitin, the main constituent of the butterfly wings, deeply affects its colors. This leads the artist to a final intervention in some parts of the wings, revealed by microscope observation.

  19. Flight testing of live Monarch butterflies to determine the aerodynamic benefit of butterfly scales

    NASA Astrophysics Data System (ADS)

    Lang, Amy; Cranford, Jacob; Conway, Jasmine; Slegers, Nathan; Dechello, Nicole; Wilroy, Jacob

    2014-11-01

    Evolutionary adaptations in the morphological structure of butterfly scales (0.1 mm in size) to develop a unique micro-patterning resulting in a surface drag alteration, stem from a probable aerodynamic benefit of minimizing the energy requirement to fly a very lightweight body with comparably large surface area in a low Re flow regime. Live Monarch butterflies were tested at UAHuntsville's Autonomous Tracking and Optical Measurement (ATOM) Laboratory, which uses 22 Vicon T40 cameras that allow for millimeter level tracking of reflective markers at 515 fps over a 4 m × 6 m × 7 m volume. Data recorded included the flight path as well as the wing flapping angle and wing-beat frequency. Insects were first tested with their scales intact, and then again with the scales carefully removed. Differences in flapping frequency and/or energy obtained during flight due to the removal of the scales will be discussed. Initial data analysis indicates that scale removal in some specimens leads to increased flapping frequencies for similar energetic flight or reduced flight speed for similar flapping frequencies. Both results point to the scales providing an aerodynamic benefit, which is hypothesized to be linked to leading-edge vortex formation and induced drag. Funding from the National Science Foundation (CBET and REU) is gratefully acknowledged.

  20. Biologically inspired flexible quasi-single-mode random laser: an integration of Pieris canidia butterfly wing and semiconductors.

    PubMed

    Wang, Cih-Su; Chang, Tsung-Yuan; Lin, Tai-Yuan; Chen, Yang-Fang

    2014-10-23

    Quasi-periodic structures of natural biomaterial membranes have great potentials to serve as resonance cavities to generate ecological friendly optoelectronic devices with low cost. To achieve the first attempt for the illustration of the underlying principle, the Pieris canidia butterfly wing was embedded with ZnO nanoparticles. Quite interestingly, it is found that the bio-inspired quasi-single-mode random laser can be achieved by the assistance of the skeleton of the membrane, in which ZnO nanoparticles act as emitting gain media. Such unique characteristics can be interpreted well by the Fabry-Perot resonance existing in the window-like quasi-periodic structure of butterfly wing. Due to the inherently promising flexibility of butterfly wing membrane, the laser action can still be maintained during the bending process. Our demonstrated approach not only indicates that the natural biological structures can provide effective scattering feedbacks but also pave a new avenue towards designing bio-controlled photonic devices.

  1. Biologically inspired flexible quasi-single-mode random laser: An integration of Pieris canidia butterfly wing and semiconductors

    PubMed Central

    Wang, Cih-Su; Chang, Tsung-Yuan; Lin, Tai-Yuan; Chen, Yang-Fang

    2014-01-01

    Quasi-periodic structures of natural biomaterial membranes have great potentials to serve as resonance cavities to generate ecological friendly optoelectronic devices with low cost. To achieve the first attempt for the illustration of the underlying principle, the Pieris canidia butterfly wing was embedded with ZnO nanoparticles. Quite interestingly, it is found that the bio-inspired quasi-single-mode random laser can be achieved by the assistance of the skeleton of the membrane, in which ZnO nanoparticles act as emitting gain media. Such unique characteristics can be interpreted well by the Fabry-Perot resonance existing in the window-like quasi-periodic structure of butterfly wing. Due to the inherently promising flexibility of butterfly wing membrane, the laser action can still be maintained during the bending process. Our demonstrated approach not only indicates that the natural biological structures can provide effective scattering feedbacks but also pave a new avenue towards designing bio-controlled photonic devices. PMID:25338507

  2. Fluid drag reduction and efficient self-cleaning with rice leaf and butterfly wing bioinspired surfaces

    NASA Astrophysics Data System (ADS)

    Bixler, Gregory D.; Bhushan, Bharat

    2013-08-01

    Researchers are continually inspired by living nature to solve complex challenges. For example, unique surface characteristics of rice leaves and butterfly wings combine the shark skin (anisotropic flow leading to low drag) and lotus leaf (superhydrophobic and self-cleaning) effects, producing the so-called rice and butterfly wing effect. In this paper, we present an overview of rice leaf and butterfly wing fluid drag and self-cleaning studies. In addition, we examine two other promising aquatic surfaces in nature known for such properties, including fish scales and shark skin. Morphology, drag, self-cleaning, contact angle, and contact angle hysteresis data are presented to understand the role of wettability, viscosity, and velocity. Liquid repellent coatings are utilized to recreate or combine various effects. Discussion is provided along with conceptual models describing the role of surface structures related to low drag, self-cleaning, and antifouling properties. Modeling provides design guidance when developing novel low drag and self-cleaning surfaces for applications in the medical, marine, and industrial fields.

  3. Biologically inspired flexible quasi-single-mode random laser: An integration of Pieris canidia butterfly wing and semiconductors

    NASA Astrophysics Data System (ADS)

    Wang, Cih-Su; Chang, Tsung-Yuan; Lin, Tai-Yuan; Chen, Yang-Fang

    2014-10-01

    Quasi-periodic structures of natural biomaterial membranes have great potentials to serve as resonance cavities to generate ecological friendly optoelectronic devices with low cost. To achieve the first attempt for the illustration of the underlying principle, the Pieris canidia butterfly wing was embedded with ZnO nanoparticles. Quite interestingly, it is found that the bio-inspired quasi-single-mode random laser can be achieved by the assistance of the skeleton of the membrane, in which ZnO nanoparticles act as emitting gain media. Such unique characteristics can be interpreted well by the Fabry-Perot resonance existing in the window-like quasi-periodic structure of butterfly wing. Due to the inherently promising flexibility of butterfly wing membrane, the laser action can still be maintained during the bending process. Our demonstrated approach not only indicates that the natural biological structures can provide effective scattering feedbacks but also pave a new avenue towards designing bio-controlled photonic devices.

  4. Hydrophobic durability characteristics of butterfly wing surface after freezing cycles towards the design of nature inspired anti-icing surfaces

    PubMed Central

    Choy, Kwang-Leong

    2018-01-01

    The hydrophobicity and anti-icing performance of the surfaces of some artificial hydrophobic coatings degraded after several icing and de-icing cycles. In this paper, the frost formation on the surfaces of butterfly wings from ten different species was observed, and the contact angles were measured after 0 to 6 frosting/defrosting cycles. The results show that no obvious changes in contact angle for the butterfly wing specimens were not obvious during the frosting/defrosting process. Further, the conclusion was inferred that the topography of the butterfly wing surface forms a special space structure which has a larger space inside that can accommodate more frozen droplets; this behavior prevents destruction of the structure. The findings of this study may provide a basis and new concepts for the design of novel industrially important surfaces to inhibit frost/ice growth, such as durable anti-icing coatings, which may decrease or prevent the socio-economic loss. PMID:29385390

  5. Hydrophobic durability characteristics of butterfly wing surface after freezing cycles towards the design of nature inspired anti-icing surfaces.

    PubMed

    Chen, Tingkun; Cong, Qian; Qi, Yingchun; Jin, Jingfu; Choy, Kwang-Leong

    2018-01-01

    The hydrophobicity and anti-icing performance of the surfaces of some artificial hydrophobic coatings degraded after several icing and de-icing cycles. In this paper, the frost formation on the surfaces of butterfly wings from ten different species was observed, and the contact angles were measured after 0 to 6 frosting/defrosting cycles. The results show that no obvious changes in contact angle for the butterfly wing specimens were not obvious during the frosting/defrosting process. Further, the conclusion was inferred that the topography of the butterfly wing surface forms a special space structure which has a larger space inside that can accommodate more frozen droplets; this behavior prevents destruction of the structure. The findings of this study may provide a basis and new concepts for the design of novel industrially important surfaces to inhibit frost/ice growth, such as durable anti-icing coatings, which may decrease or prevent the socio-economic loss.

  6. The effect of butterfly-scale inspired patterning on leading-edge vortex growth

    NASA Astrophysics Data System (ADS)

    Wilroy, Jacob; Lang, Amy

    2015-11-01

    Leading edge vortices (LEVs) are important for generating thrust and lift in flapping flight, and the surface patterning (scales) on butterfly wings is hypothesized to play a role in the vortex formation of the LEV. To simplify this complex flow problem, an experiment was designed to focus on the alteration of 2-D vortex development with a variation in surface patterning. Specifically, the secondary vorticity generated by the LEV interacting at the patterned surface was studied and the subsequent affect on the growth rate of the circulation in the LEV. For this experiment we used butterfly inspired grooves attached to a flat plate and compared the vortex formation to a smooth plate case as the plate moved vertically. The plate is impulsively started in quiescent water and flow fields at Re = 1500, 3000, and 6000 are examined using Digital Particle Image Velocimetry (DPIV). The vortex formation time is 3.0 and is based on the flat plate travel length and chord length. We would like to thank the National Science Foundation REU Site Award 1358991 for funding this research.

  7. Beyond magic traits: Multimodal mating cues in Heliconius butterflies.

    PubMed

    Mérot, Claire; Frérot, Brigitte; Leppik, Ene; Joron, Mathieu

    2015-11-01

    Species coexistence involves the evolution of reproductive barriers opposing gene flow. Heliconius butterflies display colorful patterns affecting mate choice and survival through warning signaling and mimicry. These patterns are called "magic traits" for speciation because divergent natural selection may promote mimicry shifts in pattern whose role as mating cue facilitates reproductive isolation. By contrast, between comimetic species, natural selection promotes pattern convergence. We addressed whether visual convergence interferes with reproductive isolation by testing for sexual isolation between two closely related species with similar patterns, H. timareta thelxinoe and H. melpomene amaryllis. Experiments with models confirmed visual attraction based on wing phenotype, leading to indiscriminate approach. Nevertheless, mate choice experiments showed assortative mating. Monitoring male behavior toward live females revealed asymmetry in male preference, H. melpomene males courting both species equally while H. timareta males strongly preferred conspecifics. Experiments with hybrid males suggested an important genetic component for such asymmetry. Behavioral observations support a key role for short-distance cues in determining male choice in H. timareta. Scents extracts from wings and genitalia revealed interspecific divergence in chemical signatures, and hybrid female scent composition was significantly associated with courtship intensity by H. timareta males, providing candidate chemical mating cues involved in sexual isolation. © 2015 The Author(s). Evolution © 2015 The Society for the Study of Evolution.

  8. Colour formation on the wings of the butterfly Hypolimnas salmacis by scale stacking

    PubMed Central

    Siddique, Radwanul Hasan; Vignolini, Silvia; Bartels, Carolin; Wacker, Irene; Hölscher, Hendrik

    2016-01-01

    The butterfly genus Hypolimnas features iridescent blue colouration in some areas of its dorsal wings. Here, we analyse the mechanisms responsible for such colouration on the dorsal wings of Hypolimnas salmacis and experimentally demonstrate that the lower thin lamina in the white cover scales causes the blue iridescence. This outcome contradicts other studies reporting that the radiant blue in Hypolimnas butterflies is caused by complex ridge-lamellar architectures in the upper lamina of the cover scales. Our comprehensive optical study supported by numerical calculation however shows that scale stacking primarily induces the observed colour appearance of Hypolimnas salmacis. PMID:27805005

  9. Brilliant iridescence of Morpho butterfly wing scales is due to both a thin film lower lamina and a multilayered upper lamina.

    PubMed

    Giraldo, M A; Stavenga, D G

    2016-05-01

    Butterflies belonging to the nymphalid subfamily, Morphinae, are famous for their brilliant blue wing coloration and iridescence. These striking optical phenomena are commonly explained as to originate from multilayer reflections by the ridges of the wing scales. Because the lower lamina of the scales of related nymphalid butterflies, the Nymphalinae, plays a dominant role in the wing coloration, by acting as a thin film reflector, we investigated single blue scales of three characteristic Morpho species: M. epistrophus, M. helenor and M. cypris. The experimental data obtained by spectrophotometry, scatterometry and scanning electron microscopy demonstrated that also in the Morpho genus the lower lamina of both the cover and ground scales acts as an optical thin film reflector, contributing importantly to the blue structural coloration of the wings. Melanin pigment has a contrast-enhancing function in a sub-class of ground scales.

  10. Self-assembled carbon nanotube honeycomb networks using a butterfly wing template as a multifunctional nanobiohybrid.

    PubMed

    Miyako, Eijiro; Sugino, Takushi; Okazaki, Toshiya; Bianco, Alberto; Yudasaka, Masako; Iijima, Sumio

    2013-10-22

    Insect wings have many unique and complex nano/microstructures that are presently beyond the capabilities of any current technology to reproduce them artificially. In particular, Morpho butterflies are an attractive type of insect because their multifunctional wings are composed of nano/microstructures. In this paper, we show that carbon nanotube-containing composite adopts honeycomb-shaped networks when simply self-assembled on Morpho butterfly wings used as a template. The unique nano/microstructure of the composites exhibits multifunctionalities such as laser-triggered remote-heating, high electrical conductivity, and repetitive DNA amplification. Our present study highlights the important progress that has been made toward the development of smart nanobiomaterials for various applications such as digital diagnosis, soft wearable electronic devices, photosensors, and photovoltaic cells.

  11. A lack of Wolbachia-specific DNA in samples from apollo butterfly (Parnassius apollo, Lepidoptera: Papilionidae) individuals with deformed or reduced wings.

    PubMed

    Łukasiewicz, Kinga; Sanak, Marek; Węgrzyn, Grzegorz

    2016-05-01

    Various insects contain maternally inherited endosymbiotic bacteria which can cause reproductive alterations, modulation of some physiological responses (like immunity, heat shock response, and oxidative stress response), and resistance to viral infections. In butterflies, Wolbachia sp. is the most frequent endosymbiont from this group, occurring in about 30 % of species tested to date. In this report, the presence of Wolbachia-specific DNA has been detected in apollo butterfly (Parnassius apollo). In the isolated population of this insect occurring in Pieniny National Park (Poland), malformed individuals with deformed or reduced wings appear with an exceptionally high frequency. Interestingly, while total DNA isolated from most (about 85 %) normal insects contained Wolbachia-specific sequences detected by PCR, such sequences were absent in a large fraction (70 %) of individuals with deformed wings and in all tested individuals with reduced wings. These results indicate for the first time the correlation between malformation of wings and the absence of Wolbachia sp. in insects. Although the lack of the endosymbiotic bacteria cannot be considered as the sole cause of the deformation or reduction of wings, one might suggest that Wolbachia sp. could play a protective role in the ontogenetic development of apollo butterfly.

  12. Wing patterning gene redefines the mimetic history of Heliconius butterflies.

    PubMed

    Hines, Heather M; Counterman, Brian A; Papa, Riccardo; Albuquerque de Moura, Priscila; Cardoso, Marcio Z; Linares, Mauricio; Mallet, James; Reed, Robert D; Jiggins, Chris D; Kronforst, Marcus R; McMillan, W Owen

    2011-12-06

    The mimetic butterflies Heliconius erato and Heliconius melpomene have undergone parallel radiations to form a near-identical patchwork of over 20 different wing-pattern races across the Neotropics. Previous molecular phylogenetic work on these radiations has suggested that similar but geographically disjunct color patterns arose multiple times independently in each species. The neutral markers used in these studies, however, can move freely across color pattern boundaries, and therefore might not represent the history of the adaptive traits as accurately as markers linked to color pattern genes. To assess the evolutionary histories across different loci, we compared relationships among races within H. erato and within H. melpomene using a series of unlinked genes, genes linked to color pattern loci, and optix, a gene recently shown to control red color-pattern variation. We found that although unlinked genes partition populations by geographic region, optix had a different history, structuring lineages by red color patterns and supporting a single origin of red-rayed patterns within each species. Genes closely linked (80-250 kb) to optix exhibited only weak associations with color pattern. This study empirically demonstrates the necessity of examining phenotype-determining genomic regions to understand the history of adaptive change in rapidly radiating lineages. With these refined relationships, we resolve a long-standing debate about the origins of the races within each species, supporting the hypothesis that the red-rayed Amazonian pattern evolved recently and expanded, causing disjunctions of more ancestral patterns.

  13. A Babcock-Leighton Solar Dynamo Model with Multi-cellular Meridional Circulation in Advection- and Diffusion-dominated Regimes

    NASA Astrophysics Data System (ADS)

    Belucz, Bernadett; Dikpati, Mausumi; Forgács-Dajka, Emese

    2015-06-01

    Babcock-Leighton type-solar dynamo models with single-celled meridional circulation are successful in reproducing many solar cycle features. Recent observations and theoretical models of meridional circulation do not indicate a single-celled flow pattern. We examine the role of complex multi-cellular circulation patterns in a Babcock-Leighton solar dynamo in advection- and diffusion-dominated regimes. We show from simulations that the presence of a weak, second, high-latitude reverse cell speeds up the cycle and slightly enhances the poleward branch in the butterfly diagram, whereas the presence of a second cell in depth reverses the tilt of the butterfly wing to an antisolar type. A butterfly diagram constructed from the middle of convection zone yields a solar-like pattern, but this may be difficult to realize in the Sun because of magnetic buoyancy effects. Each of the above cases behaves similarly in higher and lower magnetic diffusivity regimes. However, our dynamo with a meridional circulation containing four cells in latitude behaves distinctly differently in the two regimes, producing solar-like butterfly diagrams with fast cycles in the higher diffusivity regime, and complex branches in butterfly diagrams in the lower diffusivity regime. We also find that dynamo solutions for a four-celled pattern, two in radius and two in latitude, prefer to quickly relax to quadrupolar parity if the bottom flow speed is strong enough, of similar order of magnitude as the surface flow speed.

  14. Correlations between adult mimicry and larval host plants in ithomiine butterflies.

    PubMed

    Willmott, Keith R; Mallet, James

    2004-08-07

    The apparent paradox of multiple coexisting wing pattern mimicry 'rings' in tropical butterflies has been explained as a result of microhabitat partitioning in adults. However, very few studies have tested this hypothesis. In neotropical forests, ithomiine butterflies dominate and display the richest diversity of mimicry rings. We show that co-mimetic species occupy the same larval host-plant species significantly more often than expected in two out of five communities that we surveyed; in one of these, the effect remains significant after phylogenetic correction. This relationship is most probably a result of a third correlated variable, such as microhabitat. Host-plant microhabitat may constrain adult movement, or host-plant choice may depend on butterfly microhabitat preferences and mimicry associations. This link between mimicry and host plant could help explain some host-plant and mimicry shifts, which have been important in the radiation of this speciose tropical group.

  15. Coevolution between flight morphology, vertical stratification and sexual dimorphism: what can we learn from tropical butterflies?

    PubMed

    Graça, M B; Pequeno, P A C L; Franklin, E; Morais, J W

    2017-10-01

    Occurrence patterns are partly shaped by the affinity of species with habitat conditions. For winged organisms, flight-related attributes are vital for ecological performance. However, due to the different reproductive roles of each sex, we expect divergence in flight energy budget, and consequently different selection responses between sexes. We used tropical frugivorous butterflies as models to investigate coevolution between flight morphology, sex dimorphism and vertical stratification. We studied 94 species of Amazonian fruit-feeding butterflies sampled in seven sites across 3341 ha. We used wing-thorax ratio as a proxy for flight capacity and hierarchical Bayesian modelling to estimate stratum preference. We detected a strong phylogenetic signal in wing-thorax ratio in both sexes. Stouter fast-flying species preferred the canopy, whereas more slender slow-flying species preferred the understorey. However, this relationship was stronger in females than in males, suggesting that female phenotype associates more intimately with habitat conditions. Within species, males were stouter than females and sexual dimorphism was sharper in understorey species. Because trait-habitat relationships were independent from phylogeny, the matching between flight morphology and stratum preference is more likely to reflect adaptive radiation than shared ancestry. This study sheds light on the impact of flight and sexual dimorphism on the evolution and ecological adaptation of flying organisms. © 2017 European Society For Evolutionary Biology. Journal of Evolutionary Biology © 2017 European Society For Evolutionary Biology.

  16. Butterfly wing colours: scale beads make white pierid wings brighter.

    PubMed Central

    Stavenga, D. G.; Stowe, S.; Siebke, K.; Zeil, J.; Arikawa, K.

    2004-01-01

    The wing-scale morphologies of the pierid butterflies Pieris rapae (small white) and Delias nigrina (common jezabel), and the heliconine Heliconius melpomene are compared and related to the wing-reflectance spectra. Light scattering at the wing scales determines the wing reflectance, but when the scales contain an absorbing pigment, reflectance is suppressed in the absorption wavelength range of the pigment. The reflectance of the white wing areas of P. rapae, where the scales are studded with beads, is considerably higher than that of the white wing areas of H. melpomene, which has scales lacking beads. The beads presumably cause the distinct matt-white colour of the wings of pierids and function to increase the reflectance amplitude. This will improve the visual discrimination between conspecific males and females. PMID:15306303

  17. A Burst of miRNA Innovation in the Early Evolution of Butterflies and Moths

    PubMed Central

    Quah, Shan; Hui, Jerome H.L.; Holland, Peter W.H.

    2015-01-01

    MicroRNAs (miRNAs) are involved in posttranscriptional regulation of gene expression. Because several miRNAs are known to affect the stability or translation of developmental regulatory genes, the origin of novel miRNAs may have contributed to the evolution of developmental processes and morphology. Lepidoptera (butterflies and moths) is a species-rich clade with a well-established phylogeny and abundant genomic resources, thereby representing an ideal system in which to study miRNA evolution. We sequenced small RNA libraries from developmental stages of two divergent lepidopterans, Cameraria ohridella (Horse chestnut Leafminer) and Pararge aegeria (Speckled Wood butterfly), discovering 90 and 81 conserved miRNAs, respectively, and many species-specific miRNA sequences. Mapping miRNAs onto the lepidopteran phylogeny reveals rapid miRNA turnover and an episode of miRNA fixation early in lepidopteran evolution, implying that miRNA acquisition accompanied the early radiation of the Lepidoptera. One lepidopteran-specific miRNA gene, miR-2768, is located within an intron of the homeobox gene invected, involved in insect segmental and wing patterning. We identified cubitus interruptus (ci) as a likely direct target of miR-2768, and validated this suppression using a luciferase assay system. We propose a model by which miR-2768 modulates expression of ci in the segmentation pathway and in patterning of lepidopteran wing primordia. PMID:25576364

  18. Mueller matrix microscopy on a Morpho butterfly

    NASA Astrophysics Data System (ADS)

    Arteaga, Oriol; Kuntman, Ertan; Antó, Joan; Pascual, Esther; Canillas, Adolf; Bertran, Enric

    2015-04-01

    The brilliant iridescent colouring in male Morpho butterflies is due to the microstrutures and nanostructures present in the wing scales, rather than pigments. In this work Mueller matrix microscopy is used to investigate the polarization properties of butterfly wing scales in reflection and transmission. It is found that the top layer of more transparent scales (cover scales) have very different polarimetric properties from the ground iridescent scales. Images with high spatial resolution showing the retarding and diattenuating optical properties for both types of scales are provided.

  19. Ancient homology underlies adaptive mimetic diversity across butterflies

    PubMed Central

    Gallant, Jason R.; Imhoff, Vance E.; Martin, Arnaud; Savage, Wesley K.; Chamberlain, Nicola L.; Pote, Ben L.; Peterson, Chelsea; Smith, Gabriella E.; Evans, Benjamin; Reed, Robert D.; Kronforst, Marcus R.; Mullen, Sean P.

    2014-01-01

    Convergent evolution provides a rare, natural experiment with which to test the predictability of adaptation at the molecular level. Little is known about the molecular basis of convergence over macro-evolutionary timescales. Here we use a combination of positional cloning, population genomic resequencing, association mapping and developmental data to demonstrate that positionally orthologous nucleotide variants in the upstream region of the same gene, WntA, are responsible for parallel mimetic variation in two butterfly lineages that diverged >65 million years ago. Furthermore, characterization of spatial patterns of WntA expression during development suggests that alternative regulatory mechanisms underlie wing pattern variation in each system. Taken together, our results reveal a strikingly predictable molecular basis for phenotypic convergence over deep evolutionary time. PMID:25198507

  20. Light trapping structures in wing scales of butterfly Trogonoptera brookiana.

    PubMed

    Han, Zhiwu; Niu, Shichao; Shang, Chunhui; Liu, Zhenning; Ren, Luquan

    2012-04-28

    The fine optical structures in wing scales of Trogonoptera brookiana, a tropical butterfly exhibiting efficient light trapping effect, were carefully examined and the reflectivity was measured using reflectance spectrometry. The optimized 3D configuration of the coupling structure was determined using SEM and TEM data, and the light trapping mechanism of butterfly scales was studied. It is found that the front and back sides of butterfly wings possess different light trapping structures, but both can significantly increase the optical path and thus result in almost total absorption of all incident light. An optical model was created to check the properties of this light trapping structure. The simulated reflectance spectra are in concordance with the experimental ones. The results reliably confirm that these structures induce efficient light trapping effect. This functional "biomimetic structure" would have a potential value in wide engineering and optical applications. This journal is © The Royal Society of Chemistry 2012

  1. Iridescence and spectral filtering of the gyroid-type photonic crystals in Parides sesostris wing scales

    PubMed Central

    Wilts, Bodo D.; Michielsen, Kristel; De Raedt, Hans; Stavenga, Doekele G.

    2012-01-01

    The cover scales on the wing of the Emerald-patched Cattleheart butterfly, Parides sesostris, contain gyroid-type biological photonic crystals that brightly reflect green light. A pigment, which absorbs maximally at approximately 395 nm, is immersed predominantly throughout the elaborate upper lamina. This pigment acts as a long-pass filter shaping the reflectance spectrum of the underlying photonic crystals. The additional effect of the filtering is that the spatial distribution of the scale reflectance is approximately angle-independent, leading to a stable wing pattern contrast. The spectral tuning of the original reflectance is verified by photonic band structure modelling. PMID:24098853

  2. Evolutionary Novelty in a Butterfly Wing Pattern through Enhancer Shuffling

    PubMed Central

    Pardo-Diaz, Carolina; Hanly, Joseph J.; Martin, Simon H.; Mallet, James; Dasmahapatra, Kanchon K.; Salazar, Camilo; Joron, Mathieu; Nadeau, Nicola; McMillan, W. Owen; Jiggins, Chris D.

    2016-01-01

    An important goal in evolutionary biology is to understand the genetic changes underlying novel morphological structures. We investigated the origins of a complex wing pattern found among Amazonian Heliconius butterflies. Genome sequence data from 142 individuals across 17 species identified narrow regions associated with two distinct red colour pattern elements, dennis and ray. We hypothesise that these modules in non-coding sequence represent distinct cis-regulatory loci that control expression of the transcription factor optix, which in turn controls red pattern variation across Heliconius. Phylogenetic analysis of the two elements demonstrated that they have distinct evolutionary histories and that novel adaptive morphological variation was created by shuffling these cis-regulatory modules through recombination between divergent lineages. In addition, recombination of modules into different combinations within species further contributes to diversity. Analysis of the timing of diversification in these two regions supports the hypothesis of introgression moving regulatory modules between species, rather than shared ancestral variation. The dennis phenotype introgressed into Heliconius melpomene at about the same time that ray originated in this group, while ray introgressed back into H. elevatus much more recently. We show that shuffling of existing enhancer elements both within and between species provides a mechanism for rapid diversification and generation of novel morphological combinations during adaptive radiation. PMID:26771987

  3. Evolutionary Novelty in a Butterfly Wing Pattern through Enhancer Shuffling.

    PubMed

    Wallbank, Richard W R; Baxter, Simon W; Pardo-Diaz, Carolina; Hanly, Joseph J; Martin, Simon H; Mallet, James; Dasmahapatra, Kanchon K; Salazar, Camilo; Joron, Mathieu; Nadeau, Nicola; McMillan, W Owen; Jiggins, Chris D

    2016-01-01

    An important goal in evolutionary biology is to understand the genetic changes underlying novel morphological structures. We investigated the origins of a complex wing pattern found among Amazonian Heliconius butterflies. Genome sequence data from 142 individuals across 17 species identified narrow regions associated with two distinct red colour pattern elements, dennis and ray. We hypothesise that these modules in non-coding sequence represent distinct cis-regulatory loci that control expression of the transcription factor optix, which in turn controls red pattern variation across Heliconius. Phylogenetic analysis of the two elements demonstrated that they have distinct evolutionary histories and that novel adaptive morphological variation was created by shuffling these cis-regulatory modules through recombination between divergent lineages. In addition, recombination of modules into different combinations within species further contributes to diversity. Analysis of the timing of diversification in these two regions supports the hypothesis of introgression moving regulatory modules between species, rather than shared ancestral variation. The dennis phenotype introgressed into Heliconius melpomene at about the same time that ray originated in this group, while ray introgressed back into H. elevatus much more recently. We show that shuffling of existing enhancer elements both within and between species provides a mechanism for rapid diversification and generation of novel morphological combinations during adaptive radiation.

  4. Use of butterflies as nontarget insect test species and the acute toxicity and hazard of mosquito control insecticides.

    PubMed

    Hoang, Tham C; Pryor, Rachel L; Rand, Gary M; Frakes, Robert A

    2011-04-01

    Honeybees are the standard insect test species used for toxicity testing of pesticides on nontarget insects for the U.S. Environmental Protection Agency (U.S. EPA) under the Federal Insecticide Fungicide and Rodenticide Act (FIFRA). Butterflies are another important insect order and a valued ecological resource in pollination. The current study conducted acute toxicity tests with naled, permethrin, and dichlorvos on fifth larval instar (caterpillars) and adults of different native Florida, USA, butterfly species to determine median lethal doses (24-h LD50), because limited acute toxicity data are available with this major insect group. Thorax- and wing-only applications of each insecticide were conducted. Based on LD50s, thorax and wing application exposures were acutely toxic to both caterpillars and adults. Permethrin was the most acutely toxic insecticide after thorax exposure to fifth instars and adult butterflies. However, no generalization on acute toxicity (sensitivity) of the insecticides could be concluded based on exposures to fifth instars versus adult butterflies or on thorax versus wing exposures of adult butterflies. A comparison of LD50s of the butterflies from this study (caterpillars and adults) with honeybee LD50s for the adult mosquito insecticides on a µg/organism or µg/g basis indicates that several butterfly species are more sensitive to these insecticides than are honeybees. A comparison of species sensitivity distributions for all three insecticides shows that permethrin had the lowest 10th percentile. Using a hazard quotient approach indicates that both permethrin and naled applications in the field may present potential acute hazards to butterflies, whereas no acute hazard of dichlorvos is apparent in butterflies. Butterflies should be considered as potential test organisms when nontarget insect testing of pesticides is suggested under FIFRA. Copyright © 2011 SETAC.

  5. Vertical distribution, flight behaviour and evolution of wing morphology in Morpho butterflies.

    PubMed

    Devries, P J; Penz, Carla M; Hill, Ryan I

    2010-09-01

    1. Flight is a key innovation in the evolution of insects that is crucial to their dispersal, migration, territoriality, courtship and predator avoidance. Male butterflies have characteristic territoriality and courtship flight behaviours, and females use a characteristic flight behaviour when searching for host plants. This implies that selection acts on wing morphology to maximize flight performance for conducting important behaviours among sexes. 2. Butterflies in the genus Morpho are obvious components of neotropical forests, and many observations indicate that they show two broad categories of flight behaviour and flight height. Although species can be categorized as using gliding or flapping flight, and flying at either canopy or understorey height, the association of flight behaviour and flight height with wing shape evolution has never been explored. 3. Two clades within Morpho differ in flight behaviour and height. Males and females of one clade inhabit the forest understorey and use flapping flight, whereas in the other clade, males use gliding flight at canopy level and females use flapping flight in both canopy and understorey. 4. We used independent contrasts to answer whether wing shape is associated with flight behaviour and height. Given a single switch to canopy habitation and gliding flight, we compared contrasts for the node at which the switch to canopy flight occurred with the distribution of values in the two focal clades. We found significant changes in wing shape at the transition to canopy flight only in males, and no change in size for either sex. A second node within the canopy clade suggests that other factors may also be involved in wing shape evolution. Our results reinforce the hypothesis that natural selection acts differently on male and female butterfly wing shape and indicate that the transition to canopy flight cannot explain all wing shape diversity in Morpho. 5. This study provides a starting point for characterizing evolution of wing morphology in forest butterflies in the contexts of habitat selection and flight behaviour. Further, these observations suggest that exploring wing shape evolution for canopy and understorey species in other insects may help understand the effects of habitat destruction on biological diversity.

  6. Reversible thermochromic response based on photonic crystal structure in butterfly wing

    NASA Astrophysics Data System (ADS)

    Wang, Wanlin; Wang, Guo Ping; Zhang, Wang; Zhang, Di

    2018-01-01

    Subtle responsive properties can be achieved by the photonic crystal (PC) nanostructures of butterfly based on thermal expansion effect. The studies focused on making the sample visually distinct. However, the response is restricted by limited thermal expansion coefficients. We herein report a new class of reversible thermochromic response achieved by controlling the ambient refractive index in butterfly PC structure. The photonic ethanol-filled nanoarchitecture sample is simply assembled by sealing liquid ethanol filling Papilio ulysses butterfly wing. Volatile ethanol is used to modulate the ambient refractive index. The sample is sealed with glasses to ensure reversibility. Liquid ethanol filling butterfly wing demonstrated significant allochroic response to ambient refractive index, which can be controlled by the liquefaction and vaporization of ethanol. This design is capable of converting thermal energy into visual color signals. The mechanism of this distinct response is simulated and proven by band theory. The response properties are performed with different filled chemicals and different structure parameters. Thus, the reversible thermochromic response design might have potential use in the fields such as detection, photonic switch, displays, and so forth.

  7. Wings of the butterfly: Sunspot groups for 1826-2015

    NASA Astrophysics Data System (ADS)

    Leussu, R.; Usoskin, I. G.; Senthamizh Pavai, V.; Diercke, A.; Arlt, R.; Denker, C.; Mursula, K.

    2017-03-01

    The spatio-temporal evolution of sunspot activity, the so-called Maunder butterfly diagram, has been continously available since 1874 using data from the Royal Greenwich Observatory, extended by SOON network data after 1976. Here we present a new extended butterfly diagram of sunspot group occurrence since 1826, using the recently digitized data from Schwabe (1826-1867) and Spörer (1866-1880). The wings of the diagram are separated using a recently developed method based on an analysis of long gaps in sunspot group occurrence in different latitude bands. We define characteristic latitudes, corresponding to the start, end, and the largest extent of the wings (the F, L, and H latitudes). The H latitudes (30°-45°) are highly significantly correlated with the strength of the wings (quantified by the total sum of the monthly numbers of sunspot groups). The F latitudes (20°-30°) depict a weak tendency, especially in the southern hemisphere, to follow the wing strength. The L latitudes (2°-10°) show no clear relation to the wing strength. Overall, stronger cycle wings tend to start at higher latitudes and have a greater wing extent. A strong (5-6)-cycle periodic oscillation is found in the start and end times of the wings and in the overlap and gaps between successive wings of one hemisphere. While the average wing overlap is zero in the southern hemisphere, it is two to three months in the north. A marginally significant oscillation of about ten solar cycles is found in the asymmetry of the L latitudes. The new long database of butterfly wings provides new observational constraints to solar dynamo models that discuss the spatio-temporal distribution of sunspot occurrence over the solar cycle and longer. Digital data for Fig. 1 are available at the CDS via anonymous ftp to http://cdsarc.u-strasbg.fr (http://130.79.128.5) or via http://cdsarc.u-strasbg.fr/viz-bin/qcat?J/A+A/599/A131

  8. Enhanced thrust and speed revealed in the forward flight of a butterfly with transient body translation.

    PubMed

    Fei, Yueh-Han John; Yang, Jing-Tang

    2015-09-01

    A butterfly with broad wings, flapping at a small frequency, flies an erratic trajectory at an inconstant speed. A large variation of speed within a cycle is observed in the forward flight of a butterfly. A self-propulsion model to simulate a butterfly is thus created to investigate the transient translation of the body; the results, which are in accordance with experimental data, show that the shape of the variation of the flight speed is similar to a sinusoidal wave with a maximum (J=0.89) at the beginning of the downstroke, and a decrease to a minimum (J=0.17) during a transition from downstroke to upstroke; the difference between the extrema of the flight speed is enormous in a flapping cycle. At a high speed, a clapping motion of the butterfly wings decreases the generation of drag. At a small speed, a butterfly is able to capture the induced wakes generated in a downstroke, and effectively generates a thrust at the beginning of an upstroke. The wing motion of a butterfly skillfully interacts with its speed so as to enable an increased speed with the same motion. Considering a butterfly to fly in a constant inflow leads to either an underestimate of its speed or an overestimate of its generated lift, which yields an inaccurate interpretation of the insect's flight. Our results reveal the effect of transient translation on a butterfly in forward flight, which is especially important for an insect with a small flapping frequency.

  9. Enhanced thrust and speed revealed in the forward flight of a butterfly with transient body translation

    NASA Astrophysics Data System (ADS)

    Fei, Yueh-Han John; Yang, Jing-Tang

    2015-09-01

    A butterfly with broad wings, flapping at a small frequency, flies an erratic trajectory at an inconstant speed. A large variation of speed within a cycle is observed in the forward flight of a butterfly. A self-propulsion model to simulate a butterfly is thus created to investigate the transient translation of the body; the results, which are in accordance with experimental data, show that the shape of the variation of the flight speed is similar to a sinusoidal wave with a maximum (J =0.89 ) at the beginning of the downstroke, and a decrease to a minimum (J =0.17 ) during a transition from downstroke to upstroke; the difference between the extrema of the flight speed is enormous in a flapping cycle. At a high speed, a clapping motion of the butterfly wings decreases the generation of drag. At a small speed, a butterfly is able to capture the induced wakes generated in a downstroke, and effectively generates a thrust at the beginning of an upstroke. The wing motion of a butterfly skillfully interacts with its speed so as to enable an increased speed with the same motion. Considering a butterfly to fly in a constant inflow leads to either an underestimate of its speed or an overestimate of its generated lift, which yields an inaccurate interpretation of the insect's flight. Our results reveal the effect of transient translation on a butterfly in forward flight, which is especially important for an insect with a small flapping frequency.

  10. Deimatic display in the European swallowtail butterfly as a secondary defence against attacks from great tits.

    PubMed

    Olofsson, Martin; Eriksson, Stephan; Jakobsson, Sven; Wiklund, Christer

    2012-01-01

    Many animals reduce the risk of being attacked by a predator through crypsis, masquerade or, alternatively, by advertising unprofitability by means of aposematic signalling. Behavioural attributes in prey employed after discovery, however, signify the importance of also having an effective secondary defence if a predator uncovers, or is immune to, the prey's primary defence. In butterflies, as in most animals, secondary defence generally consists of escape flights. However, some butterfly species have evolved other means of secondary defence such as deimatic displays/startle displays. The European swallowtail, Papilio machaon, employs what appears to be a startle display by exposing its brightly coloured dorsal wing surface upon disturbance and, if the disturbance continues, by intermittently protracting and relaxing its wing muscles generating a jerky motion of the wings. This display appears directed towards predators but whether it is effective in intimidating predators so that they refrain from attacks has never been tested experimentally. In this study we staged encounters between a passerine predator, the great tit, Parus major, and live and dead swallowtail butterflies in a two-choice experiment. Results showed that the dead butterfly was virtually always attacked before the live butterfly, and that it took four times longer before a bird attacked the live butterfly. When the live butterfly was approached by a bird this generally elicited the butterfly's startle display, which usually caused the approaching bird to flee. We also performed a palatability test of the butterflies and results show that the great tits seemed to find them palatable. We conclude that the swallowtail's startle display of conspicuous coloration and jerky movements is an efficient secondary defence against small passerines. We also discuss under what conditions predator-prey systems are likely to aid the evolution of deimatic behaviours in harmless and palatable prey.

  11. Chromosomal rearrangements maintain a polymorphic supergene controlling butterfly mimicry

    PubMed Central

    Joron, Mathieu; Frezal, Lise; Jones, Robert T.; Chamberlain, Nicola L.; Lee, Siu F.; Haag, Christoph R.; Whibley, Annabel; Becuwe, Michel; Baxter, Simon W.; Ferguson, Laura; Wilkinson, Paul A.; Salazar, Camilo; Davidson, Claire; Clark, Richard; Quail, Michael A.; Beasley, Helen; Glithero, Rebecca; Lloyd, Christine; Sims, Sarah; Jones, Matthew C.; Rogers, Jane; Jiggins, Chris D.; ffrench-Constant, Richard H.

    2013-01-01

    Supergenes are tight clusters of loci that facilitate the co-segregation of adaptive variation, providing integrated control of complex adaptive phenotypes1. Polymorphic supergenes, in which specific combinations of traits are maintained within a single population, were first described for ‘pin’ and ‘thrum’ floral types in Primula1 and Fagopyrum2, but classic examples are also found in insect mimicry3–5 and snail morphology6. Understanding the evolutionary mechanisms that generate these co-adapted gene sets, as well as the mode of limiting the production of unfit recombinant forms, remains a substantial challenge7–10. Here we show that individual wing-pattern morphs in the polymorphic mimetic butterfly Heliconius numata are associated with different genomic rearrangements at the supergene locus P. These rearrangements tighten the genetic linkage between at least two colour-pattern loci that are known to recombine in closely related species9–11, with complete suppression of recombination being observed in experimental crosses across a 400-kilobase interval containing at least 18 genes. In natural populations, notable patterns of linkage disequilibrium (LD) are observed across the entire P region. The resulting divergent haplotype clades and inversion breakpoints are found in complete association with wing-pattern morphs. Our results indicate that allelic combinations at known wing-patterning loci have become locked together in a polymorphic rearrangement at the Plocus, forming a supergene that acts as a simple switch between complex adaptive phenotypes found in sympatry. These findings highlight how genomic rearrangements can have a central role in the coexistence of adaptive phenotypes involving several genes acting in concert, by locally limiting recombination and gene flow. PMID:21841803

  12. Butterfly genomics eclosing.

    PubMed

    Beldade, P; McMillan, W O; Papanicolaou, A

    2008-02-01

    Technological and conceptual advances of the last decade have led to an explosion of genomic data and the emergence of new research avenues. Evolutionary and ecological functional genomics, with its focus on the genes that affect ecological success and adaptation in natural populations, benefits immensely from a phylogenetically widespread sampling of biological patterns and processes. Among those organisms outside established model systems, butterflies offer exceptional opportunities for multidisciplinary research on the processes generating and maintaining variation in ecologically relevant traits. Here we highlight research on wing color pattern variation in two groups of Nymphalid butterflies, the African species Bicyclus anynana (subfamily Satyrinae) and species of the South American genus Heliconius (subfamily Heliconiinae), which are emerging as important systems for studying the nature and origins of functional diversity. Growing genomic resources including genomic and cDNA libraries, dense genetic maps, high-density gene arrays, and genetic transformation techniques are extending current gene mapping and expression profiling analysis and enabling the next generation of research questions linking genes, development, form, and fitness. Efforts to develop such resources in Bicyclus and Heliconius underscore the general challenges facing the larger research community and highlight the need for a community-wide effort to extend ongoing functional genomic research on butterflies.

  13. Did Adult Diurnal Activity Influence the Evolution of Wing Morphology in Opoptera Butterflies?

    PubMed

    Penz, C M; Heine, K B

    2016-02-01

    The butterfly genus Opoptera includes eight species, three of which have diurnal habits while the others are crepuscular (the usual activity period for members of the tribe Brassolini). Although never measured in the field, it is presumed that diurnal Opoptera species potentially spend more time flying than their crepuscular relatives. If a shift to diurnal habits potentially leads to a higher level of activity and energy expenditure during flight, then selection should operate on increased aerodynamic and energetic efficiency, leading to changes in wing shape. Accordingly, we ask whether diurnal habits have influenced the evolution of wing morphology in Opoptera. Using phylogenetically independent contrasts and Wilcoxon rank sum tests, we confirmed our expectation that the wings of diurnal species have higher aspect ratios (ARs) and lower wing centroids (WCs) than crepuscular congeners. These wing shape characteristics are known to promote energy efficiency during flight. Three Opoptera wing morphotypes established a priori significantly differed in AR and WC values. The crepuscular, cloud forest dweller Opoptera staudingeri (Godman & Salvin) was exceptional in having an extended forewing tip and the highest AR and lowest WC within Opoptera, possibly to facilitate flight in a cooler environment. Our study is the first to investigate how butterfly wing morphology might evolve as a response to a behavioral shift in adult time of activity.

  14. Hierarchical photonic structured stimuli-responsive materials as high-performance colorimetric sensors

    NASA Astrophysics Data System (ADS)

    Lu, Tao; Zhu, Shenmin; Chen, Zhixin; Wang, Wanlin; Zhang, Wang; Zhang, Di

    2016-05-01

    Hierarchical photonic structures in nature are of special interest because they can be used as templates for fabrication of stimuli-responsive photonic crystals (PCs) with unique structures beyond man-made synthesis. The current stimuli-responsive PCs templated directly from natural PCs showed a very weak external stimuli response and poor durability due to the limitations of natural templates. Herein, we tackle this problem by chemically coating functional polymers, polyacrylamide, on butterfly wing scales which have hierarchical photonic structures. As a result of the combination of the strong water absorption properties of the polyacrylamide and the PC structures of the butterfly wing scales, the designed materials demonstrated excellent humidity responsive properties and a tremendous colour change. The colour change is induced by the refractive index change which is in turn due to the swollen nature of the polymer when the relative humidity changes. The butterfly wing scales also showed an excellent durability which is due to the chemical bonds formed between the polymer and wing scales. The synthesis strategy provides an avenue for the promising applications of stimuli-responsive PCs with hierarchical structures.Hierarchical photonic structures in nature are of special interest because they can be used as templates for fabrication of stimuli-responsive photonic crystals (PCs) with unique structures beyond man-made synthesis. The current stimuli-responsive PCs templated directly from natural PCs showed a very weak external stimuli response and poor durability due to the limitations of natural templates. Herein, we tackle this problem by chemically coating functional polymers, polyacrylamide, on butterfly wing scales which have hierarchical photonic structures. As a result of the combination of the strong water absorption properties of the polyacrylamide and the PC structures of the butterfly wing scales, the designed materials demonstrated excellent humidity responsive properties and a tremendous colour change. The colour change is induced by the refractive index change which is in turn due to the swollen nature of the polymer when the relative humidity changes. The butterfly wing scales also showed an excellent durability which is due to the chemical bonds formed between the polymer and wing scales. The synthesis strategy provides an avenue for the promising applications of stimuli-responsive PCs with hierarchical structures. Electronic supplementary information (ESI) available. See DOI: 10.1039/c6nr01875k

  15. Effect of wing mass in free flight by a butterfly-like 3D flapping wing-body model

    NASA Astrophysics Data System (ADS)

    Suzuki, Kosuke; Okada, Iori; Yoshino, Masato

    2016-11-01

    The effect of wing mass in free flight of a flapping wing is investigated by numerical simulations based on an immersed boundary-lattice Boltzmann method. We consider a butterfly-like 3D flapping wing-model consisting of two square wings with uniform mass density connected by a rod-shaped body. We simulate free flights of the wing-body model with various mass ratios of the wing to the whole of the model. As a result, it is found that the lift and thrust forces decrease as the mass ratio increases, since the body with a large mass ratio experiences large vertical and horizontal oscillations in one period and consequently the wing tip speed relatively decreases. In addition, we find the critical mass ratio between upward flight and downward flight for various Reynolds numbers. This work was supported by JSPS KAKENHI Grant Number JP16K18012.

  16. Extended season for northern butterflies.

    PubMed

    Karlsson, Bengt

    2014-07-01

    Butterflies are like all insects in that they are temperature sensitive and a changing climate with higher temperatures might effect their phenology. Several studies have found support for earlier flight dates among the investigated species. A comparative study with data from a citizen science project, including 66 species of butterflies in Sweden, was undertaken, and the result confirms that most butterfly species now fly earlier during the season. This is especially evident for butterflies overwintering as adults or as pupae. However, the advancement in phenology is correlated with flight date, and some late season species show no advancement or have even postponed their flight dates and are now flying later in the season. The results also showed that latitude had a strong effect on the adult flight date, and most of the investigated species showed significantly later flights towards the north. Only some late flying species showed an opposite trend, flying earlier in the north. A majority of the investigated species in this study showed a general response to temperature and advanced their flight dates with warmer temperatures (on average they advanced their flight dates by 3.8 days/°C), although not all species showed this response. In essence, a climate with earlier springs and longer growing seasons seems not to change the appearance patterns in a one-way direction. We now see butterflies on the wings both earlier and later in the season and some consequences of these patterns are discussed. So far, studies have concentrated mostly on early season butterfly-plant interactions but also late season studies are needed for a better understanding of long-term population consequences.

  17. Modifying the anti-wetting property of butterfly wings and water strider legs by atomic layer deposition coating: surface materials versus geometry.

    PubMed

    Ding, Yong; Xu, Sheng; Zhang, Yue; Wang, Aurelia C; Wang, Melissa H; Xiu, Yonghao; Wong, Ching Ping; Wang, Zhong Lin

    2008-09-03

    Although butterfly wings and water strider legs have an anti-wetting property, their working conditions are quite different. Water striders, for example, live in a wet environment and their legs need to support their weight and bear the high pressure during motion. In this work, we have focused on the importance of the surface geometrical structures in determining their performance. We have applied an atomic layer deposition technique to coat the surfaces of both butterfly wings and water strider legs with a uniform 30 nm thick hydrophilic Al(2)O(3) film. By keeping the surface material the same, we have studied the effect of different surface roughness/structure on their hydrophobic property. After the surface coating, the butterfly wings changed to become hydrophilic, while the water strider legs still remained super-hydrophobic. We suggest that the super-hydrophobic property of the water strider is due to the special shape of the long inclining spindly cone-shaped setae at the surface. The roughness in the surface can enhance the natural tendency to be hydrophobic or hydrophilic, while the roughness in the normal direction of the surface is favorable for forming a composite interface.

  18. Lesions in the wingless gene of the Apollo butterfly (Parnassius apollo, Lepidoptera: Papilionidae) individuals with deformed or reduced wings, coming from the isolated population in Pieniny (Poland).

    PubMed

    Łukasiewicz, Kinga; Sanak, Marek; Węgrzyn, Grzegorz

    2016-02-01

    Parnassius apollo (Lepidoptera: Papilionidae) is a butterfly species which was common in Europe in 19th century, but now it is considered as near threatened. Various programs devoted to protect and save P. apollo have been established, between others the one in Pieniny National Park (Poland). An isolated population of this butterfly has been restored there from a small group of 20-30 individuals in early 1990s. However, deformations or reductions of wings occur in this population in a relatively large number of insects, and the cause of this phenomenon is not known. In this report, the occurrence of lesions in the wingless (wg) gene is demonstrated in most of tested butterflies with deformed or reduced wings, but not in normal insects. Although the analyses indicated that wg lesion(s) cannot be the sole cause of the deformed or reduced wings in the population of P. apollo from Pieniny, the discovery that this genetic defect occurs in most of malformed individuals, can be considered as an important step in understanding this phenomenon. Copyright © 2015 Elsevier B.V. All rights reserved.

  19. Roughness Effects on the Formation of a Leading Edge Vortex

    NASA Astrophysics Data System (ADS)

    Elliott, Cassidy; Lang, Amy; Wahidi, Redha; Wilroy, Jacob

    2016-11-01

    Microscopic scales cover the wings of Monarch butterflies, creating a patterned surface. This patterning is an important natural flow control mechanism that is thought to delay the growth of the leading edge vortex (LEV) produced by the flapping motion of a wing. The increased skin friction caused by the scales leads to a weaker LEV being shed into the butterfly's wake, lessening drag and increasing flight efficiency. To test this theory, a plate of random roughness was designed in SolidWorks and printed on the Objet 30 Pro 3D printer. A 2x3x5 cubic foot tow tank was used to test the rough plate at Reynold's numbers of 1500, 3000, and 6000 (velocities of 8, 16, and 32 mm/s) at an angle of attack of 45 degrees. Particle Image Velocimetry (PIV) captured images of the LEV generated by the plate when towed upwards through the particle-seeded flow. Codes written in MatLab were used to automatically track and determine the strength of the LEV. Circulation values for the randomly-rough plate were then compared to the same values generated in a previous experiment that used a smooth plate and a grooved plate to determine the effect of the patterning on vortex development. Funding provided by NSF REU site Grant EEC 1358991 and CBET 1628600.

  20. Mosquito control insecticides: a probabilistic ecological risk assessment on drift exposures of naled, dichlorvos (naled metabolite) and permethrin to adult butterflies.

    PubMed

    Hoang, T C; Rand, G M

    2015-01-01

    A comprehensive probabilistic terrestrial ecological risk assessment (ERA) was conducted to characterize the potential risk of mosquito control insecticide (i.e., naled, it's metabolite dichlorvos, and permethrin) usage to adult butterflies in south Florida by comparing the probability distributions of environmental exposure concentrations following actual mosquito control applications at labeled rates from ten field monitoring studies with the probability distributions of butterfly species response (effects) data from our laboratory acute toxicity studies. The overlap of these distributions was used as a measure of risk to butterflies. The long-term viability (survival) of adult butterflies, following topical (thorax/wings) exposures was the environmental value we wanted to protect. Laboratory acute toxicity studies (24-h LD50) included topical exposures (thorax and wings) to five adult butterfly species and preparation of species sensitivity distributions (SSDs). The ERA indicated that the assessment endpoint of protection, of at least 90% of the species, 90% of the time (or the 10th percentile from the acute SSDs) from acute naled and permethrin exposures, is most likely not occurring when considering topical exposures to adults. Although the surface areas for adulticide exposures are greater for the wings, exposures to the thorax provide the highest potential for risk (i.e., SSD 10th percentile is lowest) for adult butterflies. Dichlorvos appeared to present no risk. The results of this ERA can be applied to other areas of the world, where these insecticides are used and where butterflies may be exposed. Since there are other sources (e.g., agriculture) of pesticides in the environment, where butterfly exposures will occur, the ERA may under-estimate the potential risks under real-world conditions. Copyright © 2014 Elsevier B.V. All rights reserved.

  1. Butterflies' wings deformations using high speed digital holographic interferometry

    NASA Astrophysics Data System (ADS)

    Mendoza Santoyo, Fernando; Aguayo, Daniel D.; de La Torre-Ibarra, Manuel H.; Salas-Araiza, Manuel D.

    2011-08-01

    A variety of efforts in different scientific disciplines have tried to mimic the insect's in-flight complex system. The gained knowledge has been applied to improve the performance of different flying artifacts. In this research report it is presented a displacement measurement on butterflies' wings using the optical noninvasive Digital Holographic Interferometry technique with out of plane sensitivity, using a high power cw laser and a high speed CMOS camera to record the unrepeatable displacement movements on these organic tissues. A series of digital holographic interferograms were recorded and the experimental results for several butterflies during flapping events. The relative unwrapped phase maps micro-displacements over the whole wing surface are shown in a wire-mesh representation. The difference between flying modes is remarkably depicted among them.

  2. Anti-fouling properties of microstructured surfaces bio-inspired by rice leaves and butterfly wings.

    PubMed

    Bixler, Gregory D; Theiss, Andrew; Bhushan, Bharat; Lee, Stephen C

    2014-04-01

    Material scientists often look to biology for new engineering solutions to materials science problems. For example, unique surface characteristics of rice leaves and butterfly wings combine the shark skin (antifouling) and lotus leaf (self-cleaning) effects, producing the so-called rice and butterfly wing effect. In this paper, we study antifouling properties of four microstructured surfaces inspired by rice leaves and fabricated with photolithography and hot embossing techniques. Anti-biofouling effectiveness is determined with bioassays using Escherichia coli whilst inorganic fouling with simulated dirt particles. Antifouling data are presented to understand the role of surface geometrical features resistance to fouling. Conceptual modeling provides design guidance when developing novel antifouling surfaces for applications in the medical, marine, and industrial fields. Copyright © 2013 Elsevier Inc. All rights reserved.

  3. Analytical and experimental investigation of microstructural alterations in bearing steel in rolling contact fatigue

    NASA Astrophysics Data System (ADS)

    Mobasher Moghaddam, Sina

    Rolling Contact Fatigue (RCF) is one the most common failure modes in bearings. RCF is usually associated with particular microstructural alterations. Such alterations (i.e. white etching cracks, butterflies, etc.) which lead to RCF failure are known to be among the most concerning matters to bearing industry. In the current work, an analytical as well as experimental approaches are used to investigate "butterfly wing" formation, crack initiation and propagation from inclusions. A new damage evolution equation coupled with a FE model is employed to account for the effect of mean stresses and alternating stresses simultaneously to investigate butterfly formation. The proposed damage evolution law matches experimentally observed butterfly orientation, shape, and size successfully. The model is used to obtain S-N results for butterfly formation at different Hertzian load levels. The results corroborate well with the experimental data available in the open literature. The model is used to predict debonding at the inclusion/matrix interface and the most vulnerable regions for crack initiation on butterfly/matrix interface. A new variable called butterfly formation index (BFI) is introduced to manifest the dependence of wing formation on depth. The value of critical damage inside the butterfly wings was obtained experimentally and was then used to simulate damage evolution. Voronoi tessellation was used to develop the FEM domains to capture the effect of microstructural randomness on butterfly wing formation, crack initiation and propagation. Then, the effects of different inclusion characteristics such as size, depth, and stiffness on RCF life are studied. The results show that stiffness of an inclusion and its location has a significant effect on the RCF life: stiffer inclusions and inclusions located at the depth of maximum shear stress reversal are more detrimental to the RCF life. Stress concentrations are not significantly affected by inclusion size for the cases investigated; however, a stereology study showed that larger inclusions have a higher chance to be located at the critical depth and cause failure. Crack maps were recorded and compared to spall geometries observed experimentally. The results show that crack initiation locations and final spall shapes are similar to what has been observed in failed bearings.

  4. Describe Your Favorite Teacher.

    ERIC Educational Resources Information Center

    Dill, Isaac; Dill, Vicky

    1993-01-01

    A third grader describes Ms. Gonzalez, his favorite teacher, who left to accept a more lucrative teaching assignment. Ms. Gonzalez' butterflies unit covered everything from songs about social butterflies to paintings of butterfly wings, anatomy studies, and student haiku poems and biographies. Students studied biology by growing popcorn plants…

  5. The redder the better: wing color predicts flight performance in monarch butterflies.

    PubMed

    Davis, Andrew K; Chi, Jean; Bradley, Catherine; Altizer, Sonia

    2012-01-01

    The distinctive orange and black wings of monarchs (Danaus plexippus) have long been known to advertise their bitter taste and toxicity to potential predators. Recent work also showed that both the orange and black coloration of this species can vary in response to individual-level and environmental factors. Here we examine the relationship between wing color and flight performance in captive-reared monarchs using a tethered flight mill apparatus to quantify butterfly flight speed, duration and distance. In three different experiments (totaling 121 individuals) we used image analysis to measure body size and four wing traits among newly-emerged butterflies prior to flight trials: wing area, aspect ratio (length/width), melanism, and orange hue. Results showed that monarchs with darker orange (approaching red) wings flew longer distances than those with lighter orange wings in analyses that controlled for sex and other morphometric traits. This finding is consistent with past work showing that among wild monarchs, those sampled during the fall migration are darker in hue (redder) than non-migratory monarchs. Together, these results suggest that pigment deposition onto wing scales during metamorphosis could be linked with traits that influence flight, such as thorax muscle size, energy storage or metabolism. Our results reinforce an association between wing color and flight performance in insects that is suggested by past studies of wing melansim and seasonal polyphenism, and provide an important starting point for work focused on mechanistic links between insect movement and color.

  6. The Redder the Better: Wing Color Predicts Flight Performance in Monarch Butterflies

    PubMed Central

    Davis, Andrew K.; Chi, Jean; Bradley, Catherine; Altizer, Sonia

    2012-01-01

    The distinctive orange and black wings of monarchs (Danaus plexippus) have long been known to advertise their bitter taste and toxicity to potential predators. Recent work also showed that both the orange and black coloration of this species can vary in response to individual-level and environmental factors. Here we examine the relationship between wing color and flight performance in captive-reared monarchs using a tethered flight mill apparatus to quantify butterfly flight speed, duration and distance. In three different experiments (totaling 121 individuals) we used image analysis to measure body size and four wing traits among newly-emerged butterflies prior to flight trials: wing area, aspect ratio (length/width), melanism, and orange hue. Results showed that monarchs with darker orange (approaching red) wings flew longer distances than those with lighter orange wings in analyses that controlled for sex and other morphometric traits. This finding is consistent with past work showing that among wild monarchs, those sampled during the fall migration are darker in hue (redder) than non-migratory monarchs. Together, these results suggest that pigment deposition onto wing scales during metamorphosis could be linked with traits that influence flight, such as thorax muscle size, energy storage or metabolism. Our results reinforce an association between wing color and flight performance in insects that is suggested by past studies of wing melansim and seasonal polyphenism, and provide an important starting point for work focused on mechanistic links between insect movement and color. PMID:22848463

  7. Mutualistic mimicry and filtering by altitude shape the structure of Andean butterfly communities.

    PubMed

    Chazot, Nicolas; Willmott, Keith R; Santacruz Endara, Paola G; Toporov, Alexandre; Hill, Ryan I; Jiggins, Chris D; Elias, Marianne

    2014-01-01

    Both the abiotic environment and abiotic interactions among species contribute to shaping species assemblages. While the roles of habitat filtering and competitive interactions are clearly established, less is known about how positive interactions, whereby species benefit from the presence of one another, affect community structure. Here we assess the importance of positive interactions by studying Andean communities of butterflies that interact mutualistically via Müllerian mimicry. We show that communities at similar altitudes have a similar phylogenetic composition, confirming that filtering by altitude is an important process. We also provide evidence that species that interact mutualistically (i.e., species that share the same mimicry wing pattern) coexist at large scales more often than expected by chance. Furthermore, we detect an association between mimicry structure and altitude that is stronger than expected even when phylogeny is corrected for, indicating adaptive convergence for wing pattern and/or altitudinal range driven by mutualistic interactions. Positive interactions extend far beyond Müllerian mimicry, with many examples in plants and animals, and their role in the evolution and assembly of communities may be more pervasive than is currently appreciated. Our findings have strong implications for the evolution and resilience of community structure in a changing world.

  8. Biomimicry of optical microstructures of Papilio palinurus

    NASA Astrophysics Data System (ADS)

    Crne, Matija; Sharma, Vivek; Blair, John; Park, Jung Ok; Summers, Christopher J.; Srinivasarao, Mohan

    2011-01-01

    The brilliant coloration of animals in nature is sometimes based on their structure rather than on pigments. The green colour on the wings of a butterfly Papilio palinurus originates from the hierarchical microstructure of individual wing scales that are tiled on the wing. The hierarchical structure gives rise to two coloured reflections of visible light, blue and yellow which when additively mixed, produce the perception of green colour on the wing scales. We used breath figure templated assembly as the starting point for the structure and, combining it with atomic layer deposition for the multilayers necessary for the production of interference colors, we have faithfully mimicked the structure and the optical effects found on the wing scale of the butterfly Papilio palinurus.

  9. Reflection characterization of nano-sized dielectric structure in Morpho butterfly wings

    NASA Astrophysics Data System (ADS)

    Zhu, Dong

    2017-10-01

    Morpho butterflies living in Central and South America are well-known for their structural-colored blue wings. The blue coloring originates from the interaction of light with nano-sized dielectric structures that are equipped on the external surface of scales covering over their wings. The high-accuracy nonstandard finite-difference time domain (NS-FDTD) method is used to investigate the reflection characterization from the nanostructures. In the NS-FDTD calculation, a computational model is built to mimic the actual tree-like multilayered structures wherever possible using the hyperbolic tangent functions. It is generally known that both multilayer interference and diffraction grating phenomena can occur when light enters the nano-sized multilayered structure. To answer the question that which phenomenon is mainly responsible for the blue coloring, the NS-FDTD calculation is performed under various incidence angles at wavelengths from 360 to 500 nm. The calculated results at one incident wavelength under different incidence angles are visualized in a two-dimensional mapping image, where horizontal and vertical axes are incidence and reflection angles, respectively. The images demonstrate a remarkable transition from a ring-like pattern at shorter wavelengths to a retro-reflection pattern at longer wavelengths. To clarify the origin of the pattern transition, the model is separated into several simpler parts and compared their mapping images with the theoretical diffraction calculations. It can be concluded that the blue coloring at longer wavelengths is mainly caused by the cooperation of multilayer interference and retro-reflection while the effect of diffraction grating is predominant at shorter wavelengths.

  10. A BABCOCK–LEIGHTON SOLAR DYNAMO MODEL WITH MULTI-CELLULAR MERIDIONAL CIRCULATION IN ADVECTION- AND DIFFUSION-DOMINATED REGIMES

    DOE Office of Scientific and Technical Information (OSTI.GOV)

    Belucz, Bernadett; Forgács-Dajka, Emese; Dikpati, Mausumi, E-mail: bbelucz@astro.elte.hu, E-mail: dikpati@ucar.edu

    Babcock–Leighton type-solar dynamo models with single-celled meridional circulation are successful in reproducing many solar cycle features. Recent observations and theoretical models of meridional circulation do not indicate a single-celled flow pattern. We examine the role of complex multi-cellular circulation patterns in a Babcock–Leighton solar dynamo in advection- and diffusion-dominated regimes. We show from simulations that the presence of a weak, second, high-latitude reverse cell speeds up the cycle and slightly enhances the poleward branch in the butterfly diagram, whereas the presence of a second cell in depth reverses the tilt of the butterfly wing to an antisolar type. A butterflymore » diagram constructed from the middle of convection zone yields a solar-like pattern, but this may be difficult to realize in the Sun because of magnetic buoyancy effects. Each of the above cases behaves similarly in higher and lower magnetic diffusivity regimes. However, our dynamo with a meridional circulation containing four cells in latitude behaves distinctly differently in the two regimes, producing solar-like butterfly diagrams with fast cycles in the higher diffusivity regime, and complex branches in butterfly diagrams in the lower diffusivity regime. We also find that dynamo solutions for a four-celled pattern, two in radius and two in latitude, prefer to quickly relax to quadrupolar parity if the bottom flow speed is strong enough, of similar order of magnitude as the surface flow speed.« less

  11. Controlled replication of butterfly wings for achieving tunable photonic properties.

    PubMed

    Huang, Jingyun; Wang, Xudong; Wang, Zhong Lin

    2006-10-01

    The fine structure of the wing scale of a Morpho Peleides butterfly was examined carefully, and the entire configuration was completely replicated by a uniform Al(2)O(3) coating through a low-temperature ALD process. An inverted structure was achieved by removing the butterfly wing template at high temperature, forming a polycrystalline Al(2)O(3) shell structure with precisely controlled thickness. Other than the copy of the morphology of the structure, the optical property, such as the existence of PBG, was also inherited by the alumina replica. Reflection peaks at the violet/blue range were detected on both original wings and their replica, while a simple alumina coating shifted the reflection peak to longer wavelength because of the change of periodicity and refraction index. The alumina replicas also exhibited similar functional structures as waveguide and beam splitter, which may be used as the building blocks for photonic ICs with high reproducibility and lower fabrication cost compared to traditional lithography techniques.

  12. Shift from bird to butterfly pollination in Clivia (Amaryllidaceae).

    PubMed

    Kiepiel, Ian; Johnson, Steven D

    2014-01-01

    Pollinator shifts have been implicated as a driver of divergence in angiosperms. We tested the hypothesis that there was a transition from bird- to butterfly pollination in the African genus Clivia (Amaryllidaceae) and investigated how floral traits may have been either modified or retained during this transition. We identified pollinators using field observations, correlations between lepidopteran wing scales and pollen on stigmas, and single-visit and selective exclusion experiments. We also quantified floral rewards and advertising traits. The upright trumpet-shaped flowers of C. miniata were found to be pollinated effectively by swallowtail butterflies during both nectar-feeding and brush visits. These butterflies transfer pollen on their wings, as evidenced by positive correlations between wing scales and pollen loads on stigmas. All other Clivia species have narrow pendulous flowers that are visited by sunbirds. Selective exclusion of birds and large butterflies from flowers of two Clivia species resulted in a significant decline in seed production. From the distribution of pollination systems on available phylogenies, it is apparent that a shift took place from bird- to butterfly pollination in Clivia. This shift was accompanied by the evolution of trumpet-shaped flowers, smaller nectar volume, and emission of scent, while flower color and nectar chemistry do not appear to have been substantially modified. These results are consistent with the idea that pollinator shifts can explain major floral modifications during plant diversification.

  13. Identification and biosynthesis of novel male specific esters in the wings of the tropical butterfly, Bicyclus martius sanaos.

    PubMed

    Wang, Hong-Lei; Brattström, Oskar; Brakefield, Paul M; Francke, Wittko; Löfstedt, Christer

    2014-06-01

    Representatives of the highly speciose tropical butterfly genus Bicyclus (Lepidoptera: Nymphalidae) are characterized by morphological differences in the male androconia, a set of scales and hair pencils located on the surface of the wings. These androconia are assumed to be associated with the release of courtship pheromones. In the present study, we report the identification and biosynthetic pathways of several novel esters from the wings of male B. martius sanaos. We found that the volatile compounds in this male butterfly were similar to female-produced moth sex pheromones. Components associated with the male wing androconial areas were identified as ethyl, isobutyl and 2-phenylethyl hexadecanoates and (11Z)-11-hexadecenoates, among which the latter are novel natural products. By topical application of deuterium-labelled fatty acid and amino acid precursors, we found these pheromone candidates to be produced in patches located on the forewings of the males. Deuterium labels from hexadecanoic acid were incorporated into (11Z)-11-hexadecenoic acid, providing experimental evidence of a Δ11-desaturase being active in butterflies. This unusual desaturase was found previously to be involved in the biosynthesis of female-produced sex pheromones of moths. In the male butterflies, both hexadecanoic acid and (11Z)-11-hexadecenoic acid were then enzymatically esterified to form the ethyl, isobutyl and 2-phenylethyl esters, incorporating ethanol, isobutanol, and 2-phenylethanol, derived from the corresponding amino acids L-alanine, L-valine, and L-phenylalanine.

  14. Mimicking the colourful wing scale structure of the Papilio blumei butterfly.

    PubMed

    Kolle, Mathias; Salgard-Cunha, Pedro M; Scherer, Maik R J; Huang, Fumin; Vukusic, Pete; Mahajan, Sumeet; Baumberg, Jeremy J; Steiner, Ullrich

    2010-07-01

    The brightest and most vivid colours in nature arise from the interaction of light with surfaces that exhibit periodic structure on the micro- and nanoscale. In the wings of butterflies, for example, a combination of multilayer interference, optical gratings, photonic crystals and other optical structures gives rise to complex colour mixing. Although the physics of structural colours is well understood, it remains a challenge to create artificial replicas of natural photonic structures. Here we use a combination of layer deposition techniques, including colloidal self-assembly, sputtering and atomic layer deposition, to fabricate photonic structures that mimic the colour mixing effect found on the wings of the Indonesian butterfly Papilio blumei. We also show that a conceptual variation to the natural structure leads to enhanced optical properties. Our approach offers improved efficiency, versatility and scalability compared with previous approaches.

  15. Investigation of manifestation of optical properties of butterfly wings with nanoscale zinc oxide incorporation

    NASA Astrophysics Data System (ADS)

    Aideo, Swati N.; Mohanta, Dambarudhar

    2016-10-01

    In this work, microstructural and optical characteristics nanoparticles of wings of Tailed Jay (Graphium Agamemnon) butterfly were studied before and after treating it in a precursor solution of zinc acetate and ethanol. We speculate that the butterfly scales are infiltrated with ZnO nanoparticles owing to reduction of Zinc hydroxide under ambient condition. The ZnO butterfly scales so produced were characterised using optical microscopy, UV-Vis reflectance spectroscopy, and electron microscopy etc. From the reflectance spectra, we could see that after treating it in the solution, optical properties vary. We anticipate that this change may be due to the formation of ZnO nanoparticles as well as the loss in periodicity due to the chemical treatments, which could be assessed from the SEM micrographs.

  16. An introgressed wing pattern acts as a mating cue.

    PubMed

    Sánchez, Angela P; Pardo-Diaz, Carolina; Enciso-Romero, Juan; Muñoz, Astrid; Jiggins, Chris D; Salazar, Camilo; Linares, Mauricio

    2015-06-01

    Heliconius butterflies provide good examples of both homoploid hybrid speciation and ecological speciation. In particular, examples of adaptive introgression have been detected among the subspecies of Heliconius timareta, which acquired red color pattern elements from H. melpomene. We tested whether the introgression of red wing pattern elements into H. timareta florencia might also be associated with incipient reproductive isolation (RI) from its close relative, H. timareta subsp. nov., found in the eastern Andes. No choice experiments show a 50% reduction in mating between females of H. t. subsp. nov. and males of H .t. florencia, but not in the reciprocal direction. In choice experiments using wing models, males of H. timareta subsp. nov. approach and court red phenotypes less than their own, whereas males of H. t. florencia prefer models with a red phenotype. Intrinsic postzygotic isolation was not detected in crosses between these H. timareta races. These results suggest that a color pattern trait gained by introgression is triggering RI between H. timareta subsp. nov. and H. t. florencia. © 2015 The Author(s). Evolution © 2015 The Society for the Study of Evolution.

  17. AY 2002-2003 Industry Study: Final Report Strategic Materials

    DTIC Science & Technology

    2003-01-01

    Morphing Wing Project;[38] 2) an environmentally friendly, allergy-free fiber for vehicle seats patterned after the light reflection capability of the...Amazon butterfly ;[39] and 3) a camouflage net that changes color in response to its environment inspired by observations of the cuttlefish.[40] These...Biotechnology. The application of molecular level nanocircuits and nanomaterials has great potential in many areas. However, it will be

  18. Positive selection of a duplicated UV-sensitive visual pigment coincides with wing pigment evolution in Heliconius butterflies

    PubMed Central

    Briscoe, Adriana D.; Bybee, Seth M.; Bernard, Gary D.; Yuan, Furong; Sison-Mangus, Marilou P.; Reed, Robert D.; Warren, Andrew D.; Llorente-Bousquets, Jorge; Chiao, Chuan-Chin

    2010-01-01

    The butterfly Heliconius erato can see from the UV to the red part of the light spectrum with color vision proven from 440 to 640 nm. Its eye is known to contain three visual pigments, rhodopsins, produced by an 11-cis-3-hydroxyretinal chromophore together with long wavelength (LWRh), blue (BRh) and UV (UVRh1) opsins. We now find that H. erato has a second UV opsin mRNA (UVRh2)—a previously undescribed duplication of this gene among Lepidoptera. To investigate its evolutionary origin, we screened eye cDNAs from 14 butterfly species in the subfamily Heliconiinae and found both copies only among Heliconius. Phylogeny-based tests of selection indicate positive selection of UVRh2 following duplication, and some of the positively selected sites correspond to vertebrate visual pigment spectral tuning residues. Epi-microspectrophotometry reveals two UV-absorbing rhodopsins in the H. erato eye with λmax = 355 nm and 398 nm. Along with the additional UV opsin, Heliconius have also evolved 3-hydroxy-DL-kynurenine (3-OHK)-based yellow wing pigments not found in close relatives. Visual models of how butterflies perceive wing color variation indicate this has resulted in an expansion of the number of distinguishable yellow colors on Heliconius wings. Functional diversification of the UV-sensitive visual pigments may help explain why the yellow wing pigments of Heliconius are so colorful in the UV range compared to the yellow pigments of close relatives lacking the UV opsin duplicate. PMID:20133601

  19. Positive selection of a duplicated UV-sensitive visual pigment coincides with wing pigment evolution in Heliconius butterflies.

    PubMed

    Briscoe, Adriana D; Bybee, Seth M; Bernard, Gary D; Yuan, Furong; Sison-Mangus, Marilou P; Reed, Robert D; Warren, Andrew D; Llorente-Bousquets, Jorge; Chiao, Chuan-Chin

    2010-02-23

    The butterfly Heliconius erato can see from the UV to the red part of the light spectrum with color vision proven from 440 to 640 nm. Its eye is known to contain three visual pigments, rhodopsins, produced by an 11-cis-3-hydroxyretinal chromophore together with long wavelength (LWRh), blue (BRh) and UV (UVRh1) opsins. We now find that H. erato has a second UV opsin mRNA (UVRh2)-a previously undescribed duplication of this gene among Lepidoptera. To investigate its evolutionary origin, we screened eye cDNAs from 14 butterfly species in the subfamily Heliconiinae and found both copies only among Heliconius. Phylogeny-based tests of selection indicate positive selection of UVRh2 following duplication, and some of the positively selected sites correspond to vertebrate visual pigment spectral tuning residues. Epi-microspectrophotometry reveals two UV-absorbing rhodopsins in the H. erato eye with lambda(max) = 355 nm and 398 nm. Along with the additional UV opsin, Heliconius have also evolved 3-hydroxy-DL-kynurenine (3-OHK)-based yellow wing pigments not found in close relatives. Visual models of how butterflies perceive wing color variation indicate this has resulted in an expansion of the number of distinguishable yellow colors on Heliconius wings. Functional diversification of the UV-sensitive visual pigments may help explain why the yellow wing pigments of Heliconius are so colorful in the UV range compared to the yellow pigments of close relatives lacking the UV opsin duplicate.

  20. Modularity of a leaf moth-wing pattern and a versatile characteristic of the wing-pattern ground plan

    PubMed Central

    2013-01-01

    Background One of the most intriguing questions in evolutionary developmental biology is how an insect acquires a mimicry pattern within its body parts. A striking example of pattern mimicry is found in the pattern diversity of moth and butterfly wings, which is thought to evolve from preexisting elements illustrated by the nymphalid ground plan (NGP). Previous studies demonstrated that individuality of the NGP facilitates the decoupling of associated common elements, leading to divergence. In contrast, recent studies on the concept of modularity have argued the importance of a combination of coupling and decoupling of the constituent elements. Here, we examine the modularity of a mimicry wing pattern in a moth and explore an evolvable characteristic of the NGP. Results This study examined the wings of the noctuid moth Oraesia excavata, which closely resemble leaves with a leaf venation pattern. Based on a comparative morphological procedure, we found that this leaf pattern was formed by the NGP common elements. Using geometric morphometrics combined with network analysis, we found that each of the modules in the leaf pattern integrates the constituent components of the leaf venation pattern (i.e., the main and lateral veins). Moreover, the detected modules were established by coupling different common elements and decoupling even a single element into different modules. The modules of the O. excavata wing pattern were associated with leaf mimicry, not with the individuality of the NGP common elements. For comparison, we also investigated the modularity of a nonmimetic pattern in the noctuid moth Thyas juno. Quantitative analysis demonstrated that the modules of the T. juno wing pattern regularly corresponded to the individuality of the NGP common elements, unlike those in the O. excavata wing pattern. Conclusions This study provides the first evidence for modularity in a leaf mimicry pattern. The results suggest that the evolution of this pattern involves coupling and decoupling processes to originate these modules, free from the individuality of the NGP system. We propose that this evolution has been facilitated by a versatile characteristic of the NGP, allowing the association of freely modifiable subordinate common elements to make modules. PMID:23890367

  1. Wing Scale Orientation Alters Reflection Directions in the Green Hairstreak Chrysozephyrus smaragdinus (Lycaenidae; Lepidoptera).

    PubMed

    Imafuku, Michio; Ogihara, Naomichi

    2016-12-01

    There have been only a few reports on the directional reflection of light by butterfly wings. Here, we systematically investigated this phenomenon in a lycaenid butterfly, Chrysozephyrus smaragdinus,in which males have bright green wings based on structural coloration. We used a device that measures intensities of light in hemispherical space by vertical shifting of a sensor and horizontal rotation of the stage carrying the wing, which is illuminated from the top, to determine the direction of light reflected by the fore- and hindwings. The orientation and curvature of wing scales were also examined microscopically. The forewing of this species reflected light shone from the top largely forward, whereas the hindwing reflected it slightly forward. This difference was attributed to the tilt angles of the wing scales. Light reflection by the forewing was relatively weak, and widely scattered, whereas that by the hindwing was rather concentrated, resulting in higher reflectance. This difference was attributed to difference in the curvature of the wing scales on the two wings.

  2. Fabrication and structure characterization of te butterfly nanostructures.

    PubMed

    Wong, Tailun; She, Guangwei; Cheng, Chun; Li, Wei; Shi, Wensheng; Zhang, Xiaohong; Wang, Ning

    2011-12-01

    Te nanowires and butterfly nanostructures have been fabricated by template-free electrodeposition (TFED) in aqueous solution. By high-resolution transmission electron microscopy (HRTEM) study, the favored growth directions of the nanowires and the wings of the butterfly nanostructures were determined to be along the [0001] direction of trigonal Te, and the twinning plane of the butterfly nanostructures was (11-22). The cathodoluminescence measurements carried out at different positions of the butterfly nanostructure indicated that the twin boundaries influenced the photoemission efficiency.

  3. A Model for Selection of Eyespots on Butterfly Wings.

    PubMed

    Sekimura, Toshio; Venkataraman, Chandrasekhar; Madzvamuse, Anotida

    2015-01-01

    The development of eyespots on the wing surface of butterflies of the family Nympalidae is one of the most studied examples of biological pattern formation.However, little is known about the mechanism that determines the number and precise locations of eyespots on the wing. Eyespots develop around signaling centers, called foci, that are located equidistant from wing veins along the midline of a wing cell (an area bounded by veins). A fundamental question that remains unsolved is, why a certain wing cell develops an eyespot, while other wing cells do not. We illustrate that the key to understanding focus point selection may be in the venation system of the wing disc. Our main hypothesis is that changes in morphogen concentration along the proximal boundary veins of wing cells govern focus point selection. Based on previous studies, we focus on a spatially two-dimensional reaction-diffusion system model posed in the interior of each wing cell that describes the formation of focus points. Using finite element based numerical simulations, we demonstrate that variation in the proximal boundary condition is sufficient to robustly select whether an eyespot focus point forms in otherwise identical wing cells. We also illustrate that this behavior is robust to small perturbations in the parameters and geometry and moderate levels of noise. Hence, we suggest that an anterior-posterior pattern of morphogen concentration along the proximal vein may be the main determinant of the distribution of focus points on the wing surface. In order to complete our model, we propose a two stage reaction-diffusion system model, in which an one-dimensional surface reaction-diffusion system, posed on the proximal vein, generates the morphogen concentrations that act as non-homogeneous Dirichlet (i.e., fixed) boundary conditions for the two-dimensional reaction-diffusion model posed in the wing cells. The two-stage model appears capable of generating focus point distributions observed in nature. We therefore conclude that changes in the proximal boundary conditions are sufficient to explain the empirically observed distribution of eyespot focus points on the entire wing surface. The model predicts, subject to experimental verification, that the source strength of the activator at the proximal boundary should be lower in wing cells in which focus points form than in those that lack focus points. The model suggests that the number and locations of eyespot foci on the wing disc could be largely controlled by two kinds of gradients along two different directions, that is, the first one is the gradient in spatially varying parameters such as the reaction rate along the anterior-posterior direction on the proximal boundary of the wing cells, and the second one is the gradient in source values of the activator along the veins in the proximal-distal direction of the wing cell.

  4. Imaging scatterometry and microspectrophotometry of lycaenid butterfly wing scales with perforated multilayers

    PubMed Central

    Wilts, Bodo D.; Leertouwer, Hein L.; Stavenga, Doekele G.

    2008-01-01

    We studied the structural as well as spatial and spectral reflectance characteristics of the wing scales of lycaenid butterfly species, where the scale bodies consist of perforated multilayers. The extent of the spatial scattering profiles was measured with a newly built scatterometer. The width of the reflectance spectra, measured with a microspectrophotometer, decreased with the degree of perforation, in agreement with the calculations based on multilayer theory. PMID:18782721

  5. Rice- and butterfly-wing effect inspired self-cleaning and low drag micro/nanopatterned surfaces in water, oil, and air flow.

    PubMed

    Bixler, Gregory D; Bhushan, Bharat

    2014-01-07

    In search of new solutions to complex challenges, researchers are turning to living nature for inspiration. For example, special surface characteristics of rice leaves and butterfly wings combine the shark skin (anisotropic flow leading to low drag) and lotus leaf (superhydrophobic and self-cleaning) effects, producing the so-called rice and butterfly wing effect. In this paper, we study four microstructured surfaces inspired by rice leaves and fabricated with photolithography techniques. We also present a method of creating such surfaces using a hot embossing procedure for scaled-up manufacturing. Fluid drag, self-cleaning, contact angle, and contact angle hysteresis data are presented to understand the role of sample geometrical dimensions. Conceptual modeling provides design guidance when developing novel low drag, self-cleaning, and potentially antifouling surfaces for medical, marine, and industrial applications.

  6. Bioinspired Au/TiO2 photocatalyst derived from butterfly wing (Papilio Paris).

    PubMed

    Chen, Jianjun; Su, Huilan; Song, Fang; Moon, Won-Jin; Kim, Yang-Soo; Zhang, Di

    2012-03-15

    The reticular hierarchical structure of butterfly wings (Papilio Paris) is introduced as template for Au/TiO(2) photocatalyst by depositing the Au nanoparticles on TiO(2) matrix, which is carried out by a water-ethanol sol-gel procedure combined with subsequent calcination. The obtained Au/TiO(2) nanocomposites present the reticular hierarchical structure of butterfly wings, and Au nanoparticles with an average size of 7 nm are homogeneously dispersed in TiO(2) substrate. Benefiting from such unique reticular hierarchical structure and composition, the biomorphic Au/TiO(2) exhibits high-harvesting capability and presents superior photocatalytic activity. Especially, the biomorphic Au/TiO(2) at the nominal content of gold to titanium of 8 wt% shows the highest photocatalytic activity and can completely decompose methyl orange within 80 min, which is obviously higher than that of commercial Degussa P25 powders. Copyright © 2011 Elsevier Inc. All rights reserved.

  7. Rice- and butterfly-wing effect inspired self-cleaning and low drag micro/nanopatterned surfaces in water, oil, and air flow

    NASA Astrophysics Data System (ADS)

    Bixler, Gregory D.; Bhushan, Bharat

    2013-12-01

    In search of new solutions to complex challenges, researchers are turning to living nature for inspiration. For example, special surface characteristics of rice leaves and butterfly wings combine the shark skin (anisotropic flow leading to low drag) and lotus leaf (superhydrophobic and self-cleaning) effects, producing the so-called rice and butterfly wing effect. In this paper, we study four microstructured surfaces inspired by rice leaves and fabricated with photolithography techniques. We also present a method of creating such surfaces using a hot embossing procedure for scaled-up manufacturing. Fluid drag, self-cleaning, contact angle, and contact angle hysteresis data are presented to understand the role of sample geometrical dimensions. Conceptual modeling provides design guidance when developing novel low drag, self-cleaning, and potentially antifouling surfaces for medical, marine, and industrial applications.

  8. Pretreated Butterfly Wings for Tuning the Selective Vapor Sensing.

    PubMed

    Piszter, Gábor; Kertész, Krisztián; Bálint, Zsolt; Biró, László Péter

    2016-09-07

    Photonic nanoarchitectures occurring in the scales of Blue butterflies are responsible for their vivid blue wing coloration. These nanoarchitectures are quasi-ordered nanocomposites which are constituted from a chitin matrix with embedded air holes. Therefore, they can act as chemically selective sensors due to their color changes when mixing volatile vapors in the surrounding atmosphere which condensate into the nanoarchitecture through capillary condensation. Using a home-built vapor-mixing setup, the spectral changes caused by the different air + vapor mixtures were efficiently characterized. It was found that the spectral shift is vapor-specific and proportional with the vapor concentration. We showed that the conformal modification of the scale surface by atomic layer deposition and by ethanol pretreatment can significantly alter the optical response and chemical selectivity, which points the way to the efficient production of sensor arrays based on the knowledge obtained through the investigation of modified butterfly wings.

  9. Pretreated Butterfly Wings for Tuning the Selective Vapor Sensing

    PubMed Central

    Piszter, Gábor; Kertész, Krisztián; Bálint, Zsolt; Biró, László Péter

    2016-01-01

    Photonic nanoarchitectures occurring in the scales of Blue butterflies are responsible for their vivid blue wing coloration. These nanoarchitectures are quasi-ordered nanocomposites which are constituted from a chitin matrix with embedded air holes. Therefore, they can act as chemically selective sensors due to their color changes when mixing volatile vapors in the surrounding atmosphere which condensate into the nanoarchitecture through capillary condensation. Using a home-built vapor-mixing setup, the spectral changes caused by the different air + vapor mixtures were efficiently characterized. It was found that the spectral shift is vapor-specific and proportional with the vapor concentration. We showed that the conformal modification of the scale surface by atomic layer deposition and by ethanol pretreatment can significantly alter the optical response and chemical selectivity, which points the way to the efficient production of sensor arrays based on the knowledge obtained through the investigation of modified butterfly wings. PMID:27618045

  10. Lithographically-generated 3D lamella layers and their structural color

    NASA Astrophysics Data System (ADS)

    Zhang, Sichao; Chen, Yifang; Lu, Bingrui; Liu, Jianpeng; Shao, Jinhai; Xu, Chen

    2016-04-01

    Inspired by the structural color from the multilayer nanophotonic structures in Morpho butterfly wing scales, 3D lamellae layers in dielectric polymers (polymethyl methacrylate, PMMA) with n ~ 1.5 were designed and fabricated by standard top-down electron beam lithography with one-step exposure followed by an alternating development/dissolution process of PMMA/LOR (lift-off resist) multilayers. This work offers direct proof of the structural blue/green color via lithographically-replicated PMMA/air multilayers, analogous to those in real Morpho butterfly wings. The success of nanolithography in this work for the 3D lamellae structures in dielectric polymers not only enables us to gain deeper insight into the mysterious blue color of the Morpho butterfly wings, but also breaks through the bottleneck in technical development toward broad applications in gas/liquid sensors, 3D meta-materials, coloring media, and infrared imaging devices, etc.

  11. Pitching motion control of a butterfly-like 3D flapping wing-body model

    NASA Astrophysics Data System (ADS)

    Suzuki, Kosuke; Minami, Keisuke; Inamuro, Takaji

    2014-11-01

    Free flights and a pitching motion control of a butterfly-like flapping wing-body model are numerically investigated by using an immersed boundary-lattice Boltzmann method. The model flaps downward for generating the lift force and backward for generating the thrust force. Although the model can go upward against the gravity by the generated lift force, the model generates the nose-up torque, consequently gets off-balance. In this study, we discuss a way to control the pitching motion by flexing the body of the wing-body model like an actual butterfly. The body of the model is composed of two straight rigid rod connected by a rotary actuator. It is found that the pitching angle is suppressed in the range of +/-5° by using the proportional-plus-integral-plus-derivative (PID) control for the input torque of the rotary actuator.

  12. Reverse color sequence in the diffraction of white light by the wing of the male butterfly Pierella luna (Nymphalidae: Satyrinae)

    NASA Astrophysics Data System (ADS)

    Vigneron, Jean Pol; Simonis, Priscilla; Aiello, Annette; Bay, Annick; Windsor, Donald M.; Colomer, Jean-François; Rassart, Marie

    2010-08-01

    The butterfly Pierella luna (Nymphalidae) shows an intriguing rainbow iridescence effect: the forewings of the male, when illuminated along the axis from the body to the wing tip, decompose a white light beam as a diffraction grating would do. Violet light, however, emerges along a grazing angle, near the wing surface, while the other colors, from blue to red, exit respectively at angles progressively closer to the direction perpendicular to the wing plane. This sequence is the reverse of the usual decomposition of light by a grating with a periodicity parallel to the wing surface. It is shown that this effect is produced by a macroscopic deformation of the entire scale, which curls in such a way that it forms a “vertical” grating, perpendicular to the wing surface, and functions in transmission instead of reflection.

  13. Oceanography: A sea butterfly flaps its wings

    NASA Astrophysics Data System (ADS)

    Ries, Justin B.

    2012-12-01

    Ocean acidification is predicted to harm the ocean's shell-building organisms over the coming centuries. Sea butterflies, an ecologically important group of molluscs in the Arctic and Southern oceans, are already suffering the effects.

  14. Deimatic Display in the European Swallowtail Butterfly as a Secondary Defence against Attacks from Great Tits

    PubMed Central

    Olofsson, Martin; Eriksson, Stephan; Jakobsson, Sven; Wiklund, Christer

    2012-01-01

    Background Many animals reduce the risk of being attacked by a predator through crypsis, masquerade or, alternatively, by advertising unprofitability by means of aposematic signalling. Behavioural attributes in prey employed after discovery, however, signify the importance of also having an effective secondary defence if a predator uncovers, or is immune to, the prey’s primary defence. In butterflies, as in most animals, secondary defence generally consists of escape flights. However, some butterfly species have evolved other means of secondary defence such as deimatic displays/startle displays. The European swallowtail, Papilio machaon, employs what appears to be a startle display by exposing its brightly coloured dorsal wing surface upon disturbance and, if the disturbance continues, by intermittently protracting and relaxing its wing muscles generating a jerky motion of the wings. This display appears directed towards predators but whether it is effective in intimidating predators so that they refrain from attacks has never been tested experimentally. Methodology/Principal Findings In this study we staged encounters between a passerine predator, the great tit, Parus major, and live and dead swallowtail butterflies in a two-choice experiment. Results showed that the dead butterfly was virtually always attacked before the live butterfly, and that it took four times longer before a bird attacked the live butterfly. When the live butterfly was approached by a bird this generally elicited the butterfly’s startle display, which usually caused the approaching bird to flee. We also performed a palatability test of the butterflies and results show that the great tits seemed to find them palatable. Conclusions/Significance We conclude that the swallowtail’s startle display of conspicuous coloration and jerky movements is an efficient secondary defence against small passerines. We also discuss under what conditions predator-prey systems are likely to aid the evolution of deimatic behaviours in harmless and palatable prey. PMID:23056590

  15. Genomic Sequence around Butterfly Wing Development Genes: Annotation and Comparative Analysis

    PubMed Central

    Conceição, Inês C.; Long, Anthony D.; Gruber, Jonathan D.; Beldade, Patrícia

    2011-01-01

    Background Analysis of genomic sequence allows characterization of genome content and organization, and access beyond gene-coding regions for identification of functional elements. BAC libraries, where relatively large genomic regions are made readily available, are especially useful for species without a fully sequenced genome and can increase genomic coverage of phylogenetic and biological diversity. For example, no butterfly genome is yet available despite the unique genetic and biological properties of this group, such as diversified wing color patterns. The evolution and development of these patterns is being studied in a few target species, including Bicyclus anynana, where a whole-genome BAC library allows targeted access to large genomic regions. Methodology/Principal Findings We characterize ∼1.3 Mb of genomic sequence around 11 selected genes expressed in B. anynana developing wings. Extensive manual curation of in silico predictions, also making use of a large dataset of expressed genes for this species, identified repetitive elements and protein coding sequence, and highlighted an expansion of Alcohol dehydrogenase genes. Comparative analysis with orthologous regions of the lepidopteran reference genome allowed assessment of conservation of fine-scale synteny (with detection of new inversions and translocations) and of DNA sequence (with detection of high levels of conservation of non-coding regions around some, but not all, developmental genes). Conclusions The general properties and organization of the available B. anynana genomic sequence are similar to the lepidopteran reference, despite the more than 140 MY divergence. Our results lay the groundwork for further studies of new interesting findings in relation to both coding and non-coding sequence: 1) the Alcohol dehydrogenase expansion with higher similarity between the five tandemly-repeated B. anynana paralogs than with the corresponding B. mori orthologs, and 2) the high conservation of non-coding sequence around the genes wingless and Ecdysone receptor, both involved in multiple developmental processes including wing pattern formation. PMID:21909358

  16. VizieR Online Data Catalog: Butterfly diagram wings (Leussu+, 2017)

    NASA Astrophysics Data System (ADS)

    Leussu, R.; Usoskin, I. G.; Senthamizh Pavai, V.; Diercke, A.; Arlt, R.; Mursula, K.

    2016-11-01

    fig1data.dat contains the separated wings in a butterfly diagram for sunspot groups from three different origins: Sunspot observations by S.H. Schwabe and G. Spoerer, and the RGO/SOON compilation. The latitudes for sunspot groups from the Schwabe and Spoerer data are given as size-weighted averages from sunspots belonging to each group. Latitudes for the RGO compilation are given as they are stated in the original data. The columns report the year, month, day, date [yr], latitude [deg], cycle, hemisphere, and data set tag. Northern hemisphere wings are tagged with "1" and southern hemisphere wings with "2". The data set tag is "1" for Schwabe data, "2" for Spoerer data and "3" for RGO data. (1 data file).

  17. A Babcock-Leighton solar dynamo model with multi-cellular meridional circulation in advection- and diffusion-dominated regimes

    NASA Astrophysics Data System (ADS)

    Belucz, B.; Dikpati, M.; Forgacs-Dajka, E.

    2014-12-01

    Babcock-Leighton type solar dynamo models with single cell meridional circulation are successful in reproducing many solarcycle features, and recently such a model was applied for solarcycle 24 amplitude prediction. It seems that cycle 24 amplitudeforecast may not be validated. One of the reasons is the assumption of a single cell meridional circulation. Recent observations andtheoretical models of meridional circulation do not indicate a single-celledflow pattern. So it is nessecary to examine the role of complexmulti-cellular circulation patterns in a Babcock-Leighton solar dynamo model in the advection and diffusion dominated regimes.By simulating a Babcock-Leighton solar dynamo model with multi-cellularflow, we show that the presence of a weak, second, high-latitudereverse cell speeds up the cycle and slighty enhances the poleward branch in the butterfly diagram, whereas the presence of a second cellin depth reverses the tilt of the butterfly wing and leads to ananti-solar type feature. If, instead, the butterfly diagram isconstructed from the middle of the convection zone in that case, a solar-like pattern can be retrieved. All the above cases behavequalitatively similar in advection and diffusion-dominated regimes.However, our dynamo with a meridional circulation containing fourcells in latitude behaves distinctly different in the two regimes, producing a solar-like butterfly diagram with fast cycles indiffusion-dominated regime, and a complex branches in the butterflydiagram in the advection-dominated regime. Another interestingfinding from our studies is that a four-celled flow pattern containing two in radius and two in latitude always producesquadrupolar parity as the relaxed solution.

  18. Exploring structural colour in uni- and multi-coloured butterfly wings and Ag+ uptake by scales

    NASA Astrophysics Data System (ADS)

    Aideo, Swati N.; Haloi, Rajib; Mohanta, Dambarudhar

    2017-09-01

    We discuss the origin of the structural colour of different butterfly wings in the light of the typical built-in microstructural arrangement of scales that are comprised of chitin-melanin layer and air-gaps. Three specimens of butterfly wings namely, Papilio Liomedon (black), Catopsilia Pyranthe (light green) and Vanessa Cardui (multi-coloured) were chosen and diffuse reflectance characteristics have been aquired for normal incidence of p-polarized light. Moreover, the time-dependent uptake of Ag+ into scales has led to swelling and spread of the chitinous ridges and ribs, with revelation of micro-beads in Catopsilia Pyranthe specimen. The reduction of the number of air-gaps between any two parallel ridges is attributed to the merging of adjacent gaps possessing a common boundary. The availability of Ag at the centre of a chosen ridge, for every wing type, follows an exponential growing trend, ∼e0.36t . Precise inclusion of nanoscale metals into natural photonic systems would provide new insight, while applying principles of photonics and plasmonics simultaneously.

  19. Environmental effects on the shape variation of male ultraviolet patterns in the Brimstone butterfly ( Gonepteryx rhamni, Pieridae, Lepidoptera)

    NASA Astrophysics Data System (ADS)

    Pecháček, Pavel; Stella, David; Keil, Petr; Kleisner, Karel

    2014-12-01

    The males of the Brimstone butterfly ( Gonepteryx rhamni) have ultraviolet pattern on the dorsal surfaces of their wings. Using geometric morphometrics, we have analysed correlations between environmental variables (climate, productivity) and shape variability of the ultraviolet pattern and the forewing in 110 male specimens of G. rhamni collected in the Palaearctic zone. To start with, we subjected the environmental variables to principal component analysis (PCA). The first PCA axis (precipitation, temperature, latitude) significantly correlated with shape variation of the ultraviolet patterns across the Palaearctic. Additionally, we have performed two-block partial least squares (PLS) analysis to assess co-variation between intraspecific shape variation and the variation of 11 environmental variables. The first PLS axis explained 93 % of variability and represented the effect of precipitation, temperature and latitude. Along this axis, we observed a systematic increase in the relative area of ultraviolet colouration with increasing temperature and precipitation and decreasing latitude. We conclude that the shape variation of ultraviolet patterns on the forewings of male Brimstones is correlated with large-scale environmental factors.

  20. Hybridization Reveals the Evolving Genomic Architecture of Speciation

    PubMed Central

    Kronforst, Marcus R.; Hansen, Matthew E.B.; Crawford, Nicholas G.; Gallant, Jason R.; Zhang, Wei; Kulathinal, Rob J.; Kapan, Durrell D.; Mullen, Sean P.

    2014-01-01

    SUMMARY The rate at which genomes diverge during speciation is unknown, as are the physical dynamics of the process. Here, we compare full genome sequences of 32 butterflies, representing five species from a hybridizing Heliconius butterfly community, to examine genome-wide patterns of introgression and infer how divergence evolves during the speciation process. Our analyses reveal that initial divergence is restricted to a small fraction of the genome, largely clustered around known wing-patterning genes. Over time, divergence evolves rapidly, due primarily to the origin of new divergent regions. Furthermore, divergent genomic regions display signatures of both selection and adaptive introgression, demonstrating the link between microevolutionary processes acting within species and the origin of species across macroevolutionary timescales. Our results provide a uniquely comprehensive portrait of the evolving species boundary due to the role that hybridization plays in reducing the background accumulation of divergence at neutral sites. PMID:24183670

  1. Reproduction and optical analysis of Morpho-inspired polymeric nanostructures

    NASA Astrophysics Data System (ADS)

    Tippets, Cary A.; Fu, Yulan; Jackson, Anne-Martine; Donev, Eugenii U.; Lopez, Rene

    2016-06-01

    The brilliant blue coloration of the Morpho rhetenor butterfly originates from complex nanostructures found on the surface of its wings. The Morpho butterfly exhibits strong short-wavelength reflection and a unique two-lobe optical signature in the incident (θ) and reflected (ϕ) angular space. Here, we report the large-area fabrication of a Morpho-like structure and its reproduction in perfluoropolyether. Reflection comparisons of periodic and quasi-random ‘polymer butterfly’ nanostructures show similar normal-incidence spectra but differ in the angular θ-ϕ dependence. The periodic sample shows strong specular reflection and simple diffraction. However, the quasi-random sample produces a two-lobe angular reflection pattern with minimal specular refection, approximating the real butterfly’s optical behavior. Finite-difference time-domain simulations confirm that this pattern results from the quasi-random periodicity and highlights the significance of the inherent randomness in the Morpho’s photonic structure.

  2. Ecological impacts of tropical forest fragmentation: how consistent are patterns in species richness and nestedness?

    PubMed

    Hill, Jane K; Gray, Michael A; Khen, Chey Vun; Benedick, Suzan; Tawatao, Noel; Hamer, Keith C

    2011-11-27

    Large areas of tropical forest now exist as remnants scattered across agricultural landscapes, and so understanding the impacts of forest fragmentation is important for biodiversity conservation. We examined species richness and nestedness among tropical forest remnants in birds (meta-analysis of published studies) and insects (field data for fruit-feeding Lepidoptera (butterflies and moths) and ants). Species-area relationships were evident in all four taxa, and avian and insect assemblages in remnants typically were nested subsets of those in larger areas. Avian carnivores and nectarivores and predatory ants were more nested than other guilds, implying that the sequential loss of species was more predictable in these groups, and that fragmentation alters the trophic organization of communities. For butterflies, the ordering of fragments to achieve maximum nestedness was by fragment area, suggesting that differences among fragments were driven mainly by extinction. In contrast for moths, maximum nestedness was achieved by ordering species by wing length; species with longer wings (implying better dispersal) were more likely to occur at all sites, including low diversity sites, suggesting that differences among fragments were driven more strongly by colonization. Although all four taxa exhibited high levels of nestedness, patterns of species turnover were also idiosyncratic, and thus even species-poor sites contributed to landscape-scale biodiversity, particularly for insects.

  3. Ecological impacts of tropical forest fragmentation: how consistent are patterns in species richness and nestedness?

    PubMed Central

    Hill, Jane K.; Gray, Michael A.; Khen, Chey Vun; Benedick, Suzan; Tawatao, Noel; Hamer, Keith C.

    2011-01-01

    Large areas of tropical forest now exist as remnants scattered across agricultural landscapes, and so understanding the impacts of forest fragmentation is important for biodiversity conservation. We examined species richness and nestedness among tropical forest remnants in birds (meta-analysis of published studies) and insects (field data for fruit-feeding Lepidoptera (butterflies and moths) and ants). Species–area relationships were evident in all four taxa, and avian and insect assemblages in remnants typically were nested subsets of those in larger areas. Avian carnivores and nectarivores and predatory ants were more nested than other guilds, implying that the sequential loss of species was more predictable in these groups, and that fragmentation alters the trophic organization of communities. For butterflies, the ordering of fragments to achieve maximum nestedness was by fragment area, suggesting that differences among fragments were driven mainly by extinction. In contrast for moths, maximum nestedness was achieved by ordering species by wing length; species with longer wings (implying better dispersal) were more likely to occur at all sites, including low diversity sites, suggesting that differences among fragments were driven more strongly by colonization. Although all four taxa exhibited high levels of nestedness, patterns of species turnover were also idiosyncratic, and thus even species-poor sites contributed to landscape-scale biodiversity, particularly for insects. PMID:22006967

  4. The butterfly effect of the "butterfly effect".

    PubMed

    Dooley, Kevin J

    2009-07-01

    The "Butterfly Effect" metaphor states with variance that the flap of a butterfly's wings in Brazil can cause a tornado in Texas. This metaphor has become part of the common vernacular of Western culture. In this paper I discuss the origins of the metaphor, examine its current usage within popular culture, and present an argument as to why it is popular. I propose that the metaphor is a type of semantic attractor, a narrative device with invariant meaning but audience-specific contextualization. Finally I address whether the Butterfly Effect metaphor is a good example of itself.

  5. Order-disorder effects in structure and color relation of photonic-crystal-type nanostructures in butterfly wing scales.

    PubMed

    Márk, Géza I; Vértesy, Zofia; Kertész, Krisztián; Bálint, Zsolt; Biró, László P

    2009-11-01

    In order to study local and global order in butterfly wing scales possessing structural colors, we have developed a direct space algorithm, based on averaging the local environment of the repetitive units building up the structure. The method provides the statistical distribution of the local environments, including the histogram of the nearest-neighbor distance and the number of nearest neighbors. We have analyzed how the different kinds of randomness present in the direct space structure influence the reciprocal space structure. It was found that the Fourier method is useful in the case of a structure randomly deviating from an ordered lattice. The direct space averaging method remains applicable even for structures lacking long-range order. Based on the first Born approximation, a link is established between the reciprocal space image and the optical reflectance spectrum. Results calculated within this framework agree well with measured reflectance spectra because of the small width and moderate refractive index contrast of butterfly scales. By the analysis of the wing scales of Cyanophrys remus and Albulina metallica butterflies, we tested the methods for structures having long-range order, medium-range order, and short-range order.

  6. Order-disorder effects in structure and color relation of photonic-crystal-type nanostructures in butterfly wing scales

    NASA Astrophysics Data System (ADS)

    Márk, Géza I.; Vértesy, Zofia; Kertész, Krisztián; Bálint, Zsolt; Biró, László P.

    2009-11-01

    In order to study local and global order in butterfly wing scales possessing structural colors, we have developed a direct space algorithm, based on averaging the local environment of the repetitive units building up the structure. The method provides the statistical distribution of the local environments, including the histogram of the nearest-neighbor distance and the number of nearest neighbors. We have analyzed how the different kinds of randomness present in the direct space structure influence the reciprocal space structure. It was found that the Fourier method is useful in the case of a structure randomly deviating from an ordered lattice. The direct space averaging method remains applicable even for structures lacking long-range order. Based on the first Born approximation, a link is established between the reciprocal space image and the optical reflectance spectrum. Results calculated within this framework agree well with measured reflectance spectra because of the small width and moderate refractive index contrast of butterfly scales. By the analysis of the wing scales of Cyanophrys remus and Albulina metallica butterflies, we tested the methods for structures having long-range order, medium-range order, and short-range order.

  7. Latitudinal distribution of soft X-ray flares and dispairty in butterfly diagram

    NASA Astrophysics Data System (ADS)

    Pandey, K. K.; Yellaiah, G.; Hiremath, K. M.

    2015-04-01

    We present statistical analysis of about 63000 soft X-ray flare (class≥C) observed by geostationary operational environmental satellite (GOES) during the period 1976-2008. Class wise occurrence of soft X-ray (SXR) flare is in declining trend since cycle 21. The distribution pattern of cycle 21 shows the transit of hemispheric dominance of flare activity from northern to southern hemisphere and remains there during cycle 22 and 23. During the three cycles, 0-100, 21-300 latitude belts in southern hemisphere (SH) and 31-400 latitude belt in northern hemisphere (NH) are mightier. The 11-200 latitude belt of both hemisphere is mightiest. Correlation coefficient between consecutive latitude appears to be increasing from equator to poleward in northern hemisphere whereas pole to equatorward in southern hemisphere. Slope of the regression line fitted with asymmetry time series of daily flare counts is negative in all three cycles for different classes of flares. The yearly asymmetry curve fitted by a sinusoidal function varies from 5.6 to 11 years period and depends upon the intensity of flare. Variation, of curve fitted with wings of butterfly diagram, from first to second order polynomial suggests that latitudinal migration of flare activity varies from cycle to cycle, northern to southern hemisphere. The variation in slope of the butterfly wing of different flare class indicates the non uniform migration of flare activity.

  8. Bioinspired phase-separated disordered nanostructures for thin photovoltaic absorbers.

    PubMed

    Siddique, Radwanul H; Donie, Yidenekachew J; Gomard, Guillaume; Yalamanchili, Sisir; Merdzhanova, Tsvetelina; Lemmer, Uli; Hölscher, Hendrik

    2017-10-01

    The wings of the black butterfly, Pachliopta aristolochiae , are covered by micro- and nanostructured scales that harvest sunlight over a wide spectral and angular range. Considering that these properties are particularly attractive for photovoltaic applications, we analyze the contribution of these micro- and nanostructures, focusing on the structural disorder observed in the wing scales. In addition to microspectroscopy experiments, we conduct three-dimensional optical simulations of the exact scale structure. On the basis of these results, we design nanostructured thin photovoltaic absorbers of disordered nanoholes, which combine efficient light in-coupling and light-trapping properties together with a high angular robustness. Finally, inspired by the phase separation mechanism of self-assembled biophotonic nanostructures, we fabricate these bioinspired absorbers using a scalable, self-assembly patterning technique based on the phase separation of binary polymer mixture. The nanopatterned absorbers achieve a relative integrated absorption increase of 90% at a normal incident angle of light to as high as 200% at large incident angles, demonstrating the potential of black butterfly structures for light-harvesting purposes in thin-film solar cells.

  9. Bioinspired phase-separated disordered nanostructures for thin photovoltaic absorbers

    PubMed Central

    Siddique, Radwanul H.; Donie, Yidenekachew J.; Gomard, Guillaume; Yalamanchili, Sisir; Merdzhanova, Tsvetelina; Lemmer, Uli; Hölscher, Hendrik

    2017-01-01

    The wings of the black butterfly, Pachliopta aristolochiae, are covered by micro- and nanostructured scales that harvest sunlight over a wide spectral and angular range. Considering that these properties are particularly attractive for photovoltaic applications, we analyze the contribution of these micro- and nanostructures, focusing on the structural disorder observed in the wing scales. In addition to microspectroscopy experiments, we conduct three-dimensional optical simulations of the exact scale structure. On the basis of these results, we design nanostructured thin photovoltaic absorbers of disordered nanoholes, which combine efficient light in-coupling and light-trapping properties together with a high angular robustness. Finally, inspired by the phase separation mechanism of self-assembled biophotonic nanostructures, we fabricate these bioinspired absorbers using a scalable, self-assembly patterning technique based on the phase separation of binary polymer mixture. The nanopatterned absorbers achieve a relative integrated absorption increase of 90% at a normal incident angle of light to as high as 200% at large incident angles, demonstrating the potential of black butterfly structures for light-harvesting purposes in thin-film solar cells. PMID:29057320

  10. A Comparison of the Performance of 2D Square and Rectangular Dielectric Vein Structures

    DTIC Science & Technology

    2012-09-27

    are found in certain butterfly wings and in the exo-skeletal chitin of certain species of beetle.[3] It is photonic crystals that are believed to...difference of nearly 10:1. The second case sets 521. somewhat closer to a value for chitin , a material commonly found in butterfly wings. In future...studies,  is to be raised to 2.4, a common value for chitin .[12] DISTRIBUTION A DISTRIBUTION A 12 The reciprocal of the square dielectric

  11. Surface morphology of chitin highly related with the isolated body part of butterfly (Argynnis pandora).

    PubMed

    Kaya, Murat; Bitim, Betül; Mujtaba, Muhammad; Koyuncu, Turgay

    2015-11-01

    This study was conducted to understand the differences in the physicochemical properties of chitin samples isolated from the wings and the other body parts except the wings (OBP) of a butterfly species (Argynnis pandora). The same isolation method was used for obtaining chitin specimens from both types of body parts. The chitin content of the wings (22%) was recorded as being much higher than the OBP (8%). The extracted chitin samples were characterized via FT-IR, TGA, XRD, SEM, and elemental analysis techniques. Results of these characterizations revealed that the chitins from both structures (wings and OBP) were very similar, except for their surface morphologies. SEM results demonstrated one type of surface morphology for the wings and four different surface morphologies for the OBP. Therefore, it can be hypothesized that the surface morphology of the chitin is highly related with the body part. Copyright © 2015 Elsevier B.V. All rights reserved.

  12. The effect of butterfly-scale inspired patterning on leading-edge vortex growth

    NASA Astrophysics Data System (ADS)

    Wilroy, Jacob; Lang, Amy; Wahidi, Redha

    2014-11-01

    Leading edge vortices (LEVs) are important for generating thrust and lift in flapping flight, and the surface patterning (scales) on butterfly wings is hypothesized to play a role in the vortex formation of the LEV. To simplify this complex flow problem, we designed an experiment to focus on the alteration of 2-D vortex development with a variation in surface patterning. Specifically we are interested in the secondary vorticity generated by the LEV interacting at the patterned surface and how this can affect the growth rate of the circulation in the LEV. For this experiment we used rapid-prototyped longitudinal and transverse square grooves attached to a flat plate and compared the vortex formation as the plate moved vertically. The plate is impulsively started in quiescent water and flow fields at Re = 1500, 3000, and 6000 are examined using Digital Particle Image Velocimetry (DPIV). The vortex formation time is 0.6 and is based on the flat plate travel length and chord length. Support for this research came from NSF REU Grant 1358991 and CBET 1335848.

  13. Phenotypic plasticity in the range-margin population of the lycaenid butterfly Zizeeria maha

    PubMed Central

    2010-01-01

    Background Many butterfly species have been experiencing the northward range expansion and physiological adaptation, probably due to climate warming. Here, we document an extraordinary field case of a species of lycaenid butterfly, Zizeeria maha, for which plastic phenotypes of wing color-patterns were revealed at the population level in the course of range expansion. Furthermore, we examined whether this outbreak of phenotypic changes was able to be reproduced in a laboratory. Results In the recently expanded northern range margins of this species, more than 10% of the Z. maha population exhibited characteristic color-pattern modifications on the ventral wings for three years. We physiologically reproduced similar phenotypes by an artificial cold-shock treatment of a normal southern population, and furthermore, we genetically reproduced a similar phenotype after selective breeding of a normal population for ten generations, demonstrating that the cold-shock-induced phenotype was heritable and partially assimilated genetically in the breeding line. Similar genetic process might have occurred in the previous and recent range-margin populations as well. Relatively minor modifications expressed in the tenth generation of the breeding line together with other data suggest a role of founder effect in this field case. Conclusions Our results support the notion that the outbreak of the modified phenotypes in the recent range-margin population was primed by the revelation of plastic phenotypes in response to temperature stress and by the subsequent genetic process in the previous range-margin population, followed by migration and temporal establishment of genetically unstable founders in the recent range margins. This case presents not only an evolutionary role of phenotypic plasticity in the field but also a novel evolutionary aspect of range expansion at the species level. PMID:20718993

  14. Fluid Dynamics of Clap-and-Fling with Highly Flexible Wings inspired by the Locomotion of Sea Butterflies

    NASA Astrophysics Data System (ADS)

    Zhou, Zhuoyu; Shoele, Kourosh; Adhikari, Deepak; Yen, Jeannette; Webster, Donald; Mittal, Rajat; Johns Hopkins University Team; Georgia Institute of Technology Team

    2015-11-01

    This study is motivated by the locomotion of sea butterflies (L. Helicina) which propel themselves in the water column using highly flexible wing-like parapodia. These animals execute a complex clap-and-fling with their highly flexible wings that is different from that of insects, and the fluid dynamics of which is not well understood. We use two models to study the fluid dyamics of these wings. In the first, we use prescribed wing kinematics that serve as a model of those observed for these animals. The second model is a fluid-structure interaction model where wing-like parapodia are modeled as flexible but inextensible membranes. The membrane properties, such as bending and stretching stiffness are modified such that the corresponding motion qualitatively matches the kinematics of L. helicina. Both models are used to examine the fluid dynamics of the clap-and-fling and its effectiveness in generating lift for these animals. Acknowledgement - research is supported by a grant from NSF.

  15. Classical lepidopteran wing scale colouration in the giant butterfly-moth Paysandisia archon.

    PubMed

    Stavenga, Doekele G; Leertouwer, Hein L; Meglič, Andrej; Drašlar, Kazimir; Wehling, Martin F; Pirih, Primož; Belušič, Gregor

    2018-01-01

    The palm borer moth Paysandisia archon (Castniidae; giant butterfly-moths) has brown dorsal forewings and strikingly orange-coloured dorsal hindwings with white spots surrounded by black margins. Here, we have studied the structure and pigments of the wing scales in the various coloured wing areas, applying light and electron microscopy and (micro)spectrophotometry, and we analysed the spatial reflection properties with imaging scatterometry. The scales in the white spots are unpigmented, those in the black and brown wing areas contain various amounts of melanin, and the orange wing scales contain a blue-absorbing ommochrome pigment. In all scale types, the upper lamina acts as a diffuser and the lower lamina as a thin film interference reflector, with thickness of about 200 nm. Scale stacking plays an important role in creating the strong visual signals: the colour of the white eyespots is created by stacks of unpigmented blue scales, while the orange wing colour is strongly intensified by stacking the orange scales.

  16. Fabrication and good ethanol sensing of biomorphic SnO2 with architecture hierarchy of butterfly wings.

    PubMed

    Song, Fang; Su, Huilan; Han, Jie; Zhang, Di; Chen, Zhixin

    2009-12-09

    Using super-hydrophobic butterfly wings as templates, we developed an aqueous sol-gel soakage process assisted by ethanol-wetting and followed by calcination to fabricate well-organized porous hierarchical SnO(2) with connective hollow interiors and thin mesoporous walls. The exquisite hierarchical architecture of SnO(2) is faithfully replicated from the lightweight skeleton of butterfly wings at the level from nano- to macro-scales. On the basis of the self-assembly of SnO(2) nanocrystallites with diameter around 7.0 nm, the interconnected tubes (lamellas), the fastigiated hollow tubers (pillars) and the double-layered substrates further construct the biomorphic hierarchical architecture. Benefiting from the small grain size and the unique hierarchical architecture, the biomorphic SnO(2) as an ethanol sensor exhibits high sensitivity (49.8 to 50 ppm ethanol), and fast response/recovery time (11/31 s to 50 ppm ethanol) even at relatively low working temperature (170 degrees C).

  17. Bioinspired Omnidirectional Self-Stable Reflectors with Multiscale Hierarchical Structures.

    PubMed

    Han, Zhiwu; Mu, Zhengzhi; Li, Bo; Feng, Xiaoming; Wang, Ze; Zhang, Junqiu; Niu, Shichao; Ren, Luquan

    2017-08-30

    Structured surfaces, demonstrating various wondrous physicochemical performances, are ubiquitous phenomena in nature. Butterfly wings with impressive structural colors are an interesting example for multiscale hierarchical structures (MHSs). However, most natural structural colors are relatively unstable and highly sensitive to incident angles, which limit their potential practical applications to a certain extent. Here, we reported a bioinspired color reflector with omnidirectional reflective self-stable (ORS) properties, which is inspired by the wing scales of Papilio palinurus butterfly. Through experimental exploration and theoretical analysis, it was found that the vivid colors of such butterfly wings are structure-based and possess novel ORS properties, which attributes to the multiple optical actions between light and the complex structures coupling the inverse opal-like structures (IOSs) and stacked lamellar ridges (SLRs). On the basis of this, we designed and successfully fabricated the SiO 2 -based bioinspired color reflectors (BCRs) through a facile and effective biotemplate method. It was confirmed that the MHSs in biotemplate are inherited by the obtained SiO 2 -based BCRs. More importantly, the SiO 2 -based BCRs also demonstrated the similar ORS properties in a wide wavelength range. We forcefully anticipate that the reported MHS-based ORS performance discovered in butterfly wing scales here could offer new thoughts for scientists to solve unstable reflection issues in particular optical field. The involved biotemplate fabrication method offers a facile and effective strategy for fabricating functional nanomaterials or bioinspired nanodevices with 3D complex nanostructures, such as structured optical devices, displays, and optoelectronic equipment.

  18. Butterfly effects: novel functional materials inspired from the wings scales.

    PubMed

    Zhang, Wang; Gu, Jiajun; Liu, Qinglei; Su, Huilan; Fan, Tongxiang; Zhang, Di

    2014-10-07

    Through millions of years of evolutionary selection, nature has created biological materials with various functional properties for survival. Many complex natural architectures, such as shells, bones, and honeycombs, have been studied and imitated in the design and fabrication of materials with enhanced hardness and stiffness. Recently, more and more researchers have started to research the wings of butterflies, mostly because of their dazzling colors. It was found that most of these iridescent colors are caused by periodic photonic structures on the scales that make up the surfaces of these wings. These materials have recently become a focus of multidiscipline research because of their promising applications in the display of structural colors, and in advanced sensors, photonic crystals, and solar cells. This paper review aims to provide a perspective overview of the research inspired by these wing structures in recent years.

  19. Properties of sunspot cycles and hemispheric wings since the 19th century

    NASA Astrophysics Data System (ADS)

    Leussu, Raisa; Usoskin, Ilya G.; Arlt, Rainer; Mursula, Kalevi

    2016-08-01

    Aims: The latitudinal evolution of sunspot emergence over the course of the solar cycle, the so-called butterfly diagram, is a fundamental property of the solar dynamo. Here we present a study of the butterfly diagram of sunspot group occurrence for cycles 7-10 and 11-23 using data from a recently digitized sunspot drawings by Samuel Heinrich Schwabe in 1825-1867, and from RGO/USAF/NOAA(SOON) compilation of sunspot groups in 1874-2015. Methods: We developed a new, robust method of hemispheric wing separation based on an analysis of long gaps in sunspot group occurrence in different latitude bands. The method makes it possible to ascribe each sunspot group to a certain wing (solar cycle and hemisphere), and separate the old and new cycle during their overlap. This allows for an improved study of solar cycles compared to the common way of separating the cycles. Results: We separated each hemispheric wing of the butterfly diagram and analysed them with respect to the number of groups appearing in each wing, their lengths, hemispheric differences, and overlaps. Conclusions: The overlaps of successive wings were found to be systematically longer in the northern hemisphere for cycles 7-10, but in the southern hemisphere for cycles 16-22. The occurrence of sunspot groups depicts a systematic long-term variation between the two hemispheres. During Schwabe time, the hemispheric asymmetry was north-dominated during cycle 9 and south-dominated during cycle 10.

  20. Aerodynamic comparison of a butterfly-like flapping wing-body model and a revolving-wing model

    NASA Astrophysics Data System (ADS)

    Suzuki, Kosuke; Yoshino, Masato

    2017-06-01

    The aerodynamic performance of flapping- and revolving-wing models is investigated by numerical simulations based on an immersed boundary-lattice Boltzmann method. As wing models, we use (i) a butterfly-like model with a body and flapping-rectangular wings and (ii) a revolving-wing model with the same wings as the flapping case. Firstly, we calculate aerodynamic performance factors such as the lift force, the power, and the power loading of the two models for Reynolds numbers in the range of 50-1000. For the flapping-wing model, the power loading is maximal for the maximum angle of attack of 90°, a flapping amplitude of roughly 45°, and a phase shift between the flapping angle and the angle of attack of roughly 90°. For the revolving-wing model, the power loading peaks for an angle of attack of roughly 45°. In addition, we examine the ground effect on the aerodynamic performance of the revolving-wing model. Secondly, we compare the aerodynamic performance of the flapping- and revolving-wing models at their respective maximal power loadings. It is found that the revolving-wing model is more efficient than the flapping-wing model both when the body of the latter is fixed and where it can move freely. Finally, we discuss the relative agilities of the flapping- and revolving-wing models.

  1. Polymorphism at a mimicry supergene maintained by opposing frequency-dependent selection pressures.

    PubMed

    Chouteau, Mathieu; Llaurens, Violaine; Piron-Prunier, Florence; Joron, Mathieu

    2017-08-01

    Explaining the maintenance of adaptive diversity within populations is a long-standing goal in evolutionary biology, with important implications for conservation, medicine, and agriculture. Adaptation often leads to the fixation of beneficial alleles, and therefore it erodes local diversity so that understanding the coexistence of multiple adaptive phenotypes requires deciphering the ecological mechanisms that determine their respective benefits. Here, we show how antagonistic frequency-dependent selection (FDS), generated by natural and sexual selection acting on the same trait, maintains mimicry polymorphism in the toxic butterfly Heliconius numata Positive FDS imposed by predators on mimetic signals favors the fixation of the most abundant and best-protected wing-pattern morph, thereby limiting polymorphism. However, by using mate-choice experiments, we reveal disassortative mate preferences of the different wing-pattern morphs. The resulting negative FDS on wing-pattern alleles is consistent with the excess of heterozygote genotypes at the supergene locus controlling wing-pattern variation in natural populations of H. numata The combined effect of positive and negative FDS on visual signals is sufficient to maintain a diversity of morphs displaying accurate mimicry with other local prey, although some of the forms only provide moderate protection against predators. Our findings help understand how alternative adaptive phenotypes can be maintained within populations and emphasize the need to investigate interactions between selective pressures in other cases of puzzling adaptive polymorphism.

  2. Polymorphism at a mimicry supergene maintained by opposing frequency-dependent selection pressures

    PubMed Central

    Chouteau, Mathieu; Llaurens, Violaine; Piron-Prunier, Florence; Joron, Mathieu

    2017-01-01

    Explaining the maintenance of adaptive diversity within populations is a long-standing goal in evolutionary biology, with important implications for conservation, medicine, and agriculture. Adaptation often leads to the fixation of beneficial alleles, and therefore it erodes local diversity so that understanding the coexistence of multiple adaptive phenotypes requires deciphering the ecological mechanisms that determine their respective benefits. Here, we show how antagonistic frequency-dependent selection (FDS), generated by natural and sexual selection acting on the same trait, maintains mimicry polymorphism in the toxic butterfly Heliconius numata. Positive FDS imposed by predators on mimetic signals favors the fixation of the most abundant and best-protected wing-pattern morph, thereby limiting polymorphism. However, by using mate-choice experiments, we reveal disassortative mate preferences of the different wing-pattern morphs. The resulting negative FDS on wing-pattern alleles is consistent with the excess of heterozygote genotypes at the supergene locus controlling wing-pattern variation in natural populations of H. numata. The combined effect of positive and negative FDS on visual signals is sufficient to maintain a diversity of morphs displaying accurate mimicry with other local prey, although some of the forms only provide moderate protection against predators. Our findings help understand how alternative adaptive phenotypes can be maintained within populations and emphasize the need to investigate interactions between selective pressures in other cases of puzzling adaptive polymorphism. PMID:28673971

  3. Changes is genes coding for laccases 1 and 2 may contribute to deformation and reduction of wings in apollo butterfly (Parnassius apollo, Lepidoptera: Papilionidae) from the isolated population in Pieniny National Park (Poland).

    PubMed

    Łukasiewicz, Kinga; Węgrzyn, Grzegorz

    2016-01-01

    An isolated population of apollo butterfly (Parnassius apollo, Lepidoptera: Papilionidae) occurs in Pieniny National Park (Poland). Deformations and reductions of wings in a relatively large number of individuals from this population is found, yet the reasons for these defects are unknown. During studies devoted to identify cause(s) of this phenomenon, we found that specific regions of genes coding of enzymes laccases 1 and 2 could not be amplified from DNA samples isolated from large fractions of malformed insects while expected PCR products were detected in almost all (with one exception) normal butterflies. Laccases (p-diphenol:dioxygen oxidoreductases) are oxidases containing several copper atoms. They catalyse single-electron oxidations of phenolic or other compounds with concomitant reduction of oxygen to water. In insects, their enzymatic activities were found previously in epidermis, midgut, Malpighian tubules, salivary glands, and reproductive tissues. Therefore, we suggest that defects in genes coding for laccases might contribute to deformation and reduction of wings in apollo butterflies, though it seems obvious that deficiency in these enzymes could not be the sole cause of these developmental improperties in P. apollo from Pieniny National Park.

  4. Reflectivity of the gyroid biophotonic crystals in the ventral wing scales of the Green Hairstreak butterfly, Callophrys rubi.

    PubMed

    Michielsen, K; De Raedt, H; Stavenga, D G

    2010-05-06

    We present a comparison of the computer simulation data of gyroid nanostructures with optical measurements (reflectivity spectra and scattering diagrams) of ventral wing scales of the Green Hairstreak butterfly, Callophrys rubi. We demonstrate that the omnidirectional green colour arises from the gyroid cuticular structure grown in the domains of different orientation. We also show that this three-dimensional structure, operating as a biophotonic crystal, gives rise to various polarization effects. We briefly discuss the possible biological utility of the green coloration and polarization effects.

  5. Bio-sensing with butterfly wings: naturally occurring nano-structures for SERS-based malaria parasite detection.

    PubMed

    Garrett, Natalie L; Sekine, Ryo; Dixon, Matthew W A; Tilley, Leann; Bambery, Keith R; Wood, Bayden R

    2015-09-07

    Surface enhanced Raman scattering (SERS) is a powerful tool with great potential to provide improved bio-sensing capabilities. The current 'gold-standard' method for diagnosis of malaria involves visual inspection of blood smears using light microscopy, which is time consuming and can prevent early diagnosis of the disease. We present a novel surface-enhanced Raman spectroscopy substrate based on gold-coated butterfly wings, which enabled detection of malarial hemozoin pigment within lysed blood samples containing 0.005% and 0.0005% infected red blood cells.

  6. High angular and spectral selectivity of purple emperor (Lepidoptera: Apatura iris and A. ilia) butterfly wings.

    PubMed

    Pantelić, Dejan; Curčić, Srećko; Savić-Šević, Svetlana; Korać, Aleksandra; Kovačević, Aleksander; Curčić, Božidar; Bokić, Bojana

    2011-03-28

    The iridescent features of the butterfly species Apatura iris (Linnaeus, 1758) and A. ilia (Denis & Schiffermüller, 1775) were studied. We recognized the structural color of scales only on the dorsal side of both the fore and hind wings of males of both of the aforementioned butterfly species. The scale dimensions and microstructure were analyzed by a scanning electron microscope (SEM) and transmission electron microscope (TEM). The optical properties were measured and it was found that the peak reflectivity is around 380 nm, with a spectral width (full width at half maximum) of approximately 50 nm in both species. The angular selectivity is high and a purple iridescent color is observed within the angular range of only 18 degrees in both species.

  7. Taxonomy of Mechanitis (f.) (Lepidoptera: Nymphalidae) from the west Colombian Andes: an integrative approach.

    PubMed

    Giraldo, C E; Uribe, S I

    2012-12-01

    Species identification in the butterfly genus Mechanitis (F.) (Lepidoptera: Nymphalidae) becomes difficult when it is based only on wing color patterns, a common practice in butterfly taxonomy. Difficulties in Mechanitis taxonomy are related to the widespread mimicry and polymorphism among species belonging to this genus. Species recognition and inventories of Mechanitis genus in geographic areas as the Andean region of Colombia are of particular interest and the use of more than one character for taxonomic identification is desirable. In this study, we included morphological, ecological, and mitochondrial DNA data to identify the occurring species in this region. Species of Mechanitis were studied from ecological, morphological, and molecular perspectives considering host plant identification, oviposition behavior, and life cycles under laboratory conditions. Immature morphology, patterns of wing color, and genital structures of adults were also studied. The genetic barcoding region of the cytochrome oxidase I mitochondrial gene was sequenced and used to verify the limits between species previously defined by the other characters and to validate its usefulness for species delimitation in this particular genus. The integrative approach combining independent datasets successfully allowed species identification as compared to the approach based on a single dataset. Three well-differentiated species were found in the studied region, Mechanitis menapis (Hewitson), Mechanitis polymnia (Linnaeus), and Mechanitis lysimnia (Fabricius). New valuable characters that could improve taxonomic identification in this genus are considered.

  8. Fabrication of the replica templated from butterfly wing scales with complex light trapping structures

    NASA Astrophysics Data System (ADS)

    Han, Zhiwu; Li, Bo; Mu, Zhengzhi; Yang, Meng; Niu, Shichao; Zhang, Junqiu; Ren, Luquan

    2015-11-01

    The polydimethylsiloxane (PDMS) positive replica templated twice from the excellent light trapping surface of butterfly Trogonoptera brookiana wing scales was fabricated by a simple and promising route. The exact SiO2 negative replica was fabricated by using a synthesis method combining a sol-gel process and subsequent selective etching. Afterwards, a vacuum-aided process was introduced to make PDMS gel fill into the SiO2 negative replica, and the PDMS gel was solidified in an oven. Then, the SiO2 negative replica was used as secondary template and the structures in its surface was transcribed onto the surface of PDMS. At last, the PDMS positive replica was obtained. After comparing the PDMS positive replica and the original bio-template in terms of morphology, dimensions and reflectance spectra and so on, it is evident that the excellent light trapping structures of butterfly wing scales were inherited by the PDMS positive replica faithfully. This bio-inspired route could facilitate the preparation of complex light trapping nanostructure surfaces without any assistance from other power-wasting and expensive nanofabrication technologies.

  9. Micro- and nanostructures of iridescent wing scales in purple emperor butterflies (Lepidoptera: Apatura ilia and A. iris).

    PubMed

    Ćurčić, Srećko B; Pantelić, Dejan V; Ćurčić, Božidar P M; Savić-Šević, Svetlana N; Makarov, Slobodan E; Lačković, Vesna B; Labudović-Borović, Milica M; Ćurčić, Nina B; Stojanović, Dejan V

    2012-07-01

    Apatura ilia (Denis and Schiffermüller, 1775) and A. iris (Linnaeus, 1758) are fascinating butterflies found in the Palaearctic ecozone (excepting the north of Africa). The wings of these insects are covered with a great number of two types of scales positioned like roof tiles. Type I scales are on the surface, while type II scales are situated below them. The structural color of the type I scales is recognized only on the dorsal side of both the fore and hind wings of the males of the aforementioned species. Both types of scales are responsible for pigment color of the wings, but iridescence is observed only in the type I scales. The brilliant structural color is due to a multilayer structure. The features of the scales, their dimensions and fine structure were obtained using scanning electron microscopy. Cross sections of the scales were then analyzed by transmission electron microscopy. The scales of the "normal" and clytie forms of A. ilia have a different nanostructure, but are of the same type. A similar type of structure, but with a different morphology, was also noticed in A. iris. The scales of the analyzed species resemble the scales of tropical Morpho butterflies. Copyright © 2012 Wiley Periodicals, Inc.

  10. Fluid-Dynamics of Underwater Flight in Sea Butterflies: Analysis using Tomographic PIV

    NASA Astrophysics Data System (ADS)

    Adhikari, D.; Murphy, D. W.; Webster, D. R.; Yen, J.

    2014-11-01

    Sea butterflies, Limacina helicina, swim in sea water with a pair of gelatinous ``wings'' (or parapodia). Their unique propulsion mechanism has been hypothesized to consist of a combination of drag-based propulsion (rowing) and lift-based propulsion (flapping). Drag-based propulsion utilizes maximum drag on the wings during power stroke, followed by minimum drag during recovery stroke. Lift-based propulsion, in contrast, utilizes a pressure difference between the top and bottom of the wings. We present the 3D kinematics of a free-swimming sea butterfly and its induced volumetric velocity field using tomographic PIV. Both upstroke and downstroke motions propel the animal (1 - 3 mm) upward in a sawtooth-like trajectory with average speed of 5 - 15 mm/s (Re = 5 - 45) and roll the calcareous shell forwards-and-backwards at 4 - 5 Hz. The rolling motion effectively positions the wings such that they stroke downward during both the power and recovery strokes, hence inducing upward motion during both phases. A clap-and-fling mechanism is observed at the beginning of the flapping cycle. As the wings come into contact, the velocity of the organism is 2 mm/s. During fling motion, high (unsteady) lift causes the organism velocity to reach 35 mm/s. Separation vortices are observed during the fling motion, and vortices with an opposite sense of rotation form closer to the base of the wing due to the upward translation of the organism. The separation vortices shed into the wake, as the organism translates upward, in the form of separate vortex pairs.

  11. Reflectivity of the gyroid biophotonic crystals in the ventral wing scales of the Green Hairstreak butterfly, Callophrys rubi

    PubMed Central

    Michielsen, K.; De Raedt, H.; Stavenga, D. G.

    2010-01-01

    We present a comparison of the computer simulation data of gyroid nanostructures with optical measurements (reflectivity spectra and scattering diagrams) of ventral wing scales of the Green Hairstreak butterfly, Callophrys rubi. We demonstrate that the omnidirectional green colour arises from the gyroid cuticular structure grown in the domains of different orientation. We also show that this three-dimensional structure, operating as a biophotonic crystal, gives rise to various polarization effects. We briefly discuss the possible biological utility of the green coloration and polarization effects. PMID:19828506

  12. Roughness Effects on the Formation of a Leading Edge Vortex

    NASA Astrophysics Data System (ADS)

    Elliott, Cassidy; Lang, Amy; Wahidi, Redha; Wilroy, Jacob

    2017-11-01

    Microscopic scales cover the wings of Monarch butterflies, creating a patterned surface that acts as a natural energy capture mechanism. This patterning is thought to delay the growth of the leading edge vortex (LEV) produced by the flapping motion of a wing. Increased skin friction caused by the scales leads to a weaker LEV being shed into the butterfly's wake, lessening drag and increasing flight efficiency. To test how this roughness effects LEV formation, a plate of random roughness was designed in SolidWorks and printed on the Objet 30 Pro 3D printer. A 2x3x5 cubic foot tow tank was used to test the rough plate at Reynold's numbers of 1500, 3000, and 6000 (velocities of 8, 16, and 32 mm/s) at an angle of attack of 45 degrees. Images were captured of the LEV generated when the plate was towed upwards through the particle-seeded flow. These images were used to determine the XY velocity of the particles using a technique called Digital Particle Image Velocimetry (DPIV). Codes written in MATLAB were used to track and measure the strength of the LEV. Circulation values for the randomly-rough plate were then compared to the same values generated in a previous experiment that used a smooth plate and a grooved plate to determine the effect of the patterning on vortex development. Funding for this research project was provided by the National Science Foundation under the Research Experience for Undergraduates (REU) program (REU Supplement CBET 1628600 under CBET 1335848).

  13. Flexible, angle-independent, structural color reflectors inspired by morpho butterfly wings.

    PubMed

    Chung, Kyungjae; Yu, Sunkyu; Heo, Chul-Joon; Shim, Jae Won; Yang, Seung-Man; Han, Moon Gyu; Lee, Hong-Seok; Jin, Yongwan; Lee, Sang Yoon; Park, Namkyoo; Shin, Jung H

    2012-05-08

    Thin-film color reflectors inspired by Morpho butterflies are fabricated. Using a combination of directional deposition, silica microspheres with a wide size distribution, and a PDMS (polydimethylsiloxane) encasing, a large, flexible reflector is created that actually provides better angle-independent color characteristics than Morpho butterflies and which can even be bent and folded freely without losing its Morpho-mimetic photonic properties. Copyright © 2012 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim.

  14. Structure, function, and self-assembly of single network gyroid (I4132) photonic crystals in butterfly wing scales.

    PubMed

    Saranathan, Vinodkumar; Osuji, Chinedum O; Mochrie, Simon G J; Noh, Heeso; Narayanan, Suresh; Sandy, Alec; Dufresne, Eric R; Prum, Richard O

    2010-06-29

    Complex three-dimensional biophotonic nanostructures produce the vivid structural colors of many butterfly wing scales, but their exact nanoscale organization is uncertain. We used small angle X-ray scattering (SAXS) on single scales to characterize the 3D photonic nanostructures of five butterfly species from two families (Papilionidae, Lycaenidae). We identify these chitin and air nanostructures as single network gyroid (I4(1)32) photonic crystals. We describe their optical function from SAXS data and photonic band-gap modeling. Butterflies apparently grow these gyroid nanostructures by exploiting the self-organizing physical dynamics of biological lipid-bilayer membranes. These butterfly photonic nanostructures initially develop within scale cells as a core-shell double gyroid (Ia3d), as seen in block-copolymer systems, with a pentacontinuous volume comprised of extracellular space, cell plasma membrane, cellular cytoplasm, smooth endoplasmic reticulum (SER) membrane, and intra-SER lumen. This double gyroid nanostructure is subsequently transformed into a single gyroid network through the deposition of chitin in the extracellular space and the degeneration of the rest of the cell. The butterflies develop the thermodynamically favored double gyroid precursors as a route to the optically more efficient single gyroid nanostructures. Current approaches to photonic crystal engineering also aim to produce single gyroid motifs. The biologically derived photonic nanostructures characterized here may offer a convenient template for producing optical devices based on biomimicry or direct dielectric infiltration.

  15. Structure, function, and self-assembly of single network gyroid (I4132) photonic crystals in butterfly wing scales

    PubMed Central

    Saranathan, Vinodkumar; Osuji, Chinedum O.; Mochrie, Simon G. J.; Noh, Heeso; Narayanan, Suresh; Sandy, Alec; Dufresne, Eric R.; Prum, Richard O.

    2010-01-01

    Complex three-dimensional biophotonic nanostructures produce the vivid structural colors of many butterfly wing scales, but their exact nanoscale organization is uncertain. We used small angle X-ray scattering (SAXS) on single scales to characterize the 3D photonic nanostructures of five butterfly species from two families (Papilionidae, Lycaenidae). We identify these chitin and air nanostructures as single network gyroid (I4132) photonic crystals. We describe their optical function from SAXS data and photonic band-gap modeling. Butterflies apparently grow these gyroid nanostructures by exploiting the self-organizing physical dynamics of biological lipid-bilayer membranes. These butterfly photonic nanostructures initially develop within scale cells as a core-shell double gyroid (Ia3d), as seen in block-copolymer systems, with a pentacontinuous volume comprised of extracellular space, cell plasma membrane, cellular cytoplasm, smooth endoplasmic reticulum (SER) membrane, and intra-SER lumen. This double gyroid nanostructure is subsequently transformed into a single gyroid network through the deposition of chitin in the extracellular space and the degeneration of the rest of the cell. The butterflies develop the thermodynamically favored double gyroid precursors as a route to the optically more efficient single gyroid nanostructures. Current approaches to photonic crystal engineering also aim to produce single gyroid motifs. The biologically derived photonic nanostructures characterized here may offer a convenient template for producing optical devices based on biomimicry or direct dielectric infiltration. PMID:20547870

  16. A conserved supergene locus controls colour pattern diversity in Heliconius butterflies.

    PubMed

    Joron, Mathieu; Papa, Riccardo; Beltrán, Margarita; Chamberlain, Nicola; Mavárez, Jesús; Baxter, Simon; Abanto, Moisés; Bermingham, Eldredge; Humphray, Sean J; Rogers, Jane; Beasley, Helen; Barlow, Karen; ffrench-Constant, Richard H; Mallet, James; McMillan, W Owen; Jiggins, Chris D

    2006-10-01

    We studied whether similar developmental genetic mechanisms are involved in both convergent and divergent evolution. Mimetic insects are known for their diversity of patterns as well as their remarkable evolutionary convergence, and they have played an important role in controversies over the respective roles of selection and constraints in adaptive evolution. Here we contrast three butterfly species, all classic examples of Müllerian mimicry. We used a genetic linkage map to show that a locus, Yb, which controls the presence of a yellow band in geographic races of Heliconius melpomene, maps precisely to the same location as the locus Cr, which has very similar phenotypic effects in its co-mimic H. erato. Furthermore, the same genomic location acts as a "supergene", determining multiple sympatric morphs in a third species, H. numata. H. numata is a species with a very different phenotypic appearance, whose many forms mimic different unrelated ithomiine butterflies in the genus Melinaea. Other unlinked colour pattern loci map to a homologous linkage group in the co-mimics H. melpomene and H. erato, but they are not involved in mimetic polymorphism in H. numata. Hence, a single region from the multilocus colour pattern architecture of H. melpomene and H. erato appears to have gained control of the entire wing-pattern variability in H. numata, presumably as a result of selection for mimetic "supergene" polymorphism without intermediates. Although we cannot at this stage confirm the homology of the loci segregating in the three species, our results imply that a conserved yet relatively unconstrained mechanism underlying pattern switching can affect mimicry in radically different ways. We also show that adaptive evolution, both convergent and diversifying, can occur by the repeated involvement of the same genomic regions.

  17. Characterising the Phenotypic Diversity of Papilio dardanus Wing Patterns Using an Extensive Museum Collection

    PubMed Central

    Thompson, Martin J.; Timmermans, Martijn J. T. N.

    2014-01-01

    The history of 20th Century evolutionary biology can be followed through the study of mimetic butterflies. From the initial findings of discontinuous polymorphism through the debates regarding the evolution of mimicry and the step-size of evolutionary change, to the studies on supergene evolution and molecular characterisation of butterfly genomes, mimetic butterflies have been at the heart of evolutionary thought for over 100 years. During this time, few species have received as much attention and in-depth study as Papilio dardanus. To assist all aspects of mimicry research, we present a complete data-derived overview of the extent of polymorphism within this species. Using historical samples permanently held by the NHM London, we document the extent of phenotypic variation and characterise the diversity present in each of the subspecies and how it varies across Africa. We also demonstrate an association between “imperfect” mimetic forms and the transitional race formed in the area where Eastern and Western African populations meet around Lake Victoria. We present a novel portal for access to this collection, www.mimeticbutterflies.org, allowing remote access to this unique repository. It is hoped that this online resource can act as a nucleus for the sharing and dissemination of other collections databases and imagery connected with mimetic butterflies. PMID:24837717

  18. Gene regulatory networks reused to build novel traits: co-option of an eye-related gene regulatory network in eye-like organs and red wing patches on insect wings is suggested by optix expression.

    PubMed

    Monteiro, Antónia

    2012-03-01

    Co-option of the eye developmental gene regulatory network may have led to the appearance of novel functional traits on the wings of flies and butterflies. The first trait is a recently described wing organ in a species of extinct midge resembling the outer layers of the midge's own compound eye. The second trait is red pigment patches on Heliconius butterfly wings connected to the expression of an eye selector gene, optix. These examples, as well as others, are discussed regarding the type of empirical evidence and burden of proof that have been used to infer gene network co-option underlying the origin of novel traits. A conceptual framework describing increasing confidence in inference of network co-option is proposed. Novel research directions to facilitate inference of network co-option are also highlighted, especially in cases where the pre-existent and novel traits do not resemble each other. Copyright © 2012 WILEY Periodicals, Inc.

  19. Population demography of alpine butterflies: Boloria pales and Boloria napaea (Lepidoptera: Nymphalidae) and their specific adaptations to high mountain environments

    NASA Astrophysics Data System (ADS)

    Ehl, Stefan; Ebertshäuser, Marlene; Gros, Patrick; Schmitt, Thomas

    2017-11-01

    High mountain ecosystems are extreme habitats, and adaptation strategies to this ecosystem are still poorly understood in most groups. To unravel such strategies, we performed a MRR study in the Hohe Tauern National Park (Salzburg, Austria) with two nymphalid butterfly species, Boloria pales and B. napaea. We analysed their population structure over one flight period by studying the development of population size and wing wear. B. pales had more individuals and a higher survival probability than B. napaea; the sensitivity to extreme weather conditions or other external influences was higher in B. napaea. We only observed proterandry in B. pales. Imagines of both species survived under snow for at least some days. Additionally, we observed a kind of risk-spreading, in that individuals of both species, and especially B. pales, have regularly emerged throughout the flight period. This emergence pattern divided the population's age structure into three phases: an initial phase with decreasing wing quality (emergence > mortality), followed by an equilibrium phase with mostly constant average wing condition (emergence = mortality) and a final ageing phase with strongly deteriorating wing condition (mortality » emergence). Consequently, neither species would likely become extinct because of particularly unsuitable weather conditions during a single flight period. The observed differences between the two species suggest a better regional adaptation of B. pales, which is restricted to high mountain systems of Europe. In contrast, the arctic-alpine B. napaea might be best adapted to conditions in the Arctic and not the more southern high mountain systems. However, this needs to be examined during future research in the Arctic.

  20. Attack risk for butterflies changes with eyespot number and size

    PubMed Central

    Ho, Sebastian; Schachat, Sandra R.; Piel, William H.; Monteiro, Antónia

    2016-01-01

    Butterfly eyespots are known to function in predator deflection and predator intimidation, but it is still unclear what factors cause eyespots to serve one function over the other. Both functions have been demonstrated in different species that varied in eyespot size, eyespot number and wing size, leaving the contribution of each of these factors to butterfly survival unclear. Here, we study how each of these factors contributes to eyespot function by using paper butterfly models, where each factor is varied in turn, and exposing these models to predation in the field. We find that the presence of multiple, small eyespots results in high predation, whereas single large eyespots (larger than 6 mm in diameter) results in low predation. These data indicate that single large eyespots intimidate predators, whereas multiple small eyespots produce a conspicuous, but non-intimidating signal to predators. We propose that eyespots may gain an intimidation function by increasing in size. Our measurements of eyespot size in 255 nymphalid butterfly species show that large eyespots are relatively rare and occur predominantly on ventral wing surfaces. By mapping eyespot size on the phylogeny of the family Nymphalidae, we show that these large eyespots, with a potential intimidation function, are dispersed throughout multiple nymphalid lineages, indicating that phylogeny is not a strong predictor of eyespot size. PMID:26909190

  1. Artificial selection for structural color on butterfly wings and comparison with natural evolution

    PubMed Central

    Wasik, Bethany R.; Liew, Seng Fatt; Lilien, David A.; Dinwiddie, April J.; Noh, Heeso; Cao, Hui; Monteiro, Antónia

    2014-01-01

    Brilliant animal colors often are produced from light interacting with intricate nano-morphologies present in biological materials such as butterfly wing scales. Surveys across widely divergent butterfly species have identified multiple mechanisms of structural color production; however, little is known about how these colors evolved. Here, we examine how closely related species and populations of Bicyclus butterflies have evolved violet structural color from brown-pigmented ancestors with UV structural color. We used artificial selection on a laboratory model butterfly, B. anynana, to evolve violet scales from UV brown scales and compared the mechanism of violet color production with that of two other Bicyclus species, Bicyclus sambulos and Bicyclus medontias, which have evolved violet/blue scales independently via natural selection. The UV reflectance peak of B. anynana brown scales shifted to violet over six generations of artificial selection (i.e., in less than 1 y) as the result of an increase in the thickness of the lower lamina in ground scales. Similar scale structures and the same mechanism for producing violet/blue structural colors were found in the other Bicyclus species. This work shows that populations harbor large amounts of standing genetic variation that can lead to rapid evolution of scales’ structural color via slight modifications to the scales’ physical dimensions. PMID:25092295

  2. Artificial selection for structural color on butterfly wings and comparison with natural evolution.

    PubMed

    Wasik, Bethany R; Liew, Seng Fatt; Lilien, David A; Dinwiddie, April J; Noh, Heeso; Cao, Hui; Monteiro, Antónia

    2014-08-19

    Brilliant animal colors often are produced from light interacting with intricate nano-morphologies present in biological materials such as butterfly wing scales. Surveys across widely divergent butterfly species have identified multiple mechanisms of structural color production; however, little is known about how these colors evolved. Here, we examine how closely related species and populations of Bicyclus butterflies have evolved violet structural color from brown-pigmented ancestors with UV structural color. We used artificial selection on a laboratory model butterfly, B. anynana, to evolve violet scales from UV brown scales and compared the mechanism of violet color production with that of two other Bicyclus species, Bicyclus sambulos and Bicyclus medontias, which have evolved violet/blue scales independently via natural selection. The UV reflectance peak of B. anynana brown scales shifted to violet over six generations of artificial selection (i.e., in less than 1 y) as the result of an increase in the thickness of the lower lamina in ground scales. Similar scale structures and the same mechanism for producing violet/blue structural colors were found in the other Bicyclus species. This work shows that populations harbor large amounts of standing genetic variation that can lead to rapid evolution of scales' structural color via slight modifications to the scales' physical dimensions.

  3. Variability of the Structural Coloration in Two Butterfly Species with Different Prezygotic Mating Strategies

    PubMed Central

    Kertész, Krisztián; Bálint, Zsolt; Biró, László Péter

    2016-01-01

    Structural coloration variability was investigated in two Blue butterfly species that are common in Hungary. The males of Polyommatus icarus (Common Blue) and Plebejus argus (Silver-studded Blue) use their blue wing coloration for conspecific recognition. Despite living in the same type of habitat, these two species display differences in prezygotic mating strategy: the males of P. icarus are patrolling, while P. argus males have sedentary behavior. Therefore, the species-specific photonic nanoarchitecture, which is the source of the structural coloration, may have been subjected to different evolutionary effects. Despite the increasing interest in photonic nanoarchitectures of biological origin, there is a lack of studies focused on the biological variability of structural coloration that examine a statistically relevant number of individuals from the same species. To investigate possible structural color variation within the same species in populations separated by large geographical distances, climatic differences, or applied experimental conditions, one has to be able to compare these variations to the normal biological variability within a single population. The structural coloration of the four wings of 25 male individuals (100 samples for each species) was measured and compared using different light-collecting setups: perpendicular and with an integrating sphere. Significant differences were found in the near UV wavelength region that are perceptible by these polyommatine butterflies but are invisible to human observers. The differences are attributed to the differences in the photonic nanoarchitecture in the scales of these butterflies. Differences in the intensity of structural coloration were also observed and were tentatively attributed to the different prezygotic mating strategies of these insects. Despite the optical complexity of the scale covered butterfly wings, for sufficiently large sample batches, the averaged normal incidence measurements and the averaged measurements using an integrating sphere are in agreement. PMID:27832120

  4. Variability of the Structural Coloration in Two Butterfly Species with Different Prezygotic Mating Strategies.

    PubMed

    Piszter, Gábor; Kertész, Krisztián; Bálint, Zsolt; Biró, László Péter

    2016-01-01

    Structural coloration variability was investigated in two Blue butterfly species that are common in Hungary. The males of Polyommatus icarus (Common Blue) and Plebejus argus (Silver-studded Blue) use their blue wing coloration for conspecific recognition. Despite living in the same type of habitat, these two species display differences in prezygotic mating strategy: the males of P. icarus are patrolling, while P. argus males have sedentary behavior. Therefore, the species-specific photonic nanoarchitecture, which is the source of the structural coloration, may have been subjected to different evolutionary effects. Despite the increasing interest in photonic nanoarchitectures of biological origin, there is a lack of studies focused on the biological variability of structural coloration that examine a statistically relevant number of individuals from the same species. To investigate possible structural color variation within the same species in populations separated by large geographical distances, climatic differences, or applied experimental conditions, one has to be able to compare these variations to the normal biological variability within a single population. The structural coloration of the four wings of 25 male individuals (100 samples for each species) was measured and compared using different light-collecting setups: perpendicular and with an integrating sphere. Significant differences were found in the near UV wavelength region that are perceptible by these polyommatine butterflies but are invisible to human observers. The differences are attributed to the differences in the photonic nanoarchitecture in the scales of these butterflies. Differences in the intensity of structural coloration were also observed and were tentatively attributed to the different prezygotic mating strategies of these insects. Despite the optical complexity of the scale covered butterfly wings, for sufficiently large sample batches, the averaged normal incidence measurements and the averaged measurements using an integrating sphere are in agreement.

  5. Multiple factors correlating with wing malformations in the population of Parnassius apollo (Lepidoptera: Papilionidae) restituted from a low number of individuals: A mini review.

    PubMed

    Pierzynowska, Karolina; Skowron Volponi, Marta; Węgrzyn, Grzegorz

    2017-11-02

    The Apollo butterfly, Parnassius apollo (Linnaeus), was common in Europe over 100 years ago, but currently it is considered as near threatened. Different conservation programs have promoted the persistence of this species; however, it is still endangered. An example of such programs was the action devoted to reestablish the Apollo butterfly population in Pieniny National Park (Poland) from only 20-30 individuals which had survived till the last decade of the 20th century. This reintroduction has been successful; however, unexpected developmental problems appeared. Butterflies with deformed or reduced wings became frequent in the population living in the natural habitat, and particularly among those reared under seminatural conditions (in the same environment, but fenced by a net). Until recently, reasons for these malformations remained unknown. However, reports published during last months indicated that there are genetic, biochemical, and microbiological factors contributing to this phenomenon. In the malformed individuals, lesions in the wingless gene and dysfunctions of laccase 1 and 2 were found to be significantly more frequent than in normal insects. A large fraction of butterflies with deformed or reduced wings was devoid of the prokaryotic symbiont Wolbachia, which was present in most normal individuals. Moreover, Yersinia pseudotuberculosis (Pfeiffer) Smith and Thal, and Serratia sp., bacteria pathogenic to insects, were detected in the biological material from both normal and malformed butterflies from this population. These findings are summarized and discussed in this review, in the light of conservation of insects and restitution of their populations from a low number of individuals. © 2017 Institute of Zoology, Chinese Academy of Sciences.

  6. Domain morphology, boundaries, and topological defects in biophotonic gyroid nanostructures of butterfly wing scales

    PubMed Central

    Singer, Andrej; Boucheron, Leandra; Dietze, Sebastian H.; Jensen, Katharine E.; Vine, David; McNulty, Ian; Dufresne, Eric R.; Prum, Richard O.; Mochrie, Simon G. J.; Shpyrko, Oleg G.

    2016-01-01

    Many organisms in nature have evolved sophisticated cellular mechanisms to produce photonic nanostructures and, in recent years, diverse crystalline symmetries have been identified and related to macroscopic optical properties. However, because we know little about the distributions of domain sizes, the orientations of photonic crystals, and the nature of defects in these structures, we are unable to make the connection between the nanostructure and its development and functionality. We report on nondestructive studies of the morphology of chitinous photonic crystals in butterfly wing scales. Using spatially and angularly resolved x-ray diffraction, we find that the domains are highly oriented with respect to the whole scale, indicating growth from scale boundaries. X-ray coherent diffractive imaging reveals two types of crystalline domain interfaces: abrupt changes between domains emerging from distinct nucleation sites and smooth transitions with edge dislocations presumably resulting from internal stresses during nanostructure development. Our study of the scale structure reveals new aspects of photonic crystal growth in butterfly wings and shows their similarity to block copolymer materials. It opens new avenues to exploration of fundamental processes underlying the growth of biological photonic nanostructures in a variety of species. PMID:27386575

  7. Domain morphology, boundaries, and topological defects in biophotonic gyroid nanostructures of butterfly wing scales.

    PubMed

    Singer, Andrej; Boucheron, Leandra; Dietze, Sebastian H; Jensen, Katharine E; Vine, David; McNulty, Ian; Dufresne, Eric R; Prum, Richard O; Mochrie, Simon G J; Shpyrko, Oleg G

    2016-06-01

    Many organisms in nature have evolved sophisticated cellular mechanisms to produce photonic nanostructures and, in recent years, diverse crystalline symmetries have been identified and related to macroscopic optical properties. However, because we know little about the distributions of domain sizes, the orientations of photonic crystals, and the nature of defects in these structures, we are unable to make the connection between the nanostructure and its development and functionality. We report on nondestructive studies of the morphology of chitinous photonic crystals in butterfly wing scales. Using spatially and angularly resolved x-ray diffraction, we find that the domains are highly oriented with respect to the whole scale, indicating growth from scale boundaries. X-ray coherent diffractive imaging reveals two types of crystalline domain interfaces: abrupt changes between domains emerging from distinct nucleation sites and smooth transitions with edge dislocations presumably resulting from internal stresses during nanostructure development. Our study of the scale structure reveals new aspects of photonic crystal growth in butterfly wings and shows their similarity to block copolymer materials. It opens new avenues to exploration of fundamental processes underlying the growth of biological photonic nanostructures in a variety of species.

  8. Comprehensive microarray-based analysis for stage-specific larval camouflage pattern-associated genes in the swallowtail butterfly, Papilio xuthus

    PubMed Central

    2012-01-01

    Background Body coloration is an ecologically important trait that is often involved in prey-predator interactions through mimicry and crypsis. Although this subject has attracted the interest of biologists and the general public, our scientific knowledge on the subject remains fragmentary. In the caterpillar of the swallowtail butterfly Papilio xuthus, spectacular changes in the color pattern are observed; the insect mimics bird droppings (mimetic pattern) as a young larva, and switches to a green camouflage coloration (cryptic pattern) in the final instar. Despite the wide variety and significance of larval color patterns, few studies have been conducted at a molecular level compared with the number of studies on adult butterfly wing patterns. Results To obtain a catalog of genes involved in larval mimetic and cryptic pattern formation, we constructed expressed sequence tag (EST) libraries of larval epidermis for P. xuthus, and P. polytes that contained 20,736 and 5,376 clones, respectively, representing one of the largest collections available in butterflies. A comparison with silkworm epidermal EST information revealed the high expression of putative blue and yellow pigment-binding proteins in Papilio species. We also designed a microarray from the EST dataset information, analyzed more than five stages each for six markings, and confirmed spatial expression patterns by whole-mount in situ hybridization. Hence, we succeeded in elucidating many novel marking-specific genes for mimetic and cryptic pattern formation, including pigment-binding protein genes, the melanin-associated gene yellow-h3, the ecdysteroid synthesis enzyme gene 3-dehydroecdysone 3b-reductase, and Papilio-specific genes. We also found many cuticular protein genes with marking specificity that may be associated with the unique surface nanostructure of the markings. Furthermore, we identified two transcription factors, spalt and ecdysteroid signal-related E75, as genes expressed in larval eyespot markings. This finding suggests that E75 is a strong candidate mediator of the hormone-dependent coordination of larval pattern formation. Conclusions This study is one of the most comprehensive molecular analyses of complicated morphological features, and it will serve as a new resource for studying insect mimetic and cryptic pattern formation in general. The wide variety of marking-associated genes (both regulatory and structural genes) identified by our screening indicates that a similar strategy will be effective for understanding other complex traits. PMID:22651552

  9. Structural color mechanism in the Papilio blumei butterfly.

    PubMed

    Lo, Mei-Ling; Lee, Cheng-Chung

    2014-02-01

    The structural color found in biological systems has complicated nanostructure. It is very important to determine its color mechanism. In this study, the 2D photonic crystal structures of the Papilio blumei butterfly were constructed, and the corresponding reflectance spectra were simulated by the finite-difference time-domain method. The structural color of the butterfly depends on the incident angle of light, film thickness, film material (film refractive index), and the size of the air hole (effective refractive index). Analysis of simulations can help us understand the hue, brightness, and saturation of structural color on the butterfly wing. As a result, the analysis can help us fabricate expected structural color.

  10. Marginal eyespots on butterfly wings deflect bird attacks under low light intensities with UV wavelengths.

    PubMed

    Olofsson, Martin; Vallin, Adrian; Jakobsson, Sven; Wiklund, Christer

    2010-05-24

    Predators preferentially attack vital body parts to avoid prey escape. Consequently, prey adaptations that make predators attack less crucial body parts are expected to evolve. Marginal eyespots on butterfly wings have long been thought to have this deflective, but hitherto undemonstrated function. Here we report that a butterfly, Lopinga achine, with broad-spectrum reflective white scales in its marginal eyespot pupils deceives a generalist avian predator, the blue tit, to attack the marginal eyespots, but only under particular conditions-in our experiments, low light intensities with a prominent UV component. Under high light intensity conditions with a similar UV component, and at low light intensities without UV, blue tits directed attacks towards the butterfly head. In nature, birds typically forage intensively at early dawn, when the light environment shifts to shorter wavelengths, and the contrast between the eyespot pupils and the background increases. Among butterflies, deflecting attacks is likely to be particularly important at dawn when low ambient temperatures make escape by flight impossible, and when insectivorous birds typically initiate another day's search for food. Our finding that the deflective function of eyespots is highly dependent on the ambient light environment helps explain why previous attempts have provided little support for the deflective role of marginal eyespots, and we hypothesize that the mechanism that we have discovered in our experiments in a laboratory setting may function also in nature when birds forage on resting butterflies under low light intensities.

  11. Hyperspectral optical imaging of two different species of lepidoptera

    PubMed Central

    2011-01-01

    In this article, we report a hyperspectral optical imaging application for measurement of the reflectance spectra of photonic structures that produce structural colors with high spatial resolution. The measurement of the spectral reflectance function is exemplified in the butterfly wings of two different species of Lepidoptera: the blue iridescence reflected by the nymphalid Morpho didius and the green iridescence of the papilionid Papilio palinurus. Color coordinates from reflectance spectra were calculated taking into account human spectral sensitivity. For each butterfly wing, the observed color is described by a characteristic color map in the chromaticity diagram and spreads over a limited volume in the color space. The results suggest that variability in the reflectance spectra is correlated with different random arrangements in the spatial distribution of the scales that cover the wing membranes. Hyperspectral optical imaging opens new ways for the non-invasive study and classification of different forms of irregularity in structural colors. PMID:21711872

  12. Inverse Mopho butterfly: a new approach to photonic crystal.

    PubMed

    Zhang, Ji-Zhong; Gu, Zhong-Ze; Chen, Hai-Hua; Fujishima, Akira; Sato, Osamu

    2006-04-01

    In this paper we showed a new approach for the fabrication of a photonic crystal with a three-dimensional structure. By replicating biomaterials such as the wing of Mopho butterfly with TiO2 nanoparticles using the nanoparticles infiltration method, we can derive photonic crystals with unique structures, which is difficult to fabricate by other approaches. New optical properties are anticipated.

  13. Behavioural thermoregulation and the relative roles of convection and radiation in a basking butterfly.

    PubMed

    Barton, Madeleine; Porter, Warren; Kearney, Michael

    2014-04-01

    Poikilothermic animals are often reliant on behavioural thermoregulation to elevate core-body temperature above the temperature of their surroundings. Butterflies are able to do this by altering body posture and location while basking, however the specific mechanisms that achieve such regulation vary among species. The role of the wings has been particularly difficult to describe, with uncertainty surrounding whether they are positioned to reduce convective heat loss or to maximise heat gained through radiation. Characterisation of the extent to which these processes affect core-body temperature will provide insights into the way in which a species׳ thermal sensitivity and morphological traits have evolved. We conducted field and laboratory measurements to assess how basking posture affects the core-body temperature of an Australian butterfly, the common brown (Heteronympha merope). We show that, with wings held open, heat lost through convection is reduced while heat gained through radiation is simultaneously maximised. These responses have been incorporated into a biophysical model that accurately predicts the core-body temperature of basking specimens in the field, providing a powerful tool to explore how climate constrains the distribution and abundance of basking butterflies. Copyright © 2014 Elsevier Ltd. All rights reserved.

  14. The role of random nanostructures for the omnidirectional anti-reflection properties of the glasswing butterfly.

    PubMed

    Siddique, Radwanul Hasan; Gomard, Guillaume; Hölscher, Hendrik

    2015-04-22

    The glasswing butterfly (Greta oto) has, as its name suggests, transparent wings with remarkable low haze and reflectance over the whole visible spectral range even for large view angles of 80°. This omnidirectional anti-reflection behaviour is caused by small nanopillars covering the transparent regions of its wings. In difference to other anti-reflection coatings found in nature, these pillars are irregularly arranged and feature a random height and width distribution. Here we simulate the optical properties with the effective medium theory and transfer matrix method and show that the random height distribution of pillars significantly reduces the reflection not only for normal incidence but also for high view angles.

  15. Bioinspired superhydrophobic, self-cleaning and low drag surfaces

    NASA Astrophysics Data System (ADS)

    Bhushan, Bharat

    2013-09-01

    Nature has evolved objects with desired functionality using commonly found materials. Nature capitalizes on hierarchical structures to achieve functionality. The understanding of the functions provided by objects and processes found in nature can guide us to produce nanomaterials, nanodevices, and processes with desirable functionality. This article provides an overview of four topics: (1) Lotus Effect used to develop superhydrophobic and self-cleaning/antifouling surfaces with low adhesion, (2) Shark Skin Effect to develop surfaces with low fluid drag and anti-fouling characteristics, and (3-4) Rice Leaf and Butterfly Wing Effect to develop superhydrophobic and self-cleaning surfaces with low drag. Rice Leaf and Butterfly Wings combine the Shark Skin and Lotus Effects.

  16. The role of random nanostructures for the omnidirectional anti-reflection properties of the glasswing butterfly

    NASA Astrophysics Data System (ADS)

    Siddique, Radwanul Hasan; Gomard, Guillaume; Hölscher, Hendrik

    2015-04-01

    The glasswing butterfly (Greta oto) has, as its name suggests, transparent wings with remarkable low haze and reflectance over the whole visible spectral range even for large view angles of 80°. This omnidirectional anti-reflection behaviour is caused by small nanopillars covering the transparent regions of its wings. In difference to other anti-reflection coatings found in nature, these pillars are irregularly arranged and feature a random height and width distribution. Here we simulate the optical properties with the effective medium theory and transfer matrix method and show that the random height distribution of pillars significantly reduces the reflection not only for normal incidence but also for high view angles.

  17. Biotemplated Morpho Butterfly Wings for Tunable Structurally Colored Photocatalysts.

    PubMed

    Rodríguez, Robin E; Agarwal, Sneha P; An, Shun; Kazyak, Eric; Das, Debashree; Shang, Wen; Skye, Rachael; Deng, Tao; Dasgupta, Neil P

    2018-02-07

    Morpho sulkowskyi butterfly wings contain naturally occurring hierarchical nanostructures that produce structural coloration. The high aspect ratio and surface area of these wings make them attractive nanostructured templates for applications in solar energy and photocatalysis. However, biomimetic approaches to replicate their complex structural features and integrate functional materials into their three-dimensional framework are highly limited in precision and scalability. Herein, a biotemplating approach is presented that precisely replicates Morpho nanostructures by depositing nanocrystalline ZnO coatings onto wings via low-temperature atomic layer deposition (ALD). This study demonstrates the ability to precisely tune the natural structural coloration while also integrating multifunctionality by imparting photocatalytic activity onto fully intact Morpho wings. Optical spectroscopy and finite-difference time-domain numerical modeling demonstrate that ALD ZnO coatings can rationally tune the structural coloration across the visible spectrum. These structurally colored photocatalysts exhibit an optimal coating thickness to maximize photocatalytic activity, which is attributed to trade-offs between light absorption and catalytic quantum yield with increasing coating thickness. These multifunctional photocatalysts present a new approach to integrating solar energy harvesting into visually attractive surfaces that can be integrated into building facades or other macroscopic structures to impart aesthetic appeal.

  18. The chiral structure of porous chitin within the wing-scales of Callophrys rubi.

    PubMed

    Schröder-Turk, G E; Wickham, S; Averdunk, H; Brink, F; Fitz Gerald, J D; Poladian, L; Large, M C J; Hyde, S T

    2011-05-01

    The structure of the porous three-dimensional reticulated pattern in the wing scales of the butterfly Callophrys rubi (the Green Hairstreak) is explored in detail, via scanning and transmission electron microscopy. A full 3D tomographic reconstruction of a section of this material reveals that the predominantly chitin material is assembled in the wing scale to form a structure whose geometry bears a remarkable correspondence to the srs net, well-known in solid state chemistry and soft materials science. The porous solid is bounded to an excellent approximation by a parallel surface to the Gyroid, a three-periodic minimal surface with cubic crystallographic symmetry I4₁32, as foreshadowed by Stavenga and Michielson. The scale of the structure is commensurate with the wavelength of visible light, with an edge of the conventional cubic unit cell of the parallel-Gyroid of approximately 310 nm. The genesis of this structure is discussed, and we suggest it affords a remarkable example of templating of a chiral material via soft matter, analogous to the formation of mesoporous silica via surfactant assemblies in solution. In the butterfly, the templating is achieved by the lipid-protein membranes within the smooth endoplasmic reticulum (while it remains in the chrysalis), that likely form cubic membranes, folded according to the form of the Gyroid. The subsequent formation of the chiral hard chitin framework is suggested to be driven by the gradual polymerisation of the chitin precursors, whose inherent chiral assembly in solution (during growth) promotes the formation of a single enantiomer. Copyright © 2011 Elsevier Inc. All rights reserved.

  19. The effect of butterfly-scale inspired patterning on leading-edge vortex growth

    NASA Astrophysics Data System (ADS)

    Wilroy, Jacob Aaron

    Leading edge vortices (LEVs) are important for generating thrust and lift in flapping flight, and the surface patterning (scales) on butterfly wings is hypothesized to play a role in the vortex formation of the LEV. To simplify this complex flow problem, an experiment was designed to focus on the alteration of 2-D vortex development with a variation in surface patterning. Specifically, the secondary vorticity generated by the LEV interacting at the patterned surface was studied, as well as the subsequent effect on the LEV's growth rate and peak circulation. For this experiment, rapid-prototyped grooves based on the scale geometry of the Monarch butterfly (Danaus plexippus) were created using additive manufacturing and were attached to a flat plate with a chordwise orientation, thus increasing plate surface area. The vortex generated by the grooved plate was then compared to a smooth plate case in an experiment where the plate translated vertically through a 2 x 3 x 5 cubic foot tow tank. The plate was impulsively started in quiescent water and flow fields at Rec = 1416, 2833, and 5667 are examined using Digital Particle Image Velocimetry (DPIV). The maximum vortex formation number is 2.8 and is based on the flat plate travel length and chord length. Flow fields from each case show the generation of a secondary vortex whose interaction with the shear layer and LEV caused different behaviors depending upon the surface type. The vortex development process varied for each Reynolds number and it was found that for the lowest Reynolds number case a significant difference does not exist between surface types, however, for the other two cases the grooves affected the secondary vortex's behavior and the LEV's ability to grow at a rate similar to the smooth plate case.

  20. Mysterious coloring: structural origin of color mixing for two breeds of Papilio butterflies.

    PubMed

    Diao, Ying-Ying; Liu, Xiang-Yang

    2011-05-09

    The structural origin of the coloration mechanisms and related extraordinary optical properties of the wing scales of two breeds of Papilio butterflies, namely, Papilio ulysses and Papilio blumei, are explored. The precise ordered biophotonic nanostructures of the wing scales are characterized by scanning electron microscopy (SEM). Despite their structural similarities, the two breeds of Papilio butterflies do not exhibit any analogy in their optical performances. When illuminated with UV-Vis light, P. ulysses gives rise to two reflection peaks: one is from concavities, and the other is from ridges. These two spectral peaks shift their positions under different illumination angles (normal and 45° incident light). In contrast, the spectra for the green scales of P. blumei give one broad reflection peak, and the peak remains the same under normal and 45° incident light. The optical microscopy images indicate that the cap-shaped concavities on P. blumei's wing scales generate an abnormal bicolor reflection with a strong polarization effect. Both of these two breeds of butterflies take advantage of color mixing strategy: the blue color of P. ulysses is mixed by the colors reflected from concavities and ridges; the green color of P. blumei is produced by the biocolor reflection from concavities. The differences of their coloration mixing mechanisms and optical performances are due to the variations of their nanostructures. The investigation of the color mixing mechanisms of these biologically photonic nanostructures may offer a convenient way for fabricating optical devices based on biomimicry. © 2011 Optical Society of America

  1. Closed-Loop Control of Constrained Flapping Wing Micro Air Vehicles

    DTIC Science & Technology

    2014-03-27

    insects , thus concealing their appearance while also providing benefits of unsteady aerodynamics. Consider- able research has been invested in the...small visibility signature that tends to hide in plain sight by resembling insects . 1.2 Research Challenges for Flapping Wing Micro Air Vehicles There are...predicts forces and moments for the class of flapping wing fliers that makes up most insects and hummingbirds. Large bird and butterfly “clap- and

  2. On the colour of wing scales in butterflies: iridescence and preferred orientation of single gyroid photonic crystals

    PubMed Central

    2017-01-01

    Lycaenid butterflies from the genera Callophrys, Cyanophrys and Thecla have evolved remarkable biophotonic gyroid nanostructures within their wing scales that have only recently been replicated by nanoscale additive manufacturing. These nanostructures selectively reflect parts of the visible spectrum to give their characteristic non-iridescent, matte-green appearance, despite a distinct blue–green–yellow iridescence predicted for individual crystals from theory. It has been hypothesized that the organism must achieve its uniform appearance by growing crystals with some restrictions on the possible distribution of orientations, yet preferential orientation observed in Callophrys rubi confirms that this distribution need not be uniform. By analysing scanning electron microscope and optical images of 912 crystals in three wing scales, we find no preference for their rotational alignment in the plane of the scales. However, crystal orientation normal to the scale was highly correlated to their colour at low (conical) angles of view and illumination. This correlation enabled the use of optical images, each containing up to 104–105 crystals, for concluding the preferential alignment seen along the at the level of single scales, appears ubiquitous. By contrast, orientations were found to occur at no greater rate than that expected by chance. Above a critical cone angle, all crystals reflected bright green light indicating the dominant light scattering is due to the predicted band gap along the direction, independent of the domain orientation. Together with the natural variation in scale and wing shapes, we can readily understand the detailed mechanism of uniform colour production and iridescence suppression in these butterflies. It appears that the combination of preferential alignment normal to the wing scale, and uniform distribution within the plane is a near optimal solution for homogenizing the angular distribution of the band gap relative to the wings. Finally, the distributions of orientations, shapes, sizes and degree of order of crystals within single scales provide useful insights for understanding the mechanisms at play in the formation of these biophotonic nanostructures. PMID:28630678

  3. Electron tomography and computer visualisation of a three-dimensional 'photonic' crystal in a butterfly wing-scale.

    PubMed

    Argyros, A; Manos, S; Large, M C J; McKenzie, D R; Cox, G C; Dwarte, D M

    2002-01-01

    A combination of transmission electron tomography and computer modelling has been used to determine the three-dimensional structure of the photonic crystals found in the wing-scales of the Kaiser-I-Hind butterfly (Teinopalpus imperialis). These scales presented challenges for electron microscopy because the periodicity of the structure was comparable to the thickness of a section and because of the complex connectivity of the object. The structure obtained has been confirmed by taking slices of the three-dimensional computer model constructed from the tomography and comparing these with transmission electron microscope (TEM) images of microtomed sections of the actual scale. The crystal was found to have chiral tetrahedral repeating units packed in a triclinic lattice.

  4. Can butterflies cope with city life? Butterfly diversity in a young megacity in southern China.

    PubMed

    Sing, Kong-Wah; Dong, Hui; Wang, Wen-Zhi; Wilson, John-James

    2016-09-01

    During 30 years of unprecedented urbanization, plant diversity in Shenzhen, a young megacity in southern China, has increased dramatically. Although strongly associated with plant diversity, butterfly diversity generally declines with urbanization, but this has not been investigated in Shenzhen. Considering the speed of urbanization in Shenzhen and the large number of city parks, we investigated butterfly diversity in Shenzhen parks. We measured butterfly species richness in four microhabitats (groves, hedges, flowerbeds, and unmanaged areas) across 10 parks and examined the relationship with three park variables: park age, park size, and distance from the central business district. Butterflies were identified based on wing morphology and DNA barcoding. We collected 1933 butterflies belonging to 74 species from six families; 20% of the species were considered rare. Butterfly species richness showed weak negative correlations with park age and distance from the central business district, but the positive correlation with park size was statistically significant (p = 0.001). Among microhabitat types, highest species richness was recorded in unmanaged areas. Our findings are consistent with others in suggesting that to promote urban butterfly diversity it is necessary to make parks as large as possible and to set aside areas for limited management. In comparison to neighbouring cities, Shenzhen parks have high butterfly diversity.

  5. UV reflectance is associated with environmental conditions in Palaearctic Pieris napi (Lepidoptera: Pieridae).

    PubMed

    Stella, David; Pecháček, Pavel; Meyer-Rochow, Victor Benno; Kleisner, Karel

    2018-06-01

    The subject of our investigation was the visual features of wing color with special focus on the UV reflectance in the green-veined white butterfly (Pieris napi). Previous studies had concluded that UV reflectance on dorsal wing surfaces is found only in the female P. napi. Based on UV sensitive photography, we analyzed a correlation between 12 geographic and environmental factors and UV reflectance patterns on 3 patches on the forewings of 407 P. napi specimens from the Palaearctic region. Results had shown that females significantly differ from males: they exhibit a 25% higher UV reflectance. To investigate whether and how UV reflectance levels on the forewings and hindwings of both sexes are influenced by the environment, we performed a principal component analysis (PCA) with several environmental variables. For several variables (in particular, latitude and longitude, mean annual temperature and precipitation, and temperature annual range and altitude), the generalized linear model (GLM) model revealed a significant correlation in both sexes. This suggests a link between UV reflectance levels and the environment and distribution of P. napi. We found that stronger UV reflectance is associated with generally more hostile environments and concluded that large-scale environmental factors influence the UV reflectance on the forewings of both male and female green-veined white butterflies. © 2016 Institute of Zoology, Chinese Academy of Sciences.

  6. Butterfly Eyespots: Their Potential Influence on Aesthetic Preferences and Conservation Attitudes

    PubMed Central

    Manesi, Zoi; Van Lange, Paul A. M.; Pollet, Thomas V.

    2015-01-01

    Research has shown that the mere presence of stimuli that resemble eyes is sufficient to attract attention, elicit aesthetic responses, and can even enhance prosocial behavior. However, it is less clear whether eye-like stimuli could also be used as a tool for nature conservation. Several animal species, including butterflies, develop eye-like markings that are known as eyespots. In the present research, we explored whether the mere display of eyespots on butterfly wings can enhance: (a) liking for a butterfly species, and (b) attitudes and behaviors towards conservation of a butterfly species. Four online experimental studies, involving 613 participants, demonstrated that eyespots significantly increased liking for a butterfly species. Furthermore, eyespots significantly increased positive attitudes towards conservation of a butterfly species (Studies 1, 2 and 4), whereas liking mediated the eyespot effect on conservation attitudes (Study 2). However, we also found some mixed evidence for an association between eyespots and actual conservation behavior (Studies 3 and 4). Overall, these findings suggest that eyespots may increase liking for an animal and sensitize humans to conservation. We discuss possible implications for biodiversity conservation and future research directions. PMID:26544692

  7. Butterfly Eyespots: Their Potential Influence on Aesthetic Preferences and Conservation Attitudes.

    PubMed

    Manesi, Zoi; Van Lange, Paul A M; Pollet, Thomas V

    2015-01-01

    Research has shown that the mere presence of stimuli that resemble eyes is sufficient to attract attention, elicit aesthetic responses, and can even enhance prosocial behavior. However, it is less clear whether eye-like stimuli could also be used as a tool for nature conservation. Several animal species, including butterflies, develop eye-like markings that are known as eyespots. In the present research, we explored whether the mere display of eyespots on butterfly wings can enhance: (a) liking for a butterfly species, and (b) attitudes and behaviors towards conservation of a butterfly species. Four online experimental studies, involving 613 participants, demonstrated that eyespots significantly increased liking for a butterfly species. Furthermore, eyespots significantly increased positive attitudes towards conservation of a butterfly species (Studies 1, 2 and 4), whereas liking mediated the eyespot effect on conservation attitudes (Study 2). However, we also found some mixed evidence for an association between eyespots and actual conservation behavior (Studies 3 and 4). Overall, these findings suggest that eyespots may increase liking for an animal and sensitize humans to conservation. We discuss possible implications for biodiversity conservation and future research directions.

  8. Differences in the selection response of serially repeated color pattern characters: standing variation, development, and evolution.

    PubMed

    Allen, Cerisse E; Beldade, Patrícia; Zwaan, Bas J; Brakefield, Paul M

    2008-03-26

    There is spectacular morphological diversity in nature but lineages typically display a limited range of phenotypes. Because developmental processes generate the phenotypic variation that fuels natural selection, they are a likely source of evolutionary biases, facilitating some changes and limiting others. Although shifts in developmental regulation are associated with morphological differences between taxa, it is unclear how underlying mechanisms affect the rate and direction of evolutionary change within populations under selection. Here we focus on two ecologically relevant features of butterfly wing color patterns, eyespot size and color composition, which are similarly and strongly correlated across the serially repeated eyespots. Though these two characters show similar patterns of standing variation and covariation within a population, they differ in key features of their underlying development. We targeted pairs of eyespots with artificial selection for coordinated (concerted selection) versus independent (antagonistic selection) change in their color composition and size and compared evolutionary responses of the two color pattern characters. The two characters respond to selection in strikingly different ways despite initially similar patterns of variation in all directions present in the starting population. Size (determined by local properties of a diffusing inductive signal) evolves flexibly in all selected directions. However, color composition (determined by a tissue-level response to the signal concentration gradient) evolves only in the direction of coordinated change. There was no independent evolutionary change in the color composition of two eyespots in response to antagonistic selection. Moreover, these differences in the directions of short-term evolutionary change in eyespot size and color composition within a single species are consistent with the observed wing pattern diversity in the genus. Both characters respond rapidly to selection for coordinated change, but there are striking differences in their response to selection for antagonistic, independent change across eyespots. While many additional factors may contribute to both short- and long-term evolutionary response, we argue that the compartmentalization of developmental processes can influence the diversification of serial repeats such as butterfly eyespots, even under strong selection.

  9. Red & black or black & white? Phylogeny of the Araschnia butterflies (Lepidoptera: Nymphalidae) and evolution of seasonal polyphenism.

    PubMed

    Fric, Z; Konvicka, M; Zrzavy, J

    2004-03-01

    Phylogeny of the butterfly genera Araschnia, Mynes, Symbrenthia and Brensymthia (Lepidoptera: Nymphalidae: Nymphalini) is reconstructed, based on 140 morphological and ecological characters. The resulting tree shows that Araschnia is a sister group of the clade including Symbrenthia, Mynes and Brensymthia (Symbrenthia is paraphyletic in the respect of remaining genera; Symbrenthia hippalus is a derived species of Mynes). The species-level relationships within Araschnia are robustly supported as follows: (A. davidis (prorsoides ((zhangi doris) (dohertyi (levana burejana))))). Analysis of the wing colour-pattern characters linked with the seasonal polyphenism in the Araschnia species suggests that the black and white coloration of the long-day (summer) generation is apomorphic. Biogeographically, the origin of polyphenism in Araschnia predates the dispersal of some Araschnia species towards the Palaearctic temperate zone, and the ecological cause of the polyphenism itself is then probably not linked with thermoregulation. The possible mimetic/cryptic scenarios for the origin of Araschnia polyphenism are discussed.

  10. Eyespots deflect predator attack increasing fitness and promoting the evolution of phenotypic plasticity

    PubMed Central

    Prudic, Kathleen L.; Stoehr, Andrew M.; Wasik, Bethany R.; Monteiro, Antónia

    2015-01-01

    Some eyespots are thought to deflect attack away from the vulnerable body, yet there is limited empirical evidence for this function and its adaptive advantage. Here, we demonstrate the conspicuous ventral hindwing eyespots found on Bicyclus anynana butterflies protect against invertebrate predators, specifically praying mantids. Wet season (WS) butterflies with larger, brighter eyespots were easier for mantids to detect, but more difficult to capture compared to dry season (DS) butterflies with small, dull eyespots. Mantids attacked the wing eyespots of WS butterflies more frequently resulting in greater butterfly survival and reproductive success. With a reciprocal eyespot transplant, we demonstrated the fitness benefits of eyespots were independent of butterfly behaviour. Regardless of whether the butterfly was WS or DS, large marginal eyespots pasted on the hindwings increased butterfly survival and successful oviposition during predation encounters. In previous studies, DS B. anynana experienced delayed detection by vertebrate predators, but both forms suffered low survival once detected. Our results suggest predator abundance, identity and phenology may all be important selective forces for B. anynana. Thus, reciprocal selection between invertebrate and vertebrate predators across seasons may contribute to the evolution of the B. anynana polyphenism. PMID:25392465

  11. Yellow and the Novel Aposematic Signal, Red, Protect Delias Butterflies from Predators

    PubMed Central

    Wee, Jocelyn Liang Qi

    2017-01-01

    Butterflies of the South Asian and Australian genus Delias possess striking colours on the ventral wings that are presumed to serve as warning signals to predators. However, this has not been shown empirically. Here we experimentally tested whether the colours of one member of this diverse genus, Delias hyparete, function as aposematic signals. We constructed artificial paper models with either a faithful colour representation of D. hyparete, or with all of its colours converted to grey scale. We also produced models where single colours were left intact, while others were converted to grey-scale or removed entirely. We placed all model types simultaneously in the field, attached to a live mealworm, and measured relative attack rates at three separate field sites. Faithful models of D. hyparete, suffered the least amount of attacks, followed by grey-scale models with unaltered red patches, and by grey-scale models with unaltered yellow patches. We conclude that red and yellow colours function as warning signals. By mapping dorsal and ventral colouration onto a phylogeny of Delias, we observed that yellow and red colours appear almost exclusively on the ventral wing surfaces, and that basal lineages have mostly yellow, white, and black wings, whereas derived lineages contain red colour in addition to the other colours. Red appears to be, thus, a novel adaptive trait in this lineage of butterflies. PMID:28060944

  12. Yellow and the Novel Aposematic Signal, Red, Protect Delias Butterflies from Predators.

    PubMed

    Wee, Jocelyn Liang Qi; Monteiro, Antónia

    2017-01-01

    Butterflies of the South Asian and Australian genus Delias possess striking colours on the ventral wings that are presumed to serve as warning signals to predators. However, this has not been shown empirically. Here we experimentally tested whether the colours of one member of this diverse genus, Delias hyparete, function as aposematic signals. We constructed artificial paper models with either a faithful colour representation of D. hyparete, or with all of its colours converted to grey scale. We also produced models where single colours were left intact, while others were converted to grey-scale or removed entirely. We placed all model types simultaneously in the field, attached to a live mealworm, and measured relative attack rates at three separate field sites. Faithful models of D. hyparete, suffered the least amount of attacks, followed by grey-scale models with unaltered red patches, and by grey-scale models with unaltered yellow patches. We conclude that red and yellow colours function as warning signals. By mapping dorsal and ventral colouration onto a phylogeny of Delias, we observed that yellow and red colours appear almost exclusively on the ventral wing surfaces, and that basal lineages have mostly yellow, white, and black wings, whereas derived lineages contain red colour in addition to the other colours. Red appears to be, thus, a novel adaptive trait in this lineage of butterflies.

  13. Poor sequestration of toxic host plant cardenolides and their rapid loss in the milkweed butterfly Danaus chrysippus (Lepidoptera: Nymphalidae: Danainae: Danaini).

    PubMed

    Mebs, Dietrich; Wunder, Cora; Toennes, Stefan W

    2017-06-01

    Butterflies of the genus Danaus are known to sequester toxic cardenolides from milkweed host plants (Apocynaceae). In particular, Danaus plexippus efficiently sequesters and stores these compounds, whereas D. chrysippus, is considered to poorly sequester cardenolides. To estimate its sequestration capability compared with that of D. plexippus, larvae of both species were jointly reared on Asclepias curassavica and the major cardenolides of the host plant, calotropin and calactin, were analyzed in adults sampled at different time intervals after eclosion. Both cardenolides were detected in body and wings of D. plexippus. Whereas the calotropin-concentration remained constant over a period of 24 days, that of calactin steadily decreased. In the body, but not in the wings of D. chrysippus, calactin only was detected in low amounts, which was then almost completely lost during the following 8 days after eclosion, suggesting that in contrast to D. plexippus, cardenolides seem to be less important for that butterfly's defence against predators. Copyright © 2017 Elsevier Ltd. All rights reserved.

  14. Pterin pigments amplify iridescent ultraviolet signal in males of the orange sulphur butterfly, Colias eurytheme

    PubMed Central

    Rutowski, R.L; Macedonia, J.M; Morehouse, N; Taylor-Taft, L

    2005-01-01

    Animal colouration is typically the product of nanostructures that reflect or scatter light and pigments that absorb it. The interplay between these colour-producing mechanisms may influence the efficacy and potential information content of colour signals, but this notion has received little empirical attention. Wing scales in the male orange sulphur butterfly (Colias eurytheme) possess ridges with lamellae that produce a brilliant iridescent ultraviolet (UV) reflectance via thin-film interference. Curiously, these same scales contain pterin pigments that strongly absorb wavelengths below 550 nm. Given that male UV reflectance functions as a sexual signal in C. eurytheme, it is paradoxical that pigments in the wing scales are highly UV absorbing. We present spectrophotometric analyses of the wings before and after pterin removal that show that pterins both depress the amplitude of UV iridescence and suppress a diffuse UV reflectance that emanates from the scales. This latter effect enhances the directionality and spectral purity of the iridescence, and increases the signal's chromaticity and potential signal content. Our findings also suggest that pterins amplify the contrast between iridescent UV reflectance and scale background colour as a male's wings move during flight. PMID:16191648

  15. Pterin pigments amplify iridescent ultraviolet signal in males of the orange sulphur butterfly, Colias eurytheme.

    PubMed

    Rutowski, R L; Macedonia, J M; Morehouse, N; Taylor-Taft, L

    2005-11-07

    Animal colouration is typically the product of nanostructures that reflect or scatter light and pigments that absorb it. The interplay between these colour-producing mechanisms may influence the efficacy and potential information content of colour signals, but this notion has received little empirical attention. Wing scales in the male orange sulphur butterfly (Colias eurytheme) possess ridges with lamellae that produce a brilliant iridescent ultraviolet (UV) reflectance via thin-film interference. Curiously, these same scales contain pterin pigments that strongly absorb wavelengths below 550 nm. Given that male UV reflectance functions as a sexual signal in C. eurytheme, it is paradoxical that pigments in the wing scales are highly UV absorbing. We present spectrophotometric analyses of the wings before and after pterin removal that show that pterins both depress the amplitude of UV iridescence and suppress a diffuse UV reflectance that emanates from the scales. This latter effect enhances the directionality and spectral purity of the iridescence, and increases the signal's chromaticity and potential signal content. Our findings also suggest that pterins amplify the contrast between iridescent UV reflectance and scale background colour as a male's wings move during flight.

  16. Coherent array of branched filamentary scales along the wing margin of a small moth

    NASA Astrophysics Data System (ADS)

    Yoshida, Akihiro; Tejima, Shin; Sakuma, Masayuki; Sakamaki, Yositaka; Kodama, Ryuji

    2017-04-01

    In butterflies and moths, the wing margins are fringed with specialized scales that are typically longer than common scales. In the hindwings of some small moths, the posterior margins are fringed with particularly long filamentary scales. Despite the small size of these moth wings, these scales are much longer than those of large moths and butterflies. In the current study, photography of the tethered flight of a small moth, Phthorimaea operculella, revealed a wide array composed of a large number of long filamentary scales. This array did not become disheveled in flight, maintaining a coherent sheet-like structure during wingbeat. Examination of the morphology of individual scales revealed that each filamentary scale consists of a proximal stalk and distal branches. Moreover, not only long scales but also shorter scales of various lengths were found to coexist in each small section of the wing margin. Scale branches were ubiquitously and densely distributed within the scale array to form a mesh-like architecture similar to a nonwoven fabric. We propose that possible mechanical interactions among branched filamentary scales, mediated by these branches, may contribute to maintaining a coherent sheet-like structure of the scale array during wingbeat.

  17. Climate variability slows evolutionary responses of Colias butterflies to recent climate change.

    PubMed

    Kingsolver, Joel G; Buckley, Lauren B

    2015-03-07

    How does recent climate warming and climate variability alter fitness, phenotypic selection and evolution in natural populations? We combine biophysical, demographic and evolutionary models with recent climate data to address this question for the subalpine and alpine butterfly, Colias meadii, in the southern Rocky Mountains. We focus on predicting patterns of selection and evolution for a key thermoregulatory trait, melanin (solar absorptivity) on the posterior ventral hindwings, which affects patterns of body temperature, flight activity, adult and egg survival, and reproductive success in Colias. Both mean annual summer temperatures and thermal variability within summers have increased during the past 60 years at subalpine and alpine sites. At the subalpine site, predicted directional selection on wing absorptivity has shifted from generally positive (favouring increased wing melanin) to generally negative during the past 60 years, but there is substantial variation among years in the predicted magnitude and direction of selection and the optimal absorptivity. The predicted magnitude of directional selection at the alpine site declined during the past 60 years and varies substantially among years, but selection has generally been positive at this site. Predicted evolutionary responses to mean climate warming at the subalpine site since 1980 is small, because of the variability in selection and asymmetry of the fitness function. At both sites, the predicted effects of adaptive evolution on mean population fitness are much smaller than the fluctuations in mean fitness due to climate variability among years. Our analyses suggest that variation in climate within and among years may strongly limit evolutionary responses of ectotherms to mean climate warming in these habitats. © 2015 The Author(s) Published by the Royal Society. All rights reserved.

  18. Sexual Dimorphism and Retinal Mosaic Diversification following the Evolution of a Violet Receptor in Butterflies.

    PubMed

    McCulloch, Kyle J; Yuan, Furong; Zhen, Ying; Aardema, Matthew L; Smith, Gilbert; Llorente-Bousquets, Jorge; Andolfatto, Peter; Briscoe, Adriana D

    2017-09-01

    Numerous animal lineages have expanded and diversified the opsin-based photoreceptors in their eyes underlying color vision behavior. However, the selective pressures giving rise to new photoreceptors and their spectral tuning remain mostly obscure. Previously, we identified a violet receptor (UV2) that is the result of a UV opsin gene duplication specific to Heliconius butterflies. At the same time the violet receptor evolved, Heliconius evolved UV-yellow coloration on their wings, due to the pigment 3-hydroxykynurenine (3-OHK) and the nanostructure architecture of the scale cells. In order to better understand the selective pressures giving rise to the violet receptor, we characterized opsin expression patterns using immunostaining (14 species) and RNA-Seq (18 species), and reconstructed evolutionary histories of visual traits in five major lineages within Heliconius and one species from the genus Eueides. Opsin expression patterns are hyperdiverse within Heliconius. We identified six unique retinal mosaics and three distinct forms of sexual dimorphism based on ommatidial types within the genus Heliconius. Additionally, phylogenetic analysis revealed independent losses of opsin expression, pseudogenization events, and relaxation of selection on UVRh2 in one lineage. Despite this diversity, the newly evolved violet receptor is retained across most species and sexes surveyed. Discriminability modeling of behaviorally preferred 3-OHK yellow wing coloration suggests that the violet receptor may facilitate Heliconius color vision in the context of conspecific recognition. Our observations give insights into the selective pressures underlying the origins of new visual receptors. © The Author 2017. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.

  19. Blame it on the butterfly

    NASA Astrophysics Data System (ADS)

    Oliver, Kate

    2009-08-01

    Last year at a science networking event in a Central London pub, I was cornered by the manager of an "alternative healing centre", who regaled me with stories about her patients' miraculous recoveries from ailments that modern medicine had been unable to address. "After all," she said, leaning forward with the air of someone confiding an esoteric, but unassailable, argument, "if a butterfly flapping its wings in a forest can cause a hurricane, imagine what a positive attitude can do!"

  20. Force balance in the take-off of a pierid butterfly: relative importance and timing of leg impulsion and aerodynamic forces.

    PubMed

    Bimbard, Gaëlle; Kolomenskiy, Dmitry; Bouteleux, Olivier; Casas, Jérôme; Godoy-Diana, Ramiro

    2013-09-15

    Up to now, the take-off stage has remained an elusive phase of insect flight that was relatively poorly explored compared with other maneuvers. An overall assessment of the different mechanisms involved in force production during take-off has never been explored. Focusing on the first downstroke, we have addressed this problem from a force balance perspective in butterflies taking off from the ground. In order to determine whether the sole aerodynamic wing force could explain the observed motion of the insect, we have firstly compared a simple analytical model of the wing force with the acceleration of the insect's center of mass estimated from video tracking of the wing and body motions. Secondly, wing kinematics were also used for numerical simulations of the aerodynamic flow field. Similar wing aerodynamic forces were obtained by the two methods. However, neither are sufficient, nor is the inclusion of the ground effect, to predict faithfully the body acceleration. We have to resort to the leg forces to obtain a model that best fits the data. We show that the median and hind legs display an active extension responsible for the initiation of the upward motion of the insect's body, occurring before the onset of the wing downstroke. We estimate that legs generate, at various times, an upward force that can be much larger than all other forces applied to the insect's body. The relative timing of leg and wing forces explains the large variability of trajectories observed during the maneuvers.

  1. On the colour of wing scales in butterflies: iridescence and preferred orientation of single gyroid photonic crystals.

    PubMed

    Corkery, Robert W; Tyrode, Eric C

    2017-08-06

    Lycaenid butterflies from the genera Callophrys , Cyanophrys and Thecla have evolved remarkable biophotonic gyroid nanostructures within their wing scales that have only recently been replicated by nanoscale additive manufacturing. These nanostructures selectively reflect parts of the visible spectrum to give their characteristic non-iridescent, matte-green appearance, despite a distinct blue-green-yellow iridescence predicted for individual crystals from theory. It has been hypothesized that the organism must achieve its uniform appearance by growing crystals with some restrictions on the possible distribution of orientations, yet preferential orientation observed in Callophrys rubi confirms that this distribution need not be uniform. By analysing scanning electron microscope and optical images of 912 crystals in three wing scales, we find no preference for their rotational alignment in the plane of the scales. However, crystal orientation normal to the scale was highly correlated to their colour at low (conical) angles of view and illumination. This correlation enabled the use of optical images, each containing up to 10 4 -10 5 crystals, for concluding the preferential alignment seen along the [Formula: see text] at the level of single scales, appears ubiquitous. By contrast, [Formula: see text] orientations were found to occur at no greater rate than that expected by chance. Above a critical cone angle, all crystals reflected bright green light indicating the dominant light scattering is due to the predicted band gap along the [Formula: see text] direction, independent of the domain orientation. Together with the natural variation in scale and wing shapes, we can readily understand the detailed mechanism of uniform colour production and iridescence suppression in these butterflies. It appears that the combination of preferential alignment normal to the wing scale, and uniform distribution within the plane is a near optimal solution for homogenizing the angular distribution of the [Formula: see text] band gap relative to the wings. Finally, the distributions of orientations, shapes, sizes and degree of order of crystals within single scales provide useful insights for understanding the mechanisms at play in the formation of these biophotonic nanostructures.

  2. Mutations in CTNNA1 cause butterfly-shaped pigment dystrophy and perturbed retinal pigment epithelium integrity

    PubMed Central

    Saksens, Nicole T.M.; Krebs, Mark P.; Schoenmaker-Koller, Frederieke E.; Hicks, Wanda; Yu, Minzhong; Shi, Lanying; Rowe, Lucy; Collin, Gayle B.; Charette, Jeremy R.; Letteboer, Stef J.; Neveling, Kornelia; van Moorsel, Tamara W.; Abu-Ltaif, Sleiman; De Baere, Elfride; Walraedt, Sophie; Banfi, Sandro; Simonelli, Francesca; Cremers, Frans P.M.; Boon, Camiel J.F.; Roepman, Ronald; Leroy, Bart P.; Peachey, Neal S.; Hoyng, Carel B.; Nishina, Patsy M.; den Hollander, Anneke I.

    2015-01-01

    Butterfly-shaped pigment dystrophy is an eye disease characterized by lesions in the macula that can resemble the wings of a butterfly. Here, we report the identification of heterozygous missense mutations in the α-catenin 1 (CTNNA1) gene in three families with butterfly-shaped pigment dystrophy. In addition, we identified a Ctnna1 missense mutation in a chemically induced mouse mutant, tvrm5. Parallel clinical phenotypes were observed in the retinal pigment epithelium (RPE) of individuals with butterfly-shaped pigment dystrophy and in tvrm5 mice, including pigmentary abnormalities, focal thickening and elevated lesions, and decreased light-activated responses. Morphological studies in tvrm5 mice revealed increased cell shedding and large multinucleated RPE cells, suggesting defects in intercellular adhesion and cytokinesis. This study identifies CTNNA1 gene variants as a cause of macular dystrophy, suggests that CTNNA1 is involved in maintaining RPE integrity, and suggests that other components that participate in intercellular adhesion may be implicated in macular disease. PMID:26691986

  3. Wing scale microstructures and nanostructures in butterflies--natural photonic crystals.

    PubMed

    Vértesy, Z; Bálint, Zs; Kertész, K; Vigneron, J P; Lousse, V; Biró, L P

    2006-10-01

    The aim of our study was to investigate the correlation between structural colour and scale morphology in butterflies. Detailed correlations between blue colour and structure were investigated in three lycaenid subfamilies, which represent a monophylum in the butterfly family Lycaenidae (Lepidoptera): the Coppers (Lycaeninae), the Hairstreaks (Theclinae) and the Blues (Polyommatinae). Complex investigations such as spectral measurements and characterization by means of light microscopy, scanning electron microscopy and transmission electron microscopy enabled us to demonstrate that: (i) a wide array of nanostructures generate blue colours; (ii) monophyletic groups use qualitatively similar structures; and (iii) the hue of the blue colour is characteristic for the microstructure and nanostructure of the body of the scales.

  4. Mechanisms of structural colour in the Morpho butterfly: cooperation of regularity and irregularity in an iridescent scale.

    PubMed Central

    Kinoshita, Shuichi; Yoshioka, Shinya; Kawagoe, Kenji

    2002-01-01

    Structural colour in the Morpho butterfly originates from submicron structure within a scale and, for over a century, its colour and reflectivity have been explained as interference of light due to the multilayer of cuticle and air. However, this model fails to explain the extraordinarily uniform colour of the wing with respect to the observation direction. We have performed microscopic, optical and theoretical investigations, and have found that the separate lamellar structure with irregular heights is extremely important. Using a simple model, we have shown that the combined action of interference and diffraction is essential for the structural colour of the Morpho butterfly. PMID:12137569

  5. Two lactones in the androconial scent of the lycaenid butterfly Celastrina argiolus ladonides

    NASA Astrophysics Data System (ADS)

    Ômura, Hisashi; Yakumaru, Kazuhisa; Honda, Keiichi; Itoh, Takao

    2013-04-01

    Male adult butterflies of many species have characteristic odors originating from the disseminating organs known as androconia. Despite the fact that androconia exist in several species, there have been few investigations on adult scents from the lycaenid species. Celastrina argiolus ladonides (Lycaenidae) is a common species in Eurasia. We have reported that male adults of this species emit a faint odor, and the major components causing this odor have been newly found in the Insecta. By using field-caught individuals, we determined the chemical nature and location of this odor in the butterfly. Gas chromatography-mass spectrometry (GC-MS) analyses revealed that two lactone compounds, lavender lactone and δ-decalactone, are present in the extracts of males but absent in those of the females. On an average, approximately 50 ng of each compound was found per male. Chiral GC analyses performed using enantiomerically pure standards revealed that the natural lavender lactone was a mixture of two enantiomers with an R/ S ratio of 32:68, whereas the natural δ-decalactone contained only the R-enantiomer. When the analyses were conducted using different parts—forewings, hindwings, and body—of three males, the lactones were more abundantly found on the forewings and hindwings than on the body. Microscopic observation of the wings demonstrated that battledore scales known as androconia are scattered on the upper surface of both the wings of C. argiolus ladonides males. These results indicate that the specialized scales on the wings of males serve as scent-disseminating organs.

  6. Selection of Valid Reference Genes for Reverse Transcription Quantitative PCR Analysis in Heliconius numata (Lepidoptera: Nymphalidae)

    PubMed Central

    Chouteau, Mathieu; Whibley, Annabel; Joron, Mathieu; Llaurens, Violaine

    2016-01-01

    Identifying the genetic basis of adaptive variation is challenging in non-model organisms and quantitative real time PCR. is a useful tool for validating predictions regarding the expression of candidate genes. However, comparing expression levels in different conditions requires rigorous experimental design and statistical analyses. Here, we focused on the neotropical passion-vine butterflies Heliconius, non-model species studied in evolutionary biology for their adaptive variation in wing color patterns involved in mimicry and in the signaling of their toxicity to predators. We aimed at selecting stable reference genes to be used for normalization of gene expression data in RT-qPCR analyses from developing wing discs according to the minimal guidelines described in Minimum Information for publication of Quantitative Real-Time PCR Experiments (MIQE). To design internal RT-qPCR controls, we studied the stability of expression of nine candidate reference genes (actin, annexin, eF1α, FK506BP, PolyABP, PolyUBQ, RpL3, RPS3A, and tubulin) at two developmental stages (prepupal and pupal) using three widely used programs (GeNorm, NormFinder and BestKeeper). Results showed that, despite differences in statistical methods, genes RpL3, eF1α, polyABP, and annexin were stably expressed in wing discs in late larval and pupal stages of Heliconius numata. This combination of genes may be used as a reference for a reliable study of differential expression in wings for instance for genes involved in important phenotypic variation, such as wing color pattern variation. Through this example, we provide general useful technical recommendations as well as relevant statistical strategies for evolutionary biologists aiming to identify candidate-genes involved adaptive variation in non-model organisms. PMID:27271971

  7. A new Hermeuptychia (Lepidoptera, Nymphalidae, Satyrinae) is sympatric and synchronic with H. sosybius in southeast US coastal plains, while another new Hermeuptychia species – not hermes – inhabits south Texas and northeast Mexico

    PubMed Central

    Cong, Qian; Grishin, Nick V.

    2014-01-01

    Abstract Hermeuptychia intricata Grishin, sp. n. is described from the Brazos Bend State Park in Texas, United States, where it flies synchronously with Hermeuptychia sosybius (Fabricius, 1793). The two species differ strongly in both male and female genitalia and exhibit 3.5% difference in the COI barcode sequence of mitochondrial DNA. Setting such significant genitalic and genotypic differences aside, we were not able to find reliable wing pattern characters to tell a difference between the two species. This superficial similarity may explain why H. intricata, only distantly related to H. sosybius, has remained unnoticed until now, despite being widely distributed in the coastal plains from South Carolina to Texas, USA (and possibly to Costa Rica). Obscuring the presence of a cryptic species even further, wing patterns are variable in both butterflies and ventral eyespots vary from large to almost absent. To avoid confusion with the new species, neotype for Papilio sosybius Fabricius, 1793, a common butterfly that occurs across northeast US, is designated from Savannah, Georgia, USA. It secures the universally accepted traditional usage of this name. Furthermore, we find that DNA barcodes of Hermeuptychia specimens from the US, even those from extreme south Texas, are at least 4% different from those of H. hermes (Fabricius, 1775)—type locality Brazil: Rio de Janeiro—and suggest that the name H. hermes should not be used for USA populations, but rather reserved for the South American species. This conclusion is further supported by comparison of male genitalia. However, facies, genitalia and 2.1% different DNA barcodes set Hermeuptychia populations in the lower Rio Grande Valley of Texas apart from H. sosybius. These southern populations, also found in northeastern Mexico, are described here as Hermeuptychia hermybius Grishin, sp. n. (type locality Texas: Cameron County). While being phylogenetically closer to H. sosybius than to any other Hermeuptychia species, H. hermybius can usually be recognized by wing patterns, such as the size of eyespots and the shape of brown lines on hindwing. “Intricate Satyr” and “South Texas Satyr” are proposed as the English names for H. intricata and H. hermybius, respectively. PMID:24574857

  8. Morpho peleides butterfly wing imprints as structural colour stamp.

    PubMed

    Zobl, Sigrid; Salvenmoser, Willi; Schwerte, Thorsten; Gebeshuber, Ille C; Schreiner, Manfred

    2016-02-02

    This study presents the replication of a color-causing nanostructure based on the upper laminae of numerous cover scales of Morpho peleides butterfly wings and obtained solely by imprinting their upper-wing surfaces. Our results indicate that a simple casting technique using a novel integrated release agent can obtain a large positive replica using negative imprints via Polyvinylsiloxane. The developed method is low-tech and high-yield and is thus substantially easier and less expensive than previous methods. The microstructures were investigated with light microscopy, the nanostructures with both scanning and transmission electron microscopy, and the reflections with UV visible spectrometry. The influence of the release agent and the quality of the master stamp were determined by comparing measurements of the cover-scale sizes and their chromaticity values obtained by their images and with their positive imprints. The master stamp provided multiple positive replicas up to 3 cm(2) in just 1 h with structural coloration effects visible to the naked eye. Thus, the developed method proves the accuracy of the replicated nanostructure and its potential industrial application as a color-producing nanostamp.

  9. Magnetic intermittency of solar wind turbulence in the dissipation range

    NASA Astrophysics Data System (ADS)

    Pei, Zhongtian; He, Jiansen; Tu, Chuanyi; Marsch, Eckart; Wang, Linghua

    2016-04-01

    The feature, nature, and fate of intermittency in the dissipation range are an interesting topic in the solar wind turbulence. We calculate the distribution of flatness for the magnetic field fluctuations as a functionof angle and scale. The flatness distribution shows a "butterfly" pattern, with two wings located at angles parallel/anti-parallel to local mean magnetic field direction and main body located at angles perpendicular to local B0. This "butterfly" pattern illustrates that the flatness profile in (anti-) parallel direction approaches to the maximum value at larger scale and drops faster than that in perpendicular direction. The contours for probability distribution functions at different scales illustrate a "vase" pattern, more clear in parallel direction, which confirms the scale-variation of flatness and indicates the intermittency generation and dissipation. The angular distribution of structure function in the dissipation range shows an anisotropic pattern. The quasi-mono-fractal scaling of structure function in the dissipation range is also illustrated and investigated with the mathematical model for inhomogeneous cascading (extended p-model). Different from the inertial range, the extended p-model for the dissipation range results in approximate uniform fragmentation measure. However, more complete mathematicaland physical model involving both non-uniform cascading and dissipation is needed. The nature of intermittency may be strong structures or large amplitude fluctuations, which may be tested with magnetic helicity. In one case study, we find the heating effect in terms of entropy for large amplitude fluctuations seems to be more obvious than strong structures.

  10. Butterflies: Photonic Crystals on the Wing

    DTIC Science & Technology

    2007-03-22

    Coliadinae), Eronia leda, Colotis danae, and Pieris rapae rapae (all Pierinae), respectively. The left and right column of the photographs are taken...phylogenetic tree: Colias electo belongs to the Coliadinae, Eronia leda and Colotis danae belong to the Colotis group, and Pieris rapae rapae to the Pierini...the droplet is brought into contact with a piece of wing (here of a large white, Pieris brassicae). 3 Fig. 4. Electron microscopic photographs

  11. Contribution of both the upperside and the underside of the wing on the iridescence in the male butterfly Troïdes magellanus (Papilionidae)

    NASA Astrophysics Data System (ADS)

    Van Hooijdonk, Eloise; Berthier, Serge; Vigneron, Jean-Pol

    2012-10-01

    Until now, the existing literature suggests that the color effects on the butterflies wings come from the scales exposed to an outside observer. The particularity of the present work lies in the consideration of both sides of a wing. The male Troïdes magellanus concentrates on its hindwings distinct but complementary optical effects contributing to its exceptional visual attraction: (i) a uniform yellow coloration under daylight, (ii) a bright glint showing bluish or greenish hues under grazing illumination and observation, and (iii) the presence of fluorescent molecules, producing a yellow-green coloration when irradiated by ultraviolet light, embedded in a three-dimensional photonic structure. Our experimental examination reveals a decomposition of the visual aspect in terms of a pigmentary, a structural, and a fluorescent component, as well as a contribution of the scales of the upperside and the underside of the wing on this observation. Our results highlight the role of the scales photonic structure—from both sides of the organ—on this correlation. The investigation was realized by means of spectrophotometric measurements, bidirectional reflectance distribution function mapping, and electron microscopy. Analyzing the scattered light according to the observation direction proved to be crucial in this work.

  12. Hierarchical organization of butterfly gyroid nanostructures provide a time-frozen glimpse of intracellular membrane development

    NASA Astrophysics Data System (ADS)

    Wilts, Bodo; Winter, Benjamin; Klatt, Michael; Butz, Benjamin; Fischer, Michael; Kelly, Stephen; Spieker, Erdmann; Steiner, Ullrich; Schroeder-Turk, Gerd

    The formation of the biophotonic gyroid material in butterfly wing scales is an exceptional feat of evolutionary engineering of functional nanostructures. Previous work hypothesized that this nanostructure forms by chitin polymerization inside a convoluted membrane of corresponding shape in the endoplasmic reticulum. In vivo imaging however cannot yet elucidate this dynamic formation process, including whether membrane folding and chitin expression are simultaneous or subsequent processes. Here we show an unusual hierarchical ultrastructure in a Hairstreak butterfly that allows high-resolution 3D microscopy. Rather than the conventional polycrystalline space-filling arrangement, the gyroid occurs in isolated facetted crystallites with a pronounced size-gradient. This arrangement is interpreted as a sequence of time-frozen snapshots of the morphogenesis. This provides insight into the formation mechanisms of the nanoporous gyroid material, especially when compared among other butterflies with different arrangements. Financially supported through DFG, the NCCR Bio-inspired Mateirals and the SNF Ambizione programme.

  13. Complex Population Patterns of Eunica tatila Herrich-Schäffer (Lepidoptera: Nymphalidae), with Special Emphasis on Sexual Dimorphism.

    PubMed

    Cavanzón-Medrano, L; Pozo, C; Hénaut, Y; Legal, L; Salas-Suárez, N; Machkour-M'Rabet, S

    2016-04-01

    The species Eunica tatila (Herrich-Schäffer) is present in the Neotropical region and comprises three subspecies. In Mexico, only one subspecies is reported: E. t. tatila (Herrich-Schäffer). The Yucatan Peninsula, in southeastern Mexico, is located in a transitional geographical position, between southern Florida, the West Indies and Central America. It is part of a transitional region, important for the dispersion of insects from southern Florida via Cuba and the Yucatan Peninsula. Considering the possibility of the overlapping and delimitation of described subspecies, we sampled different populations in the Yucatan Peninsula to possibly assign a subspecies name and evaluate the magnitude of sexual dimorphism. We collected 591 individuals (♀284, ♂307) in conserved areas. The study of male genitalia led to the identification of Eunica tatila tatilista (Kaye) as a subspecies; however, hypandrium structure and wing pattern analysis suggest a mix of E. t. tatila and E. t. tatilista characteristics. The analysis of sexual dimorphism provided evidence of more complex wing morphs for females, with 12 patterns instead of four as previously described. Our results demonstrate the complexity of characterizing E. tatila and suggest that the Yucatan Peninsula is a transitional zone for subspecies of some butterflies.

  14. Documentation of roller-bearing effect on butterfly inspired grooves

    NASA Astrophysics Data System (ADS)

    Gautam, Sashank; Lang, Amy

    2017-11-01

    Butterfly wings are covered with scales in a roof shingle pattern which align together to form grooves. The increase or decrease of laminar friction drag depends on the flow orientation to the scales. Flow in the longitudinal direction to the grooves encounters increased surface area which increases the friction drag. However, in the transverse direction, for low Re laminar flow, a single vortex is formed inside each groove and is predicted to remain stable due to the very low Re of the flow in each cavity. These embedded vortices act as roller bearings to the flow above, such that the fluid from the outer boundary layer does not mix with fluid inside the cavities. This leads to a reduction of skin friction drag when compared to a smooth surface. When the cavity flow Re is increased beyond a critical point, the vortex becomes unstable and the low-momentum fluid in the grooves mixes with the outer boundary layer flow, increasing the drag. The objective of this experiment is to determine the critical Re where the embedded vortex transitions from a stable to an unstable state using DPIV. Subsequently, for steady vortex conditions, a comparison of skin friction drag between the grooved and flat plate can show that the butterfly scaled surface can result in sub-laminar friction drag. The National Science Foundation (Grant No. 1335848).

  15. The Hofstadter Butterfly and some physical consequences

    NASA Astrophysics Data System (ADS)

    Claro, Francisco

    Opening its beautiful wings for the first time four decades ago, the Hofstadter Butterfly emerged as a self-similar pattern of bands and gaps displaying the allowed energies for two dimensional crystalline electrons in a perpendicular magnetic field. Within the Harper model, as the external field parameter is varied well defined gaps traverse the spectrum, some closing at a Dirac point where two approaching bands touch. Such band edges degeneracy is lifted in more realistic models. Gaps have a unique label that determines the Hall conductivity of a noninteracting electron system, as observed in recent experiments. When the 2D electron assembly is allowed to interact in the absence of an underlying periodic potential, the mean field approximation predicts a liquid at integer filling fractions and electron density fluctuations otherwise, which if periodic may be represented again by a Harper equation. The intriguing odd denominator rule observed in experiment in the fractional quantum Hall regime is then a natural prediction of the model. Although I have an affiliation (lifetime Granted) I am actually retired (do not have a paid contract).

  16. Photonic effects in natural nanostructures

    NASA Astrophysics Data System (ADS)

    Rey GonzáLez, Rafael Ramón; Barrera Patiã+/-O, Claudia Patricia

    Nature exhibits a great variety of structures and nanostructures. In particular the interaction light-matter has a strong dependence with the shape of the nanostructures. In some cases, in the so called structural color, ordered arrays of nanostructures play a very critical role. One of the most interesting color effects is the iridescence, the angular dependence of the observed color in some species of butterflies, insects, plants, beetles, fishes, birds and even in minerals. In the last years, iridescence has been related with photonic properties. In the present work, we present a theoretical study of the photonic properties for different patterns that exist in natural nanostructures present in wings of butterflies that exhibit iridescence. The nanostructures observed in these cases present spatial variations of the dielectric constant that are possible to model them as 1D and 2D photonic crystal. Partial photonic gaps are found as function of lattice constant, dielectric contrast and geometrical configuration. Also, disordered effects are considered. Authors would like to thank the División de Investigación Sede Bogotá for their financial support at Universidad Nacional de Colombia.

  17. [Study of Japanese anatomical terms, such as 'sphenoid bone'].

    PubMed

    Sawai, Tadashi

    2008-12-01

    Japanese anatomical terms (butterfly-shaped bone) have an interesting history. Galen named a bone (wedge-like). This Greek term was introduced into Latin anatomical texts by transcribing into 'os sphnoides' or translating it as 'os cuneiforme'. Both terms mean equally wedge-like bone. From 16th century on, these two terms prevailed in European anatomical textbooks, but in 18th century some anatomists merged this bone with some kinds of winged creatures and named their wings "Ala major' and 'Ala minor'. In mid-19th century English-Chinese anatomical book, this bone was named (butterfly bone) by a medical missionary Benjamin Hobson. This term was introduced into Japanese textbooks. In Meiji Era both terms were used in Japanese textbooks, and (wedged-like bone). Some anatomists insisted on using because this echoed original Latin term's sense. Eventually, Japanese Associations of Anatomists adopted in 1943.

  18. Coloration mechanisms and phylogeny of Morpho butterflies.

    PubMed

    Giraldo, M A; Yoshioka, S; Liu, C; Stavenga, D G

    2016-12-15

    Morpho butterflies are universally admired for their iridescent blue coloration, which is due to nanostructured wing scales. We performed a comparative study on the coloration of 16 Morpho species, investigating the morphological, spectral and spatial scattering properties of the differently organized wing scales. In numerous previous studies, the bright blue Morpho coloration has been fully attributed to the multi-layered ridges of the cover scales' upper laminae, but we found that the lower laminae of the cover and ground scales play an important additional role, by acting as optical thin film reflectors. We conclude that Morpho coloration is a subtle combination of overlapping pigmented and/or unpigmented scales, multilayer systems, optical thin films and sometimes undulated scale surfaces. Based on the scales' architecture and their organization, five main groups can be distinguished within the genus Morpho, largely agreeing with the accepted phylogeny. © 2016. Published by The Company of Biologists Ltd.

  19. High Selectivity Toward C2H4 Production over Cu Particles Supported by Butterfly-Wing-Derived Carbon Frameworks.

    PubMed

    Huo, Yajiao; Peng, Xianyun; Liu, Xijun; Li, Huaiyu; Luo, Jun

    2018-04-18

    Converting carbon dioxide to useful C2 chemicals in a selective and efficient manner remains a major challenge in renewable and sustainable energy research. Herein, we adopt butterfly wings to assist the preparation of an electrocatalyst containing monodispersed Cu particles supported by nitrogen-doped carbon frameworks for an efficient reduction of CO 2 . Benefiting from structure advantages and the synergistic effect between nitrogen dopants and stepped surface-rich Cu particles, the resulting catalyst exhibited a high faradic efficiency of 63.7 ± 1.4% for ethylene production (corresponding to an ethylene/methane products' ratio of 57.9 ± 5.4) and an excellent durability (∼100% retention after 24 h). This work presents some guidelines for the rational design and accurate modulation of metal heterocatalysts for high selectivity toward ethylene from CO 2 electroreduction.

  20. An illustrated checklist of the genus Elymnias Hübner, 1818 (Nymphalidae, Satyrinae)

    PubMed Central

    Wei, Chia-Hsuan; Lohman, David J.; Peggie, Djunijanti; Yen, Shen-Horn

    2017-01-01

    Abstract We review the genus Elymnias Hübner, 1818, a morphologically diverse satyrine butterfly clade involved in multifarious Batesian mimicry relationships throughout Asia and Africa. A variety of different model species are mimicked, and many Elymnias species are sexually dimorphic mimics, with males and females resembling different model species. We revise species and subspecies delimitations in light of an integrative taxonomic investigation using external morphology, male and female genital morphology, and a multi-locus molecular phylogeny. There is little interspecific genitalic variation among species in this group, and previous taxonomists therefore relied almost entirely on wing patterns. Our molecular phylogenetic analysis reveals several examples of polymorphism or wing pattern divergence within a single species currently classified as two or more different species. We also found examples of wing pattern convergence among disparate lineages that mimic the same widespread model species. Frequently, two or more phenotypically similar species were classified as a single species. This comprehensive checklist reviews all names associated with Elymnias to align its taxonomy with the evolutionary history of the group. All available information on nomenclature, type localities, repositories of type specimens, and geographical distributions is summarized, and images of adult specimens and genitalia are provided along with distribution maps of all species and selected subspecies. We identify 2 species incertae sedis, establish 15 monophyletic species groups (including 1 species unplaced in any species group), and make 49 taxonomic changes, including 35 new synonyms, 7 new combinations (2 of which have new status), 1 resurrected combination, 1 resurrected subspecies, and 7 status changes. PMID:28769686

  1. Coexistence of both gyroid chiralities in individual butterfly wing scales of Callophrys rubi

    PubMed Central

    Winter, Benjamin; Butz, Benjamin; Dieker, Christel; Schröder-Turk, Gerd E.; Mecke, Klaus; Spiecker, Erdmann

    2015-01-01

    The wing scales of the Green Hairstreak butterfly Callophrys rubi consist of crystalline domains with sizes of a few micrometers, which exhibit a congenitally handed porous chitin microstructure identified as the chiral triply periodic single-gyroid structure. Here, the chirality and crystallographic texture of these domains are investigated by means of electron tomography. The tomograms unambiguously reveal the coexistence of the two enantiomeric forms of opposite handedness: the left- and right-handed gyroids. These two enantiomers appear with nonequal probabilities, implying that molecularly chiral constituents of the biological formation process presumably invoke a chiral symmetry break, resulting in a preferred enantiomeric form of the gyroid structure. Assuming validity of the formation model proposed by Ghiradella H (1989) J Morphol 202(1):69–88 and Saranathan V, et al. (2010) Proc Natl Acad Sci USA 107(26):11676–11681, where the two enantiomeric labyrinthine domains of the gyroid are connected to the extracellular and intra-SER spaces, our findings imply that the structural chirality of the single gyroid is, however, not caused by the molecular chirality of chitin. Furthermore, the wing scales are found to be highly textured, with a substantial fraction of domains exhibiting the <001> directions of the gyroid crystal aligned parallel to the scale surface normal. Both findings are needed to completely understand the photonic purpose of the single gyroid in gyroid-forming butterflies. More importantly, they show the level of control that morphogenesis exerts over secondary features of biological nanostructures, such as chirality or crystallographic texture, providing inspiration for biomimetic replication strategies for synthetic self-assembly mechanisms. PMID:26438839

  2. Coexistence of both gyroid chiralities in individual butterfly wing scales of Callophrys rubi.

    PubMed

    Winter, Benjamin; Butz, Benjamin; Dieker, Christel; Schröder-Turk, Gerd E; Mecke, Klaus; Spiecker, Erdmann

    2015-10-20

    The wing scales of the Green Hairstreak butterfly Callophrys rubi consist of crystalline domains with sizes of a few micrometers, which exhibit a congenitally handed porous chitin microstructure identified as the chiral triply periodic single-gyroid structure. Here, the chirality and crystallographic texture of these domains are investigated by means of electron tomography. The tomograms unambiguously reveal the coexistence of the two enantiomeric forms of opposite handedness: the left- and right-handed gyroids. These two enantiomers appear with nonequal probabilities, implying that molecularly chiral constituents of the biological formation process presumably invoke a chiral symmetry break, resulting in a preferred enantiomeric form of the gyroid structure. Assuming validity of the formation model proposed by Ghiradella H (1989) J Morphol 202(1):69-88 and Saranathan V, et al. (2010) Proc Natl Acad Sci USA 107(26):11676-11681, where the two enantiomeric labyrinthine domains of the gyroid are connected to the extracellular and intra-SER spaces, our findings imply that the structural chirality of the single gyroid is, however, not caused by the molecular chirality of chitin. Furthermore, the wing scales are found to be highly textured, with a substantial fraction of domains exhibiting the <001> directions of the gyroid crystal aligned parallel to the scale surface normal. Both findings are needed to completely understand the photonic purpose of the single gyroid in gyroid-forming butterflies. More importantly, they show the level of control that morphogenesis exerts over secondary features of biological nanostructures, such as chirality or crystallographic texture, providing inspiration for biomimetic replication strategies for synthetic self-assembly mechanisms.

  3. AFM imaging of natural optical structures

    NASA Astrophysics Data System (ADS)

    Dallaeva, Dinara; Tománek, Pavel; Prokopyeva, Elena; Kaspar, Pavel; Grmela, Lubomír.; Škarvada, Pavel

    2015-01-01

    The colors of some living organisms assosiated with the surface structure. Irridesence butterfly wings is an example of such coloration. Optical effects such as interference, diffraction, polarization are responsible for physical colors appearance. Alongside with amazing beauty this structure represent interest for design of optical devices. Here we report the results of morphology investigation by atomic force microscopy. The difference in surface structure of black and blue wings areas is clearly observed. It explains the angle dependence of the wing blue color, since these micrometer and sub-micrometer quasiperiodical structures could control the light propagation, absorption and reflection.

  4. Swimming of a Sea Butterfly with an Elongated Shell

    NASA Astrophysics Data System (ADS)

    Karakas, Ferhat; Maas, Amy E.; Murphy, David W.

    2017-11-01

    Sea butterflies (pteropods) are small, zooplanktonic marine snails which swim by flapping highly flexible parapodia. Previous studies show that the swimming hydrodynamics of Limacina helicina, a polar pteropod with a spiraled shell, is similar to tiny insect flight aerodynamics and that forward-backward pitching is key for lift generation. However, swimming by diverse pteropod species with different shell shapes has not been examined. We present measurements of the swimming of Cuvierina columnella, a warm water species with an elongated non-spiraled shell collected off the coast of Bermuda. With a body length of 9 mm, wing beat frequency of 4-6 Hz and swimming speed of 35 mm/s, these organisms swim at a Reynolds number of approximately 300, larger than that of L. helicina. High speed 3D kinematics acquired via two orthogonal cameras reveals that the elongated shell correlates with reduced body pitching and that the wings bend approximately 180 degrees in each direction, overlapping at the end of each half-stroke. Time resolved 2D flow measurements collected with a micro-PIV system reveal leading edge vortices present in both power and recovery strokes. Interactions between the overlapping wings and the shell also likely play a role in lift generation.

  5. On the Total Edge Irregularity Strength of Generalized Butterfly Graph

    NASA Astrophysics Data System (ADS)

    Dwi Wahyuna, Hafidhyah; Indriati, Diari

    2018-04-01

    Let G(V, E) be a connected, simple, and undirected graph with vertex set V and edge set E. A total k-labeling is a map that carries vertices and edges of a graph G into a set of positive integer labels {1, 2, …, k}. An edge irregular total k-labeling λ: V(G) ∪ E(G) → {1, 2, …, k} of a graph G is a total k-labeling such that the weights calculated for all edges are distinct. The weight of an edge uv in G, denoted by wt(uv), is defined as the sum of the label of u, the label of v, and the label of uv. The total edge irregularity strength of G, denoted by tes(G), is the minimum value of the largest label k over all such edge irregular total k-labelings. A generalized butterfly graph, BFn , obtained by inserting vertices to every wing with assumption that sum of inserting vertices to every wing are same then it has 2n + 1 vertices and 4n ‑ 2 edges. In this paper, we investigate the total edge irregularity strength of generalized butterfly graph, BFn , for n > 2. The result is tes(B{F}n)=\\lceil \\frac{4n}{3}\\rceil .

  6. The enigmatic fast leaflet rotation in Desmodium motorium: butterfly mimicry for defense?

    PubMed

    Lev-Yadun, Simcha

    2013-06-01

    I propose that the enigmatic leaflet movements in elliptical circles every few minutes of the Indian telegraph (semaphore) plant Desmodium motorium ( = D. gyrans = Hedysarum gyrans = Codariocalyx motorius), which has intrigued scientists for centuries, is a new type of butterfly or general winged arthropod mimicry by this plant. Such leaflet movement may deceive a passing butterfly searching for an un-occupied site suitable to deposit its eggs, that the plant is already occupied. It may also attract insectivorous birds, reptiles or arthropods to the plant because it looks as if it is harboring a potential prey and while they patrol there, they can find insects or other invertebrates that indeed attack the plant. The possibility that diurnal mammalian herbivores may also be deterred by these movements should not be dismissed.

  7. Exaggeration and suppression of iridescence: the evolution of two-dimensional butterfly structural colours

    PubMed Central

    Wickham, Shelley; Large, Maryanne C.J; Poladian, Leon; Jermiin, Lars S

    2005-01-01

    Many butterfly species possess ‘structural’ colour, where colour is due to optical microstructures found in the wing scales. A number of such structures have been identified in butterfly scales, including three variations on a simple multi-layer structure. In this study, we optically characterize examples of all three types of multi-layer structure, as found in 10 species. The optical mechanism of the suppression and exaggeration of the angle-dependent optical properties (iridescence) of these structures is described. In addition, we consider the phylogeny of the butterflies, and are thus able to relate the optical properties of the structures to their evolutionary development. By applying two different types of analysis, the mechanism of adaptation is addressed. A simple parsimony analysis, in which all evolutionary changes are given an equal weighting, suggests convergent evolution of one structure. A Dollo parsimony analysis, in which the evolutionary ‘cost’ of losing a structure is less than that of gaining it, implies that ‘latent’ structures can be reused. PMID:16849221

  8. An Exploratory Study of the Butterfly Effect Using Agent-Based Modeling

    NASA Technical Reports Server (NTRS)

    Khasawneh, Mahmoud T.; Zhang, Jun; Shearer, Nevan E. N.; Rodriquez-Velasquez, Elkin; Bowling, Shannon R.

    2010-01-01

    This paper provides insights about the behavior of chaotic complex systems, and the sensitive dependence of the system on the initial starting conditions. How much does a small change in the initial conditions of a complex system affect it in the long term? Do complex systems exhibit what is called the "Butterfly Effect"? This paper uses an agent-based modeling approach to address these questions. An existing model from NetLogo library was extended in order to compare chaotic complex systems with near-identical initial conditions. Results show that small changes in initial starting conditions can have a huge impact on the behavior of chaotic complex systems. The term the "butterfly effect" is attributed to the work of Edward Lorenz [1]. It is used to describe the sensitive dependence of the behavior of chaotic complex systems on the initial conditions of these systems. The metaphor refers to the notion that a butterfly flapping its wings somewhere may cause extreme changes in the ecological system's behavior in the future, such as a hurricane.

  9. Divergence with gene flow across a speciation continuum of Heliconius butterflies.

    PubMed

    Supple, Megan A; Papa, Riccardo; Hines, Heather M; McMillan, W Owen; Counterman, Brian A

    2015-09-24

    A key to understanding the origins of species is determining the evolutionary processes that drive the patterns of genomic divergence during speciation. New genomic technologies enable the study of high-resolution genomic patterns of divergence across natural speciation continua, where taxa pairs with different levels of reproductive isolation can be used as proxies for different stages of speciation. Empirical studies of these speciation continua can provide valuable insights into how genomes diverge during speciation. We examine variation across a handful of genomic regions in parapatric and allopatric populations of Heliconius butterflies with varying levels of reproductive isolation. Genome sequences were mapped to 2.2-Mb of the H. erato genome, including 1-Mb across the red color pattern locus and multiple regions unlinked to color pattern variation. Phylogenetic analyses reveal a speciation continuum of pairs of hybridizing races and incipient species in the Heliconius erato clade. Comparisons of hybridizing pairs of divergently colored races and incipient species reveal that genomic divergence increases with ecological and reproductive isolation, not only across the locus responsible for adaptive variation in red wing coloration, but also at genomic regions unlinked to color pattern. We observe high levels of divergence between the incipient species H. erato and H. himera, suggesting that divergence may accumulate early in the speciation process. Comparisons of genomic divergence between the incipient species and allopatric races suggest that limited gene flow cannot account for the observed high levels of divergence between the incipient species. Our results provide a reconstruction of the speciation continuum across the H. erato clade and provide insights into the processes that drive genomic divergence during speciation, establishing the H. erato clade as a powerful framework for the study of speciation.

  10. Fine structures of wing scales in Sasakia charonda butterflies as photonic crystals.

    PubMed

    Matejková-Plskova, J; Shiojiri, S; Shiojiri, M

    2009-11-01

    We investigate the microstructure of scales in the wings of male Sasakia charonda charonda butterflies by scanning electron microscopy with the aid of optical microscopy. Six types of scales are identified: B1, W1 and R1 in brown background yellow spots and red spots, respectively; B2 in iridescent purple-blue and W2 in white pearl, both of which characterize the male and B3 in the wing edges. The B1, W1 and R1 scales are almost the same in structure and the B2 and W2 scales are almost the same. The difference among the B, W and R scales is in species and content of pigment. The B1, W1 and R1 scales have only two layers of cuticle lapped on the ridges. In contrast with them, the B2 and W2 scales have seven multilayers of cuticle piled on the ridge. The multiple interference of light that occurs among these cuticle layers, spaced with air layers, generates the significant iridescence of the B2 and W2 scales. Thus, the characteristic purple-blue of the male wings is ascribed to the combination of the structural and chemical colouration in the B2 scales with melanin. The photonic crystals of these scales may be applicable to fine light manipulators such as reflection elements in laser diodes. B3 has many holes between the ridges and no multilayers of cuticle on the ridges. These structures may play any role in aerodynamically easy flight and/or in drainage of wet wings.

  11. Butterfly scale form birefringence related to photonics.

    PubMed

    Vidal, Benedicto de Campos

    2011-12-01

    Wings of the butterflies Morpho aega and Eryphanis reevesi were investigated in the present study by fluorescence, polarization and infra-red (IR) spectroscopic microscopy with the aim of identifying the oriented organization of their components and morphological details of their substructures. These wings were found to exhibit a strong iridescent glow depending on the angle of the incident light; their isolated scales exhibited blue fluorescence. Parallel columns or ridges extend from the pad and sockets to the dented apical scale's region, and they are perpendicular to the ribs that connect the columnar ridges. The scales reveal linear dichroism (LD) visually, when attached on the wing matrix or isolated on slides. The LD was inferred to be textural and positive and was also demonstrated with IR microscopy. The scale columns and ribs are birefringent structures. Images obtained before and after birefringence compensation allowed a detailed study of the scale morphology. Form and intrinsic birefringence findings here estimated and discussed in the context of nonlinear optical properties, bring to the level of morphology the state of molecular order and periodicity of the wing structure. FT-IR absorption peaks were found at wavenumbers which correspond to symmetric and asymmetric (-N-H) stretching, symmetric (-C-H) stretching, amide I (-CO) stretching, amide II(-N-H), and β-linking. Based on LD results obtained with polarized IR the molecular vibrations of the wing scales of M. aega and E. reevesi are assumed to be oriented with respect to the long axis of these structures. Copyright © 2011 Elsevier Ltd. All rights reserved.

  12. Maintaining mimicry diversity: optimal warning colour patterns differ among microhabitats in Amazonian clearwing butterflies.

    PubMed

    Willmott, Keith R; Robinson Willmott, Julia C; Elias, Marianne; Jiggins, Chris D

    2017-05-31

    Mimicry is one of the best-studied examples of adaptation, and recent studies have provided new insights into the role of mimicry in speciation and diversification. Classical Müllerian mimicry theory predicts convergence in warning signal among protected species, yet tropical butterflies are exuberantly diverse in warning colour patterns, even within communities. We tested the hypothesis that microhabitat partitioning in aposematic butterflies and insectivorous birds can lead to selection for different colour patterns in different microhabitats and thus help maintain mimicry diversity. We measured distribution across flight height and topography for 64 species of clearwing butterflies (Ithomiini) and their co-mimics, and 127 species of insectivorous birds, in an Amazon rainforest community. For the majority of bird species, estimated encounter rates were non-random for the two most abundant mimicry rings. Furthermore, most butterfly species in these two mimicry rings displayed the warning colour pattern predicted to be optimal for anti-predator defence in their preferred microhabitats. These conclusions were supported by a field trial using butterfly specimens, which showed significantly different predation rates on colour patterns in two microhabitats. We therefore provide the first direct evidence to support the hypothesis that different mimicry patterns can represent stable, community-level adaptations to differing biotic environments. © 2017 The Author(s).

  13. Maintaining mimicry diversity: optimal warning colour patterns differ among microhabitats in Amazonian clearwing butterflies

    PubMed Central

    Robinson Willmott, Julia C.

    2017-01-01

    Mimicry is one of the best-studied examples of adaptation, and recent studies have provided new insights into the role of mimicry in speciation and diversification. Classical Müllerian mimicry theory predicts convergence in warning signal among protected species, yet tropical butterflies are exuberantly diverse in warning colour patterns, even within communities. We tested the hypothesis that microhabitat partitioning in aposematic butterflies and insectivorous birds can lead to selection for different colour patterns in different microhabitats and thus help maintain mimicry diversity. We measured distribution across flight height and topography for 64 species of clearwing butterflies (Ithomiini) and their co-mimics, and 127 species of insectivorous birds, in an Amazon rainforest community. For the majority of bird species, estimated encounter rates were non-random for the two most abundant mimicry rings. Furthermore, most butterfly species in these two mimicry rings displayed the warning colour pattern predicted to be optimal for anti-predator defence in their preferred microhabitats. These conclusions were supported by a field trial using butterfly specimens, which showed significantly different predation rates on colour patterns in two microhabitats. We therefore provide the first direct evidence to support the hypothesis that different mimicry patterns can represent stable, community-level adaptations to differing biotic environments. PMID:28539522

  14. eButterfly: Leveraging Massive Online Citizen Science for Butterfly Conservation

    PubMed Central

    Prudic, Kathleen L.; McFarland, Kent P.; Oliver, Jeffrey C.; Hutchinson, Rebecca A.; Long, Elizabeth C.; Kerr, Jeremy T.; Larrivée, Maxim

    2017-01-01

    Data collection, storage, analysis, visualization, and dissemination are changing rapidly due to advances in new technologies driven by computer science and universal access to the internet. These technologies and web connections place human observers front and center in citizen science-driven research and are critical in generating new discoveries and innovation in such fields as astronomy, biodiversity, and meteorology. Research projects utilizing a citizen science approach address scientific problems at regional, continental, and even global scales otherwise impossible for a single lab or even a small collection of academic researchers. Here we describe eButterfly an integrative checklist-based butterfly monitoring and database web-platform that leverages the skills and knowledge of recreational butterfly enthusiasts to create a globally accessible unified database of butterfly observations across North America. Citizen scientists, conservationists, policy makers, and scientists are using eButterfly data to better understand the biological patterns of butterfly species diversity and how environmental conditions shape these patterns in space and time. eButterfly in collaboration with thousands of butterfly enthusiasts has created a near real-time butterfly data resource producing tens of thousands of observations per year open to all to share and explore. PMID:28524117

  15. eButterfly: Leveraging Massive Online Citizen Science for Butterfly Consevation.

    PubMed

    Prudic, Kathleen L; McFarland, Kent P; Oliver, Jeffrey C; Hutchinson, Rebecca A; Long, Elizabeth C; Kerr, Jeremy T; Larrivée, Maxim

    2017-05-18

    Data collection, storage, analysis, visualization, and dissemination are changing rapidly due to advances in new technologies driven by computer science and universal access to the internet. These technologies and web connections place human observers front and center in citizen science-driven research and are critical in generating new discoveries and innovation in such fields as astronomy, biodiversity, and meteorology. Research projects utilizing a citizen science approach address scientific problems at regional, continental, and even global scales otherwise impossible for a single lab or even a small collection of academic researchers. Here we describe eButterfly an integrative checklist-based butterfly monitoring and database web-platform that leverages the skills and knowledge of recreational butterfly enthusiasts to create a globally accessible unified database of butterfly observations across North America. Citizen scientists, conservationists, policy makers, and scientists are using eButterfly data to better understand the biological patterns of butterfly species diversity and how environmental conditions shape these patterns in space and time. eButterfly in collaboration with thousands of butterfly enthusiasts has created a near real-time butterfly data resource producing tens of thousands of observations per year open to all to share and explore.

  16. Revised species definitions and nomenclature of the rose colored Cithaerias butterflies (Lepidoptera, Nymphalidae, Satyrinae).

    PubMed

    Penz, Carla M; Alexander, Laura G; Devries, Philip J

    2014-10-20

    This study provides updated species definitions for five rose-colored Cithaerias butterflies, starting with a historical overview of their taxonomy. Given their mostly transparent wings, genitalia morphology yielded the most reliable characters for species definition and identification. Genitalic divergence is more pronounced when multiple species occur in sympatry than between parapatric taxa. Cithaerias aurorina is granted full species status, C. cliftoni is reinstated as a full species, and one new combination is proposed, i.e. C. aurora tambopata. Two new synonyms are proposed, Callitaera phantoma and Callitaera aura = Cithaerias aurora. 

  17. Helium ion microscopy of Lepidoptera scales.

    PubMed

    Boden, Stuart A; Asadollahbaik, Asa; Rutt, Harvey N; Bagnall, Darren M

    2012-01-01

    In this report, helium ion microscopy (HIM) is used to study the micro and nanostructures responsible for structural color in the wings of two species of Lepidotera from the Papilionidae family: Papilio ulysses (Blue Mountain Butterfly) and Parides sesostris (Emerald-patched Cattleheart). Electronic charging of uncoated scales from the wings of these butterflies, due to the incident ion beam, is successfully neutralized, leading to images displaying a large depth-of-field and a high level of surface detail, which would normally be obscured by traditional coating methods used for scanning electron microscopy (SEM). The images are compared with those from variable pressure SEM, demonstrating the superiority of HIM at high magnifications. In addition, the large depth-of-field capabilities of HIM are exploited through the creation of stereo pairs that allows the exploration of the third dimension. Furthermore, the extraction of quantitative height information which matches well with cross-sectional transmission electron microscopy measurements from the literature is demonstrated. © Wiley Periodicals, Inc.

  18. Biased learning affects mate choice in a butterfly.

    PubMed

    Westerman, Erica L; Hodgins-Davis, Andrea; Dinwiddie, April; Monteiro, Antónia

    2012-07-03

    Early acquisition of mate preferences or mate-preference learning is associated with signal diversity and speciation in a wide variety of animal species. However, the diversity of mechanisms of mate-preference learning across taxa remains poorly understood. Using the butterfly Bicyclus anynana we uncover a mechanism that can lead to directional sexual selection via mate-preference learning: a bias in learning enhanced ornamentation, which is independent of preexisting mating biases. Naïve females mated preferentially with wild-type males over males with enhanced wing ornamentation, but females briefly exposed to enhanced males mated significantly more often with enhanced males. In contrast, females exposed to males with reduced wing ornamentation did not learn to prefer drab males. Thus, we observe both a learned change of a preexisting mating bias, and a bias in ability to learn enhanced male ornaments over reduced ornaments. Our findings demonstrate that females are able to change their preferences in response to a single social event, and suggest a role for biased learning in the evolution of visual sexual ornamentation.

  19. Near-Infrared Trigged Stimulus-Responsive Photonic Crystals with Hierarchical Structures.

    PubMed

    Lu, Tao; Pan, Hui; Ma, Jun; Li, Yao; Zhu, Shenmin; Zhang, Di

    2017-10-04

    Stimuli-responsive photonic crystals (PCs) trigged by light would provide a novel intuitive and quantitative method for noninvasive detection. Inspired by the flame-detecting aptitude of fire beetles and the hierarchical photonic structures of butterfly wings, we herein developed near-infrared stimuli-responsive PCs through coupling photothermal Fe 3 O 4 nanoparticles with thermoresponsive poly(N-isopropylacrylamide) (PNIPAM), with hierarchical photonic structured butterfly wing scales as the template. The nanoparticles within 10 s transferred near-infrared radiation into heat that triggered the phase transition of PNIPAM; this almost immediately posed an anticipated effect on the PNIPAM refractive index and resulted in a composite spectrum change of ∼26 nm, leading to the direct visual readout. It is noteworthy that the whole process is durable and stable mainly owing to the chemical bonding formed between PNIPAM and the biotemplate. We envision that this biologically inspired approach could be utilized in a broad range of applications and would have a great impact on various monitoring processes and medical sensing.

  20. Polymorphic butterfly reveals the missing link in ecological speciation.

    PubMed

    Chamberlain, Nicola L; Hill, Ryan I; Kapan, Durrell D; Gilbert, Lawrence E; Kronforst, Marcus R

    2009-11-06

    Ecological speciation occurs when ecologically based, divergent selection causes the evolution of reproductive isolation. There are many empirical examples of this process; however, there exists a poorly characterized stage during which the traits that distinguish species ecologically and reproductively segregate in a single population. By using a combination of genetic mapping, mate-choice experiments, field observations, and population genetics, we studied a butterfly population with a mimetic wing color polymorphism and found that the butterflies exhibited partial, color-based, assortative mate preference. These traits represent the divergent, ecologically based signal and preference components of sexual isolation that usually distinguish incipient and sibling species. The association between behavior and recognition trait in a single population may enhance the probability of speciation and provides an example of the missing link between an interbreeding population and isolated species.

  1. Combining Taxonomic and Functional Approaches to Unravel the Spatial Distribution of an Amazonian Butterfly Community.

    PubMed

    Graça, Márlon B; Morais, José W; Franklin, Elizabeth; Pequeno, Pedro A C L; Souza, Jorge L P; Bueno, Anderson Saldanha

    2016-04-01

    This study investigated the spatial distribution of an Amazonian fruit-feeding butterfly assemblage by linking species taxonomic and functional approaches. We hypothesized that: 1) vegetation richness (i.e., resources) and abundance of insectivorous birds (i.e., predators) should drive changes in butterfly taxonomic composition, 2) larval diet breadth should decrease with increase of plant species richness, 3) small-sized adults should be favored by higher abundance of birds, and 4) communities with eyespot markings should be able to exploit areas with higher predation pressure. Fruit-feeding butterflies were sampled with bait traps and insect nets across 25 km(2) of an Amazonian ombrophilous forest in Brazil. We measured larval diet breadth, adult body size, and wing marking of all butterflies. Our results showed that plant species richness explained most of the variation in butterfly taxonomic turnover. Also, community average diet breadth decreased with increase of plant species richness, which supports our expectations. In contrast, community average body size increased with the abundance of birds, refuting our hypothesis. We detected no influence of environmental gradients on the occurrence of species with eyespot markings. The association between butterfly taxonomic and functional composition points to a mediator role of the functional traits in the environmental filtering of butterflies. The incorporation of the functional approach into the analyses allowed for the detection of relationships that were not observed using a strictly taxonomic perspective and provided an extra insight into comprehending the potential adaptive strategies of butterflies. © The Authors 2015. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For Permissions, please email: journals.permissions@oup.com.

  2. The Butterflies of Barro Colorado Island, Panama: Local Extinction since the 1930s.

    PubMed

    Basset, Yves; Barrios, Héctor; Segar, Simon; Srygley, Robert B; Aiello, Annette; Warren, Andrew D; Delgado, Francisco; Coronado, James; Lezcano, Jorge; Arizala, Stephany; Rivera, Marleny; Perez, Filonila; Bobadilla, Ricardo; Lopez, Yacksecari; Ramirez, José Alejandro

    2015-01-01

    Few data are available about the regional or local extinction of tropical butterfly species. When confirmed, local extinction was often due to the loss of host-plant species. We used published lists and recent monitoring programs to evaluate changes in butterfly composition on Barro Colorado Island (BCI, Panama) between an old (1923-1943) and a recent (1993-2013) period. Although 601 butterfly species have been recorded from BCI during the 1923-2013 period, we estimate that 390 species are currently breeding on the island, including 34 cryptic species, currently only known by their DNA Barcode Index Number. Twenty-three butterfly species that were considered abundant during the old period could not be collected during the recent period, despite a much higher sampling effort in recent times. We consider these species locally extinct from BCI and they conservatively represent 6% of the estimated local pool of resident species. Extinct species represent distant phylogenetic branches and several families. The butterfly traits most likely to influence the probability of extinction were host growth form, wing size and host specificity, independently of the phylogenetic relationships among butterfly species. On BCI, most likely candidates for extinction were small hesperiids feeding on herbs (35% of extinct species). However, contrary to our working hypothesis, extinction of these species on BCI cannot be attributed to loss of host plants. In most cases these host plants remain extant, but they probably subsist at lower or more fragmented densities. Coupled with low dispersal power, this reduced availability of host plants has probably caused the local extinction of some butterfly species. Many more bird than butterfly species have been lost from BCI recently, confirming that small preserves may be far more effective at conserving invertebrates than vertebrates and, therefore, should not necessarily be neglected from a conservation viewpoint.

  3. Investigating Nanoscopic Structures on a Butterfly Wing to Explore Solvation and Coloration

    ERIC Educational Resources Information Center

    Bober, Brittany A.; Ogata, Jennifer K.; Martinez, Veronica E.; Hallinan, Janae J.; Leach, Taylor A.; Negru, Bogdan

    2018-01-01

    Surface structures on the nanometer size scale can impart new and exciting properties to bulk materials. Nanoscopic structures on hydrophobic materials can result in superhydrophobicity and structural coloration. We present an interdisciplinary experiment that introduces undergraduate students to nanotechnology by manipulating the…

  4. Genetic resources in the USDA, ARS, PGRCU legume crop germplasm collections with phyto-pharmaceutical uses

    USDA-ARS?s Scientific Manuscript database

    Seventeen health functional legumes including butterfly pea (Clitoria ternatea L.), Indigofera cassioides Rottler ex DC., I. linnaei Ali, I. suffruticosa Mill., hyacinth bean [Lablab purpureus (L.) Sweet], velvetbean [Mucuna pruriens (L.) DC], jicama [Pachyrhizus erosus (L.) Urb.], winged bean [Psop...

  5. Swimming of a Tiny Subtropical Sea Butterfly with Coiled Shell

    NASA Astrophysics Data System (ADS)

    Murphy, David; Karakas, Ferhat; Maas, Amy

    2017-11-01

    Sea butterflies, also known as pteropods, include a variety of small, zooplanktonic marine snails. Thecosomatous pteropods possess a shell and swim at low Reynolds numbers by beating their wing-like parapodia in a manner reminiscent of insect flight. In fact, previous studies of the pteropod Limacina helicina have shown that pteropod swimming hydrodynamics and tiny insect flight aerodynamics are dynamically similar. Studies of L. helicina swimming have been performed in polar (0 degrees C) and temperate conditions (12 degrees C). Here we present measurements of the swimming of Heliconoides inflatus, a smaller yet morphologically similar pteropod that lives in warm Bermuda seawater (21 degrees C) with a viscosity almost half that of the polar seawater. The collected H. inflatus have shell sizes less than 1.5 mm in diameter, beat their wings at frequencies up to 11 Hz, and swim upwards in sawtooth trajectories at speeds up to approximately 25 mm/s. Using three-dimensional wing and body kinematics collected with two orthogonal high speed cameras and time-resolved, 2D flow measurements collected with a micro-PIV system, we compare the effects of smaller body size and lower water viscosity on the flow physics underlying flapping-based swimming by pteropods and flight by tiny insects.

  6. Observations on Leading-Edge Vortex Development

    NASA Astrophysics Data System (ADS)

    Glenn, Michael; Lang, Amy; Wahidi, Redha; Wilroy, Jacob

    2016-11-01

    Most of an insect's lift comes from the leading edge vortex (LEV) that they produce when flapping their wings. There are many variables that make a LEV either stronger or weaker such as: roughness from the scales on their wings, angle of attack (AoA) of wing, size of the wing, and speed of the wing during flapping motion. Experiments were conducted to study LEV development to gain a better understanding of butterfly flight and the importance of LEV formation. The variables emphasized in this particular experiment were the chord length Reynolds numbers. Two smooth plates of 4 inches and 7 inches were compared in this experiment with Re of 1500 and 3000. Matlab was used to track the LEV location and calculate the vorticity and circulation magnitudes. Differences in LEV vortex strength as a function of chord length will be presented. Funding was provided by NSF REU site Grant EEC 1358991 and CBET Grant 1628600.

  7. Effects of spatial heterogeneity on butterfly species richness in Rocky Mountain National Park, CO, USA

    USGS Publications Warehouse

    Kumar, S.; Simonson, S.E.; Stohlgren, T.J.

    2009-01-01

    We investigated butterfly responses to plot-level characteristics (plant species richness, vegetation height, and range in NDVI [normalized difference vegetation index]) and spatial heterogeneity in topography and landscape patterns (composition and configuration) at multiple spatial scales. Stratified random sampling was used to collect data on butterfly species richness from seventy-six 20 ?? 50 m plots. The plant species richness and average vegetation height data were collected from 76 modified-Whittaker plots overlaid on 76 butterfly plots. Spatial heterogeneity around sample plots was quantified by measuring topographic variables and landscape metrics at eight spatial extents (radii of 300, 600 to 2,400 m). The number of butterfly species recorded was strongly positively correlated with plant species richness, proportion of shrubland and mean patch size of shrubland. Patterns in butterfly species richness were negatively correlated with other variables including mean patch size, average vegetation height, elevation, and range in NDVI. The best predictive model selected using Akaike's Information Criterion corrected for small sample size (AICc), explained 62% of the variation in butterfly species richness at the 2,100 m spatial extent. Average vegetation height and mean patch size were among the best predictors of butterfly species richness. The models that included plot-level information and topographic variables explained relatively less variation in butterfly species richness, and were improved significantly after including landscape metrics. Our results suggest that spatial heterogeneity greatly influences patterns in butterfly species richness, and that it should be explicitly considered in conservation and management actions. ?? 2008 Springer Science+Business Media B.V.

  8. Earthworms, Stamps and Butterfly Wings: Encouraging Children's Interests and Collections.

    ERIC Educational Resources Information Center

    McGreevy, Ann

    2000-01-01

    This article examines the importance of encouraging children's interests and the pursuit of collections and hobbies as strategies for developing talent and abilities. Excerpts are cited from eminent people's lives as examples of early interests/collections and eventual success. Letters from children on their collections are included. (Contains…

  9. Butterfly gyroid nanostructures as a time-frozen glimpse of intracellular membrane development

    PubMed Central

    Wilts, Bodo D.; Apeleo Zubiri, Benjamin; Klatt, Michael A.; Butz, Benjamin; Fischer, Michael G.; Kelly, Stephen T.; Spiecker, Erdmann; Steiner, Ullrich; Schröder-Turk, Gerd E.

    2017-01-01

    The formation of the biophotonic gyroid material in butterfly wing scales is an exceptional feat of evolutionary engineering of functional nanostructures. It is hypothesized that this nanostructure forms by chitin polymerization inside a convoluted membrane of corresponding shape in the endoplasmic reticulum. However, this dynamic formation process, including whether membrane folding and chitin expression are simultaneous or sequential processes, cannot yet be elucidated by in vivo imaging. We report an unusual hierarchical ultrastructure in the butterfly Thecla opisena that, as a solid material, allows high-resolution three-dimensional microscopy. Rather than the conventional polycrystalline space-filling arrangement, a gyroid occurs in isolated facetted crystallites with a pronounced size gradient. When interpreted as a sequence of time-frozen snapshots of the morphogenesis, this arrangement provides insight into the formation mechanisms of the nanoporous gyroid material as well as of the intracellular organelle membrane that acts as the template. PMID:28508050

  10. Origin, development, and evolution of butterfly eyespots.

    PubMed

    Monteiro, Antónia

    2015-01-07

    This article reviews the latest developments in our understanding of the origin, development, and evolution of nymphalid butterfly eyespots. Recent contributions to this field include insights into the evolutionary and developmental origin of eyespots and their ancestral deployment on the wing, the evolution of eyespot number and eyespot sexual dimorphism, and the identification of genes affecting eyespot development and black pigmentation. I also compare features of old and more recently proposed models of eyespot development and propose a schematic for the genetic regulatory architecture of eyespots. Using this schematic I propose two hypotheses for why we observe limits to morphological diversity across these serially homologous traits.

  11. It's All in the Mix: Blend-Specific Behavioral Response to a Sexual Pheromone in a Butterfly

    PubMed Central

    Larsdotter-Mellström, Helena; Eriksson, Kerstin; Liblikas I, Ilme; Wiklund, Christer; Borg-Karlson, Anna K.; Nylin, Sören; Janz, Niklas; Carlsson, Mikael A.

    2016-01-01

    Among insects, sexual pheromones are typically mixtures of two to several components, all of which are generally required to elicit a behavioral response. Here we show for the first time that a complete blend of sexual pheromone components is needed to elicit a response also in a butterfly. Males of the Green-veined White, Pieris napi, emit an aphrodisiac pheromone, citral, from wing glands. This pheromone is requisite for females to accept mating with a courting male. Citral is a mixture of the two geometric isomers geranial (E-isomer) and neral (Z-isomer) in an approximate 1:1 ratio. We found that both these compounds are required to elicit acceptance behavior, which indicates synergistic interaction between processing of the isomers. Using functional Ca2+ imaging we found that geranial and neral evoke significantly different but overlapping glomerular activity patterns in the antennal lobe, which suggests receptors with different affinity for the two isomers. However, these glomeruli were intermingled with glomeruli responding to, for example, plant-related compounds, i.e., no distinct subpopulation of pheromone-responding glomeruli as in moths and other insects. In addition, these glomeruli showed lower specificity than pheromone-activated glomeruli in moths. We could, however, not detect any mixture interactions among four identified glomeruli, indicating that the synergistic effect may be generated at a higher processing level. Furthermore, correlations between glomerular activity patterns evoked by the single isomers and the blend did not change over time. PMID:26973536

  12. Winging It: Using Digital Imaging To Investigate Butterfly Metamorphosis

    ERIC Educational Resources Information Center

    Bowen, Anne; Bell, Randy L.

    2004-01-01

    One of the best ways to inspire interest in biology is through observations of living things. Unfortunately, this important component of science methodology is often left out because of the difficulty of including it in the classroom. Additionally, amazing processes occur in nature that few have the chance to observe. This article reviews a…

  13. From the Meticulous to the Sublime: Transformations in Ink.

    ERIC Educational Resources Information Center

    King, Steve

    1998-01-01

    Provides a lesson that gives students the opportunity to master the medium of pen and ink through a project called "Transformations." Explains that students choose two pictures, copying each one, and then combine them to create a transformation where one object transforms into the other, such as a bolt with butterfly wings. (CMK)

  14. The Wind beneath Our Wings: Chaos Theory and the Butterfly Effect in Curriculum Design.

    ERIC Educational Resources Information Center

    Grover, Robert; Achleitner, Herbert; Thomas, Nancy; Wyatt, Roger; Vowell, Faye N.

    1997-01-01

    Discussion of change and its influence on information professionals focuses on curriculum changes needed and implemented in library and information science education at Emporia State University. Topics include social context and shifts in perspective; postmodernism; dynamics of information technology and the global information economy; and a…

  15. "All Flying Insects with Big, Beautiful Wings are Butterflies!" A Study in Challenging This Misconception

    ERIC Educational Resources Information Center

    Tsoi, Kwok-Ho

    2013-01-01

    This study investigated the level of understanding among student teachers in differentiating lepidopterans. It adopted a constructive approach to promoting conceptual change in students on the issue of animal classification by generating cognitive conflict. Most of the students used inaccurate morphological traits for identification, such as wing…

  16. Geometry in Biomimetic Network: Double Gyroid to Pseudo-Single Gyroid in Nanohybrid Materials

    NASA Astrophysics Data System (ADS)

    Hsueh, Han-Yu; Ho, Rong-Ming; Hung, Yu-Chueh; Ling, Yi-Chun; Hasegawa, Hirokazu

    2013-03-01

    Biological systems have developed delicately arranged micro- and architectures to produce striking optical effects since millions of years ago. Inspired by the textures of butterfly wings with single gyroid (SG) structure, herein, we aim to fabricate biocompatible and robust materials with SG-like structure in nanometer size so as to give new materials with unprecedented optical properties for applications. Biommicking from the biological photonic structures of butterfly wings, a double gyroid (DG) structure in nanometer size is obtained from the self-assembly of polystyrene-b-poly(L-lactide) (PS-PLLA). To acquire robust backbone networks, inorganic networks in polymer matrix are fabricated by using the hydrolyzed PS-PLLA with DG structure as a template for sol-gel reaction. Owing to the soft polymer matrix, two co-continuous inorganic networks embedded in the polymer matrix can be rearranged by thermal annealing at temperature above the glass transition of the polymer. Consequently, the rearrangement of these inorganic networks leads the formation of SG-like structure possessing unique nanohybrids with ordered texture. This unique nanomaterials with SG-like structure is referred as a pseudo-SG (p-SG) nanohybrids.

  17. A simple and effective approach towards biomimetic replication of photonic structures from butterfly wings.

    PubMed

    Zhu, Shenmin; Zhang, Di; Chen, Zhixin; Gu, Jiajun; Li, Wenfei; Jiang, Haibo; Zhou, Gang

    2009-08-05

    A general sonochemical process is reported for the replication of photonic structures from Morpho butterfly wings in several hours. By selecting appropriate precursors, we can achieve exact replications of photonic structures in a variety of transparent metal oxides, such as titania, tin oxide and silica. The exact replications at the micro- and nanoscales were characterized by a combination of FE-SEM, TEM, EDX and Raman measurements. The optical properties of the replicas were investigated by using reflectance spectroscopy, and it was found that the interesting chromaticity of the reflected light could be adjusted simply by tuning the replica materials. An ultrasensitive SnO(2)-based chemical sensor was prepared from the SnO(2) replica. The sensor has a sensitivity of 35.3-50 ppm ethanol at 300 degrees C, accompanied by a rapid response and recovery (around 8-15 s), owing to its large surface area and photonic structure. Thus, this process could be developed to produce photonic structural ceramics which could be used in many passive and active infrared devices, especially high performance optical components and sensors.

  18. The Butterflies of Barro Colorado Island, Panama: Local Extinction since the 1930s

    PubMed Central

    Basset, Yves; Barrios, Héctor; Segar, Simon; Srygley, Robert B.; Aiello, Annette; Warren, Andrew D.; Delgado, Francisco; Coronado, James; Lezcano, Jorge; Arizala, Stephany; Rivera, Marleny; Perez, Filonila; Bobadilla, Ricardo; Lopez, Yacksecari; Ramirez, José Alejandro

    2015-01-01

    Few data are available about the regional or local extinction of tropical butterfly species. When confirmed, local extinction was often due to the loss of host-plant species. We used published lists and recent monitoring programs to evaluate changes in butterfly composition on Barro Colorado Island (BCI, Panama) between an old (1923–1943) and a recent (1993–2013) period. Although 601 butterfly species have been recorded from BCI during the 1923–2013 period, we estimate that 390 species are currently breeding on the island, including 34 cryptic species, currently only known by their DNA Barcode Index Number. Twenty-three butterfly species that were considered abundant during the old period could not be collected during the recent period, despite a much higher sampling effort in recent times. We consider these species locally extinct from BCI and they conservatively represent 6% of the estimated local pool of resident species. Extinct species represent distant phylogenetic branches and several families. The butterfly traits most likely to influence the probability of extinction were host growth form, wing size and host specificity, independently of the phylogenetic relationships among butterfly species. On BCI, most likely candidates for extinction were small hesperiids feeding on herbs (35% of extinct species). However, contrary to our working hypothesis, extinction of these species on BCI cannot be attributed to loss of host plants. In most cases these host plants remain extant, but they probably subsist at lower or more fragmented densities. Coupled with low dispersal power, this reduced availability of host plants has probably caused the local extinction of some butterfly species. Many more bird than butterfly species have been lost from BCI recently, confirming that small preserves may be far more effective at conserving invertebrates than vertebrates and, therefore, should not necessarily be neglected from a conservation viewpoint. PMID:26305111

  19. Biomimetic gyroid nanostructures exceeding their natural origins.

    PubMed

    Gan, Zongsong; Turner, Mark D; Gu, Min

    2016-05-01

    Using optical two-beam lithography with improved resolution and enhanced mechanical strength, we demonstrate the replication of gyroid photonic nanostructures found in the butterfly Callophrys rubi. These artificial structures are shown to have size, controllability, and uniformity that are superior to those of their biological counterparts. In particular, the elastic Young's modulus of fabricated nanowires is enhanced by up to 20%. As such, the circular dichroism enabled by the gyroid nanostructures can operate in the near-ultraviolet wavelength region, shorter than that supported by the natural butterfly wings of C. rubi. This fabrication technique provides a unique tool for extracting three-dimensional photonic designs from nature and will aid the investigation of biomimetic nanostructures.

  20. Biomimetic gyroid nanostructures exceeding their natural origins

    PubMed Central

    Gan, Zongsong; Turner, Mark D.; Gu, Min

    2016-01-01

    Using optical two-beam lithography with improved resolution and enhanced mechanical strength, we demonstrate the replication of gyroid photonic nanostructures found in the butterfly Callophrys rubi. These artificial structures are shown to have size, controllability, and uniformity that are superior to those of their biological counterparts. In particular, the elastic Young’s modulus of fabricated nanowires is enhanced by up to 20%. As such, the circular dichroism enabled by the gyroid nanostructures can operate in the near-ultraviolet wavelength region, shorter than that supported by the natural butterfly wings of C. rubi. This fabrication technique provides a unique tool for extracting three-dimensional photonic designs from nature and will aid the investigation of biomimetic nanostructures. PMID:27386542

  1. Insect Thin Films as Sun Blocks, Not Solar Collectors

    NASA Astrophysics Data System (ADS)

    Koon, Daniel W.; Crawford, Andrew B.

    2000-05-01

    We measured the visible reflectance spectra of whole wing sections from three species of iridescent butterflies and moths, for normal incidence, integrated over all reflected angles. In this manner, we separated the optics of the thin films causing the iridescence from the optics of the rest of the scale. We found that iridescence reduces solar absorption by the wing in all cases, typically by approximately 20% or less, in contrast to claims by Miaoulis and Heilman Ann. Entomol. Soc. Am. 91, 122 (1998) that the thin-film structures that produce iridescence act as solar collectors.

  2. Spectroscopy on the wing: naturally inspired SERS substrates for biochemical analysis.

    PubMed

    Garrett, Natalie L; Vukusic, Peter; Ogrin, Feodor; Sirotkin, Evgeny; Winlove, C Peter; Moger, Julian

    2009-03-01

    We show that naturally occurring chitinous nanostructures found on the wings of the Graphium butterfly can be used as substrates for surface-enhanced Raman scattering when coated with a thin film of gold or silver. The substrates were found to exhibit excellent biocompatibility and sensitivity, making them ideal for protein assaying. An assay using avidin/biotin binding showed that the substrates could be used to quantify protein binding directly from changes in the surface-enhanced Raman scattering (SERS) spectra and were sensitive over a concentration range comparable with a typical enzyme-linked immunosorbent assays (ELISA) assay. A biomimetic version of the wing nanostructures produced using a highly reproducible, large-scale fabrication process, yielded comparable enhancement factors and biocompatibility. The excellent biocompatibility of the wings and biomimetic substrates is unparalleled by other lithographically produced substrates, and this could pave the way for widespread application of ultrasensitive SERS-based bioassays.

  3. Variation in wing pattern and palatability in a female-limited polymorphic mimicry system

    PubMed Central

    Long, Elizabeth C; Hahn, Thomas P; Shapiro, Arthur M

    2014-01-01

    Checkerspot butterflies in the genera Euphydryas and Chlosyne exhibit phenotypic polymorphisms along a well-defined latitudinal and elevational gradient in California. The patterns of phenotypic variation in Euphydryas chalcedona, Chlosyne palla, and Chlosyne hoffmanni suggest a mimetic relationship; in addition, the specific patterns of variation in C. palla suggest a female-limited polymorphic mimicry system (FPM). However, the existence of polymorphic models runs counter to predictions of mimicry theory. Palatability trials were undertaken to assess whether or not the different color morphs of each species were distasteful or toxic to a generalized avian predator, the European starling (Sturnus vulgaris). Results indicate that the black morph of E. chalcedona is distasteful, but not toxic, to predators, while the red morph is palatable. C . hoffmanni and both color morphs of C. palla are palatable to predators. Predators that learn to reject black E. chalcedona also reject black C. palla, suggesting that the latter is a FPM of the former. C. hoffmanni does not appear to be involved in this mimetic relationship. PMID:25512850

  4. Predator mimicry, not conspicuousness, explains the efficacy of butterfly eyespots

    PubMed Central

    De Bona, Sebastiano; Valkonen, Janne K.; López-Sepulcre, Andrés; Mappes, Johanna

    2015-01-01

    Large conspicuous eyespots on butterfly wings have been shown to deter predators. This has been traditionally explained by mimicry of vertebrate eyes, but recently the classic eye-mimicry hypothesis has been challenged. It is proposed that the conspicuousness of the eyespot, not mimicry, is what causes aversion due to sensory biases, neophobia or sensory overloads. We conducted an experiment to directly test whether the eye-mimicry or the conspicuousness hypothesis better explain eyespot efficacy. We used great tits (Parus major) as model predator, and tested their reaction towards animated images on a computer display. Birds were tested against images of butterflies without eyespots, with natural-looking eyespots, and manipulated spots with the same contrast but reduced resemblance to an eye, as well as images of predators (owls) with and without eyes. We found that mimetic eyespots were as effective as true eyes of owls and more efficient in eliciting an aversive response than modified, less mimetic but equally contrasting eyespots. We conclude that the eye-mimicry hypothesis explains our results better than the conspicuousness hypothesis and is thus likely to be an important mechanism behind the evolution of butterfly eyespots. PMID:25854889

  5. The effect of female wings on male courtship behavior in the cricket Gryllus bimaculatus

    NASA Astrophysics Data System (ADS)

    Itoh, Masanori; Murakami, Shio

    2002-03-01

    Sexual dimorphism exists in the shape and the structure of the forewings of the cricket (Gryllus bimaculatus). However, the functional significance of the wings in the G. bimaculatus female has been unclear. In common blue butterflies (Polyommatus icarus), wings in females have been suggested as being important for attracting males. To test whether female crickets need wings for conspecific males to recognize them and initiate mating behavior, we removed all wings from females and observed the behavior of males towards them. Most males (87.5%) showed mating behavior towards the wingless females: they produced courtship song and transferred spermatophores to the wingless females. Similarly, 88.5% of the males showed mating behavior towards intact females. When males were placed with both a wingless female and an intact female, no significant difference was detected in male mate choice. The findings demonstrate that the wing of the G. bimaculatus female is not necessary for female recognition by conspecific males and the initiation of male mating behavior, and that it is not important in male mate choice.

  6. Bauhinia monandra Kurz.

    Treesearch

    K.F Connor

    2002-01-01

    Bauhinia monandra is a small fast growing, evergreen tree or shrub that commonly reaches 3 to 15.2 m in height and 0.5 m in diameter. Its smooth, gray bark can become scaly and red brown on older trees. The leaves of B. monandra are shaped like butterfly wings, rounded, and split 1/3 to 1/2 their length, forming two equal lobes...

  7. Photonic Crystals on the Wing

    DTIC Science & Technology

    2009-04-30

    The Green Hairstreak butterfly has ventrally scales consisting of 3D gyroid domains acting as photonic crystals, which can be easily...study on the structural coloration of the Green Hairstreak , Callophrys rubi. Extensive experimental measurements have been complemented with three...colorations, produced by purely dielectric materials are widely encountered [1-3]. Famous examples are the metallic- green buprestid and golden scarab

  8. Strapping Wings on a Caterpillar and Calling It a Butterfly: When Systemic Change Is Not Systemic

    ERIC Educational Resources Information Center

    Duffy, Francis M.

    2008-01-01

    This article challenges misperceptions about the definition of systemic change in school districts. While many contemporary change efforts are described as "systemic," in fact, they are not. The author also argues that a special instance of systemic change known as systemic transformational change is required to transform entire school…

  9. Thermoregulation and the determinants of heat transfer in Colias butterflies.

    PubMed

    Kingsolver, Joel G; Moffat, Robert J

    1982-04-01

    As a means of exploring behavioral and morphological adaptations for thermoregulation in Colias butterflies, convective heat transfer coefficients of real and model butterflies were measured in a wind tunnel as a function of wind speed and body orientation (yaw angle). Results are reported in terms of a dimensionless heat transfer coefficient (Nusselt number, Nu) and a dimensionless wind speed (Reynolds number, Re), for a wind speed range typical of that experienced by basking Colias in the field. The resultant Nusselt-Reynolds (Nu-Re) plots thus indicate the rates of heat transfer by forced convection as a function of wind speed for particular model geometries.For Reynolds numbers throughout the measured range, Nusselt numbers for C. eurytheme butterflies are consistently lower than those for long cylinders, and are independent of yaw angle. There is significant variation among individual butterflies in heat transfer coefficients throughout the Re range. Model butterflies without artificial fur have Nu-Re relations similar to those for cylinders. Heat transfer in these models depends upon yaw angle, with higher heat transfer at intermediate yaw angles (30-60°); these yaw effects increase with increasing Reynolds number. Models with artificial fur, like real Colias, have Nusselt numbers which are consistently lower than those for models without fur at given Reynolds numbers throughout the Re range. Unlike real Colias, however, the models with fur do show yaw angle effects similar to those for models without fur.The independence of heat loss from yaw angle for real Colias is consistent with field observations indicating no behavioral orientation to wind direction. The presence of fur on the models reduces heat loss but does not affect yaw dependence. The large individual variation in heat transfer coefficients among butterflies is probably due to differences in fur characteristics rather than to differences in wing morphology.Finally, a physical model of a butterfly was constructed which accurately simulates the body temperatures of basking Colias in the field for a variety of radiation and wind velocity conditions. The success of the butterfly simulator in mimicking Colias thermal characteristics confirms our preliminary understanding of the physical bases for and heat transfer mechanisms underlying thermoregulatory adaptations in these butterflies.

  10. The evolution of movements and behaviour at boundaries in different landscapes: a common arena experiment with butterflies.

    PubMed Central

    Merckx, Thomas; Van Dyck, Hans; Karlsson, Bengt; Leimar, Olof

    2003-01-01

    As landscapes change, mobility patterns of species may alter. Different mechanistic scenarios may, however, lead to particular patterns. Here, we tested conflicting predictions from two hypotheses on butterfly movements in relation to habitat fragmentation. According to the resource distribution hypothesis, butterflies in more fragmented landscapes would have higher levels of mobility as resources are more scattered. However, these butterflies could have lower levels of mobility as they experience 'hard' habitat boundaries more frequently (i.e. higher crossing costs) compared with butterflies in landscapes with continuous habitat; i.e. the behaviour-at-boundaries hypothesis. We studied movements, habitat boundary crossing and habitat preference of laboratory-reared individuals of Pararge aegeria that originated from woodland and agricultural landscapes, by using an experimental landscape as a common environment (outdoor cages) to test the predictions, taking into account sexual differences and weather. Woodland butterflies covered longer distances, were more prone to cross open-shade boundaries, travelled more frequently between woodland parts of the cages and were more at flight than agricultural butterflies. Our results support the behaviour-at-boundaries hypothesis, with 'softer' boundaries for woodland landscapes. Because the butterflies were reared in a common environment, the observed behavioural differences rely on heritable variation between populations from woodland and agricultural landscapes. PMID:12964984

  11. Adaptive developmental plasticity: compartmentalized responses to environmental cues and to corresponding internal signals provide phenotypic flexibility.

    PubMed

    Mateus, Ana Rita A; Marques-Pita, Manuel; Oostra, Vicencio; Lafuente, Elvira; Brakefield, Paul M; Zwaan, Bas J; Beldade, Patrícia

    2014-11-21

    The environmental regulation of development can result in the production of distinct phenotypes from the same genotype and provide the means for organisms to cope with environmental heterogeneity. The effect of the environment on developmental outcomes is typically mediated by hormonal signals which convey information about external cues to the developing tissues. While such plasticity is a wide-spread property of development, not all developing tissues are equally plastic. To understand how organisms integrate environmental input into coherent adult phenotypes, we must know how different body parts respond, independently or in concert, to external cues and to the corresponding internal signals. We quantified the effect of temperature and ecdysone hormone manipulations on post-growth tissue patterning in an experimental model of adaptive developmental plasticity, the butterfly Bicyclus anynana. Following a suite of traits evolving by natural or sexual selection, we found that different groups of cells within the same tissue have sensitivities and patterns of response that are surprisingly distinct for the external environmental cue and for the internal hormonal signal. All but those wing traits presumably involved in mate choice responded to developmental temperature and, of those, all but the wing traits not exposed to predators responded to hormone manipulations. On the other hand, while patterns of significant response to temperature contrasted traits on autonomously-developing wings, significant response to hormone manipulations contrasted neighboring groups of cells with distinct color fates. We also showed that the spatial compartmentalization of these responses cannot be explained by the spatial or temporal compartmentalization of the hormone receptor protein. Our results unravel the integration of different aspects of the adult phenotype into developmental and functional units which both reflect and impact evolutionary change. Importantly, our findings underscore the complexity of the interactions between environment and physiology in shaping the development of different body parts.

  12. Contribution of urban expansion and a changing climate to decline of a butterfly fauna.

    PubMed

    Casner, Kayce L; Forister, Matthew L; O'Brien, Joshua M; Thorne, James; Waetjen, David; Shapiro, Arthur M

    2014-06-01

    Butterfly populations are naturally patchy and undergo extinctions and recolonizations. Analyses based on more than 2 decades of data on California's Central Valley butterfly fauna show a net loss in species richness through time. We analyzed 22 years of phenological and faunistic data for butterflies to investigate patterns of species richness over time. We then used 18-22 years of data on changes in regional land use and 37 years of seasonal climate data to develop an explanatory model. The model related the effects of changes in land-use patterns, from working landscapes (farm and ranchland) to urban and suburban landscapes, and of a changing climate on butterfly species richness. Additionally, we investigated local trends in land use and climate. A decline in the area of farmland and ranchland, an increase in minimum temperatures during the summer and maximum temperatures in the fall negatively affected net species richness, whereas increased minimum temperatures in the spring and greater precipitation in the previous summer positively affected species richness. According to the model, there was a threshold between 30% and 40% working-landscape area below which further loss of working-landscape area had a proportionally greater effect on butterfly richness. Some of the isolated effects of a warming climate acted in opposition to affect butterfly richness. Three of the 4 climate variables that most affected richness showed systematic trends (spring and summer mean minimum and fall mean maximum temperatures). Higher spring minimum temperatures were associated with greater species richness, whereas higher summer temperatures in the previous year and lower rainfall were linked to lower richness. Patterns of land use contributed to declines in species richness (although the pattern was not linear), but the net effect of a changing climate on butterfly richness was more difficult to discern. © 2014 Society for Conservation Biology.

  13. Do monarch butterflies use polarized skylight for migratory orientation?

    PubMed

    Stalleicken, Julia; Mukhida, Maya; Labhart, Thomas; Wehner, Rüdiger; Frost, Barrie; Mouritsen, Henrik

    2005-06-01

    To test if migratory monarch butterflies use polarized light patterns as part of their time-compensated sun compass, we recorded their virtual flight paths in a flight simulator while the butterflies were exposed to patches of naturally polarized blue sky, artificial polarizers or a sunny sky. In addition, we tested butterflies with and without the polarized light detectors of their compound eye being occluded. The monarchs' orientation responses suggested that the butterflies did not use the polarized light patterns as a compass cue, nor did they exhibit a specific alignment response towards the axis of polarized light. When given direct view of the sun, migratory monarchs with their polarized light detectors painted out were still able to use their time-compensated compass: non-clockshifted butterflies, with their dorsal rim area occluded, oriented in their typical south-southwesterly migratory direction. Furthermore, they shifted their flight course clockwise by the predicted approximately 90 degrees after being advance clockshifted 6 h. We conclude that in migratory monarch butterflies, polarized light cues are not necessary for a time-compensated celestial compass to work and that the azimuthal position of the sun disc and/or the associated light-intensity and spectral gradients seem to be the migrants' major compass cue.

  14. Rapid assessment of butterfly diversity in a montane landscape

    USGS Publications Warehouse

    Simonson, S.E.; Opler, P.A.; Stohlgren, T.J.; Chong, G.W.

    2001-01-01

    We present the results of a rapid assessment of butterfly diversity in the 754 ha Beaver Meadows study area in Rocky Mountain National Park, Larimer County, Colorado. We measured butterfly species richness and relative abundance as part of a landscape-scale investigation of diversity patterns involving several groups of organisms. A stratified random sampling design was used to include replication in both rare and common vegetation types. We recorded 49 butterfly species from the twenty-four 0.1 ha plots that were sampled four times during June, July, and August 1996. Butterfly species richness, diversity, and uniqueness were highest in quaking aspen (Populus tremuloides Michaux) groves and wet meadows, which occupy only a small proportion of the studied landscape. This result supports the suggestion that aspen areas represent ‘hotspots’ of biological diversity in this montane landscape. Patterns of butterfly species richness were positively correlated with total vascular plant species richness (r = 0.69; P < 0.001), and native plant species richness (r = 0.64; P < 0.001). However, exotic plant species richness (r = 0.70; P < 0.001) and the cover of exotic plant species (r = 0.70; P < 0.001) were the best predictors of butterfly species richness.

  15. 25th anniversary article: scalable multiscale patterned structures inspired by nature: the role of hierarchy.

    PubMed

    Bae, Won-Gyu; Kim, Hong Nam; Kim, Doogon; Park, Suk-Hee; Jeong, Hoon Eui; Suh, Kahp-Yang

    2014-02-01

    Multiscale, hierarchically patterned surfaces, such as lotus leaves, butterfly wings, adhesion pads of gecko lizards are abundantly found in nature, where microstructures are usually used to strengthen the mechanical stability while nanostructures offer the main functionality, i.e., wettability, structural color, or dry adhesion. To emulate such hierarchical structures in nature, multiscale, multilevel patterning has been extensively utilized for the last few decades towards various applications ranging from wetting control, structural colors, to tissue scaffolds. In this review, we highlight recent advances in scalable multiscale patterning to bring about improved functions that can even surpass those found in nature, with particular focus on the analogy between natural and synthetic architectures in terms of the role of different length scales. This review is organized into four sections. First, the role and importance of multiscale, hierarchical structures is described with four representative examples. Second, recent achievements in multiscale patterning are introduced with their strengths and weaknesses. Third, four application areas of wetting control, dry adhesives, selectively filtrating membranes, and multiscale tissue scaffolds are overviewed by stressing out how and why multiscale structures need to be incorporated to carry out their performances. Finally, we present future directions and challenges for scalable, multiscale patterned surfaces. © 2013 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim.

  16. Congruence and diversity of butterfly-host plant associations at higher taxonomic levels.

    PubMed

    Ferrer-Paris, José R; Sánchez-Mercado, Ada; Viloria, Ángel L; Donaldson, John

    2013-01-01

    We aggregated data on butterfly-host plant associations from existing sources in order to address the following questions: (1) is there a general correlation between host diversity and butterfly species richness?, (2) has the evolution of host plant use followed consistent patterns across butterfly lineages?, (3) what is the common ancestral host plant for all butterfly lineages? The compilation included 44,148 records from 5,152 butterfly species (28.6% of worldwide species of Papilionoidea) and 1,193 genera (66.3%). The overwhelming majority of butterflies use angiosperms as host plants. Fabales is used by most species (1,007 spp.) from all seven butterfly families and most subfamilies, Poales is the second most frequently used order, but is mostly restricted to two species-rich subfamilies: Hesperiinae (56.5% of all Hesperiidae), and Satyrinae (42.6% of all Nymphalidae). We found a significant and strong correlation between host plant diversity and butterfly species richness. A global test for congruence (Parafit test) was sensitive to uncertainty in the butterfly cladogram, and suggests a mixed system with congruent associations between Papilionidae and magnoliids, Hesperiidae and monocots, and the remaining subfamilies with the eudicots (fabids and malvids), but also numerous random associations. The congruent associations are also recovered as the most probable ancestral states in each node using maximum likelihood methods. The shift from basal groups to eudicots appears to be more likely than the other way around, with the only exception being a Satyrine-clade within the Nymphalidae that feed on monocots. Our analysis contributes to the visualization of the complex pattern of interactions at superfamily level and provides a context to discuss the timing of changes in host plant utilization that might have promoted diversification in some butterfly lineages.

  17. Congruence and Diversity of Butterfly-Host Plant Associations at Higher Taxonomic Levels

    PubMed Central

    Ferrer-Paris, José R.; Sánchez-Mercado, Ada; Viloria, Ángel L.; Donaldson, John

    2013-01-01

    We aggregated data on butterfly-host plant associations from existing sources in order to address the following questions: (1) is there a general correlation between host diversity and butterfly species richness?, (2) has the evolution of host plant use followed consistent patterns across butterfly lineages?, (3) what is the common ancestral host plant for all butterfly lineages? The compilation included 44,148 records from 5,152 butterfly species (28.6% of worldwide species of Papilionoidea) and 1,193 genera (66.3%). The overwhelming majority of butterflies use angiosperms as host plants. Fabales is used by most species (1,007 spp.) from all seven butterfly families and most subfamilies, Poales is the second most frequently used order, but is mostly restricted to two species-rich subfamilies: Hesperiinae (56.5% of all Hesperiidae), and Satyrinae (42.6% of all Nymphalidae). We found a significant and strong correlation between host plant diversity and butterfly species richness. A global test for congruence (Parafit test) was sensitive to uncertainty in the butterfly cladogram, and suggests a mixed system with congruent associations between Papilionidae and magnoliids, Hesperiidae and monocots, and the remaining subfamilies with the eudicots (fabids and malvids), but also numerous random associations. The congruent associations are also recovered as the most probable ancestral states in each node using maximum likelihood methods. The shift from basal groups to eudicots appears to be more likely than the other way around, with the only exception being a Satyrine-clade within the Nymphalidae that feed on monocots. Our analysis contributes to the visualization of the complex pattern of interactions at superfamily level and provides a context to discuss the timing of changes in host plant utilization that might have promoted diversification in some butterfly lineages. PMID:23717448

  18. [History and present status of butterfly monitoring in Europe and related development strategies for China].

    PubMed

    Fang, Li-Jun; Xu, Hai-Gen; Guan, Jian-Ling

    2013-09-01

    Butterfly is an important bio-indicator for biodiversity monitoring and ecological environment assessment. In Europe, the species composition, population dynamics, and distribution pattern of butterfly have been monitored for decades, and many long-term monitoring schemes with international effects have been implemented. These schemes are aimed to assess the regional and national variation trends of butterfly species abundance, and to analyze the relationships of this species abundance with habitat, climate change, and other environmental factors, providing basic data for researching, protecting, and utilizing butterfly resources and predicting environmental changes, and playing important roles in the division of butterfly' s threatened level, the formulation of related protection measures, and the protection and management of ecological environment. This paper reviewed the history and present status of butterfly monitoring in Europe, with the focus on the well-known long-term monitoring programs, e. g. , the UK Butterfly Monitoring Scheme and the Germany and European Union Butterfly Monitoring Scheme. Some specific proposals for conducting butterflies monitoring in China were suggested.

  19. Estimating Brownian motion dispersal rate, longevity and population density from spatially explicit mark-recapture data on tropical butterflies.

    PubMed

    Tufto, Jarle; Lande, Russell; Ringsby, Thor-Harald; Engen, Steinar; Saether, Bernt-Erik; Walla, Thomas R; DeVries, Philip J

    2012-07-01

    1. We develop a Bayesian method for analysing mark-recapture data in continuous habitat using a model in which individuals movement paths are Brownian motions, life spans are exponentially distributed and capture events occur at given instants in time if individuals are within a certain attractive distance of the traps. 2. The joint posterior distribution of the dispersal rate, longevity, trap attraction distances and a number of latent variables representing the unobserved movement paths and time of death of all individuals is computed using Gibbs sampling. 3. An estimate of absolute local population density is obtained simply by dividing the Poisson counts of individuals captured at given points in time by the estimated total attraction area of all traps. Our approach for estimating population density in continuous habitat avoids the need to define an arbitrary effective trapping area that characterized previous mark-recapture methods in continuous habitat. 4. We applied our method to estimate spatial demography parameters in nine species of neotropical butterflies. Path analysis of interspecific variation in demographic parameters and mean wing length revealed a simple network of strong causation. Larger wing length increases dispersal rate, which in turn increases trap attraction distance. However, higher dispersal rate also decreases longevity, thus explaining the surprising observation of a negative correlation between wing length and longevity. © 2012 The Authors. Journal of Animal Ecology © 2012 British Ecological Society.

  20. Biological Inspiration for Agile Autonomous Air Vehicles

    DTIC Science & Technology

    2007-11-01

    rhythmically contract the thorax; the hind wings have become specialized as small body rotation sensors (halteres). Butterflies and moths have two pairs of...orthogonal pairs of power muscles that produce alternating dorso-ventral and longitudinal flexure of the thorax from rhythmic contractions similar to...other physical sciences lend themselves to somewhat reductionist approaches for both analysis and synthesis. Complex engineered systems are built from

  1. Q&A: The nanomaterials designer

    NASA Astrophysics Data System (ADS)

    Gibney, Elizabeth

    2015-10-01

    Ali Yetisen's research includes using nanotechnology and biosensors to make environmentally responsive materials for clothes, tattoos, accessories and contact lenses -- materials that could be the future of fashion. Here, Yetisen, who works at Harvard University and Massachusetts General Hospital in Cambridge, talks about mimicking the diffraction in butterfly wings, transforming gowns, and what fashion designers and materials scientists can learn from each other.

  2. Temperature and saturation dependence in the vapor sensing of butterfly wing scales.

    PubMed

    Kertész, K; Piszter, G; Jakab, E; Bálint, Zs; Vértesy, Z; Biró, L P

    2014-06-01

    The sensing of gasses/vapors in the ambient air is the focus of attention due to the need to monitor our everyday environment. Photonic crystals are sensing materials of the future because of their strong light-manipulating properties. Natural photonic structures are well-suited materials for testing detection principles because they are significantly cheaper than artificial photonic structures and are available in larger sizes. Additionally, natural photonic structures may provide new ideas for developing novel artificial photonic nanoarchitectures with improved properties. In the present paper, we discuss the effects arising from the sensor temperature and the vapor concentration in air during measurements with a photonic crystal-type optical gas sensor. Our results shed light on the sources of discrepancy between simulated and experimental sensing behaviors of photonic crystal-type structures. Through capillary condensation, the vapors will condensate to a liquid state inside the nanocavities. Due to the temperature and radius of curvature dependence of capillary condensation, the measured signals are affected by the sensor temperature as well as by the presence of a nanocavity size distribution. The sensing materials used are natural photonic nanoarchitectures present in the wing scales of blue butterflies. Copyright © 2014 Elsevier B.V. All rights reserved.

  3. Color change of Blue butterfly wing scales in an air - Vapor ambient

    NASA Astrophysics Data System (ADS)

    Kertész, Krisztián; Piszter, Gábor; Jakab, Emma; Bálint, Zsolt; Vértesy, Zofia; Biró, László Péter

    2013-09-01

    Photonic crystals are periodic dielectric nanocomposites, which have photonic band gaps that forbid the propagation of light within certain frequency ranges. The optical response of such nanoarchitectures on chemical changes in the environment is determined by the spectral change of the reflected light, and depends on the composition of the ambient atmosphere and on the nanostructure characteristics. We carried out reflectance measurements on closely related Blue lycaenid butterfly males possessing so-called "pepper-pot" type photonic nanoarchitecture in their scales covering their dorsal wing surfaces. Experiments were carried out changing the concentration and nature of test vapors while monitoring the spectral variations in time. All the tests were done with the sample temperature set at, and below the room temperature. The spectral changes were found to be linear with the increasing of concentration and the signal amplitude is higher at lower temperatures. The mechanism of reflectance spectra modification is based on capillary condensation of the vapors penetrating in the nanostructure. These structures of natural origin may serve as cheap, environmentally free and biodegradable sensor elements. The study of these nanoarchitectures of biologic origin could be the source of various new bioinspired systems.

  4. Polarization-sensitive color in butterfly scales: polarization conversion from ridges with reflecting elements.

    PubMed

    Zhang, Ke; Tang, Yiwen; Meng, Jinsong; Wang, Ge; Zhou, Han; Fan, Tongxiang; Zhang, Di

    2014-11-03

    Polarization-sensitive color originates from polarization-dependent reflection or transmission, exhibiting abundant light information, including intensity, spectral distribution, and polarization. A wide range of butterflies are physiologically sensitive to polarized light, but the origins of polarized signal have not been fully understood. Here we systematically investigate the colorful scales of six species of butterfly to reveal the physical origins of polarization-sensitive color. Microscopic optical images under crossed polarizers exhibit their polarization-sensitive characteristic, and micro-structural characterizations clarify their structural commonality. In the case of the structural scales that have deep ridges, the polarization-sensitive color related with scale azimuth is remarkable. Periodic ridges lead to the anisotropic effective refractive indices in the parallel and perpendicular grating orientations, which achieves form-birefringence, resulting in the phase difference of two different component polarized lights. Simulated results show that ridge structures with reflecting elements reflect and rotate the incident p-polarized light into s-polarized light. The dimensional parameters and shapes of grating greatly affect the polarization conversion process, and the triangular deep grating extends the outstanding polarization conversion effect from the sub-wavelength period to the period comparable to visible light wavelength. The parameters of ridge structures in butterfly scales have been optimized to fulfill the polarization-dependent reflection for secret communication. The structural and physical origin of polarization conversion provides a more comprehensive perspective on the creation of polarization-sensitive color in butterfly wing scales. These findings show great potential in anti-counterfeiting technology and advanced optical material design.

  5. DNA barcodes and citizen science provoke a diversity reappraisal for the "ring" butterflies of Peninsular Malaysia (Ypthima: Satyrinae: Nymphalidae: Lepidoptera).

    PubMed

    Jisming-See, Shi-Wei; Sing, Kong-Wah; Wilson, John-James

    2016-10-01

    The "rings" belonging to the genus Ypthima are amongst the most common butterflies in Peninsular Malaysia. However, the species can be difficult to tell apart, with keys relying on minor and often non-discrete ring characters found on the hindwing. Seven species have been reported from Peninsular Malaysia, but this is thought to be an underestimate of diversity. DNA barcodes of 165 individuals, and wing and genital morphology, were examined to reappraise species diversity of this genus in Peninsular Malaysia. DNA barcodes collected during citizen science projects-School Butterfly Project and Peninsular Malaysia Butterfly Count-recently conducted in Peninsular Malaysia were included. The new DNA barcodes formed six groups with different Barcode Index Numbers (BINs) representing four species reported in Peninsular Malaysia. When combined with public DNA barcodes from the Barcode Of Life Datasystems, several taxonomic issues arose. We consider the taxon Y. newboldi, formerly treated as a subspecies of Y. baldus, as a distinct species. DNA barcodes also supported an earlier suggestion that Y. nebulosa is a synonym under Y. horsfieldii humei. Two BINs of the genus Ypthima comprising DNA barcodes collected during citizen science projects did not correspond to any species previously reported in Peninsular Malaysia.

  6. Long-distance autumn migration across the Sahara by painted lady butterflies: exploiting resource pulses in the tropical savannah.

    PubMed

    Stefanescu, Constantí; Soto, David X; Talavera, Gerard; Vila, Roger; Hobson, Keith A

    2016-10-01

    The painted lady, Vanessa cardui, is a migratory butterfly that performs an annual multi-generational migration between Europe and North Africa. Its seasonal appearance south of the Sahara in autumn is well known and has led to the suggestion that it results from extremely long migratory flights by European butterflies to seasonally exploit the Sahel and the tropical savannah. However, this possibility has remained unproven. Here, we analyse the isotopic composition of butterflies from seven European and seven African countries to provide new support for this hypothesis. Each individual was assigned a geographical natal origin, based on its wing stable hydrogen isotope (δ 2 H w ) value and a predicted δ 2 H w basemap for Europe and northern Africa. Natal assignments of autumn migrants collected south of the Sahara confirmed long-distance movements (of 4000 km or more) starting in Europe. Samples from Maghreb revealed a mixed origin of migrants, with most individuals with a European origin, but others having originated in the Sahel. Therefore, autumn movements are not only directed to northwestern Africa, but also include southward and northward flights across the Sahara. Through this remarkable behaviour, the productive but highly seasonal region south of the Sahara is incorporated into the migratory circuit of V. cardui. © 2016 The Author(s).

  7. Structural characterization of Papilio kotzebuea (Eschscholtz 1821) butterfly wings

    NASA Astrophysics Data System (ADS)

    Sackey, J.; Nuru, Z. Y.; Berthier, S.; Maaza, M.

    2018-05-01

    The `plain black' forewings and black with `red spot' hindwings of the Papilio kotzebuea (Eschscholtz, 1821) were characterized by Scanning Electron Microscopy (SEM), Energy-Dispersive x-ray Spectroscopy (EDS), Atomic Force Microscopy (AFM), Fourier transform Infrared spectroscopy (FT-IR), UV-Vis spectrophometer and NIRQuest spectrometer. SEM images showed that the two sections of wings have different structures. The black with `red spot' hindwings have `hair-like' structures attached to the ridges and connected to the lamellae. On the contrary, the `plain black' forewings have holes that separate the ridges. AFM analysis unveiled that the `plain black' forewings have higher average surfaces roughness values as compared with the black with `red spot' hindwing. EDS and FT-IR results confirmed the presence of naturally hydrophobic materials on the wings. The `plain black' forewing exhibited strong absorptance (97%) throughout the solar spectrum range, which is attributed to the high melanin concentration as well as to the presence of holes in the scales. Biomimicking this wing could serves as equivalent solar absorber material.

  8. A dual layer hair array of the brown lacewing: repelling water at different length scales.

    PubMed

    Watson, Jolanta A; Cribb, Bronwen W; Hu, Hsuan-Ming; Watson, Gregory S

    2011-02-16

    Additional weight due to contamination (water and/or contaminating particles) can potentially have a detrimental effect on the flight capabilities of large winged insects such as butterflies and dragonflies. Insects where the wing surface area-body mass ratio is very high will be even more susceptible to these effects. Water droplets tend to move spontaneously off the wing surface of these insects. In the case of the brown lacewing, the drops effectively encounter a dual bed of hair springs with a topographical structure which aids in the hairs resisting penetration into water bodies. In this article, we demonstrate experimentally how this protective defense system employed by the brown lacewing (Micromus tasmaniae) aids in resisting contamination from water and how the micro- and nanostructures found on these hairs are responsible for quickly shedding water from the wing which demonstrates an active liquid-repelling surface. Copyright © 2011 Biophysical Society. Published by Elsevier Inc. All rights reserved.

  9. The Return of the Blue Butterfly

    NASA Astrophysics Data System (ADS)

    Santos, Anabela

    2014-05-01

    The Return of the Blue Butterfly The English writer Charles Dickens once wrote: "I only ask to be free. The butterflies are free". But are they really? The work that I performed with a group of students from 8th grade, had a starting point of climate change and the implications it has on ecosystems. Joining the passion I have for butterflies, I realized that they are also in danger of extinction due to these climatic effects. Thus, it was easy to seduce my students wanting to know more. Luckily I found Dr. Paula Seixas Arnaldo, a researcher at the University of Trás-os-Montes and Alto Douro, who has worked on butterflies and precisely investigated this issue. Portugal is the southern limit of butterfly-blue (Phengaris alcon), and has been many years in the red book of endangered species. Butterfly-blue is very demanding of their habitat, and disappears very easily if ideal conditions are not satisfied. Increased fragmentation of landscapes and degradation of suitable habitats, are considered the greatest challenges of the conservation of Phengaris butterfly in Portugal. In recent decades, climate change has also changed butterfly-blue spatial distribution with a movement of the species northward to colder locations, and dispersion in latitude. Butterflies of Europe must escape to the North because of the heat. Dr. Paula Seixas Arnaldo and her research team began a project, completed in December 2013, wanted to preserve and restore priority habitats recognized by the European Union to help species in danger of disappearing with increasing temperature. The blue butterfly is extremely important because it is a key indicator of the quality of these habitats. In the field, the butterflies are monitored to collect all possible data in order to identify the key species. Butterflies start flying in early July and cease in late August. Mating takes about an hour and occurs in the first days of life. The gentian-peat (Gentiana pneumonanthe) serves as the host plant for laying eggs. Each female lays an average of 60 eggs. Larva must grow in a plant near an anthill of Myrmica aloba species. This is important because butterfly larvae are myrmecophilous, living with ants that feed the butterfly larvae for 11 months, because the ants think the butterfly larvae are ant larvae. In early summer the larvae pupate in the nest of ants. Before expanding their wings, they have to leave quickly to avoid being killed by ants when the ants discover have been deceived. My students became aware of this research; we studied and prepared in order to carry out fieldwork. Thus students learn the content and curricular in a scientifically fun way, first with group work in the classroom with my guidance and in a second stage carry knowledge to the field under the guidance of Dra Paula Seixas Arnaldo. We know where we started ... where we arrives is success!

  10. Lattice Boltzmann simulations of flapping wings: The flock effect and the lateral wind effect

    NASA Astrophysics Data System (ADS)

    de Rosis, Alessandro

    2014-02-01

    In this paper, numerical analysis aiming at simulating biological organisms immersed in a fluid are carried out. The fluid domain is modeled through the lattice Boltzmann (LB) method, while the immersed boundary method is used to account for the position of the organisms idealized as rigid bodies. The time discontinuous Galerkin method is employed to compute body motion. An explicit coupling strategy to combine the adopted numerical methods is proposed. The vertical take-off of a couple of butterflies is numerically simulated in different scenarios, showing the mutual interaction that a butterfly exerts on the other one. Moreover, the effect of lateral wind is investigated. A critical threshold value of the lateral wind is defined, thus corresponding to an increasing arduous take-off.

  11. The enigmatic fast leaflet rotation in Desmodium motorium

    PubMed Central

    Lev-Yadun, Simcha

    2013-01-01

    I propose that the enigmatic leaflet movements in elliptical circles every few minutes of the Indian telegraph (semaphore) plant Desmodium motorium ( = D. gyrans = Hedysarum gyrans = Codariocalyx motorius), which has intrigued scientists for centuries, is a new type of butterfly or general winged arthropod mimicry by this plant. Such leaflet movement may deceive a passing butterfly searching for an un-occupied site suitable to deposit its eggs, that the plant is already occupied. It may also attract insectivorous birds, reptiles or arthropods to the plant because it looks as if it is harboring a potential prey and while they patrol there, they can find insects or other invertebrates that indeed attack the plant. The possibility that diurnal mammalian herbivores may also be deterred by these movements should not be dismissed. PMID:23603964

  12. Species delimitation in the Grayling genus Pseudochazara (Lepidoptera, Nymphalidae, Satyrinae) supported by DNA barcodes

    PubMed Central

    Verovnik, Rudi; Wiemers, Martin

    2016-01-01

    Abstract The Palaearctic Grayling genus Pseudochazara encompasses a number of petrophilous butterfly species, most of which are local endemics especially in their centre of radiation in SW Asia and the Balkans. Due to a lack of consistent morphological characters, coupled with habitat induced variability, their taxonomy is poorly understood and species delimitation is hampered. We employed a DNA barcoding approach to address the question of separate species status for several European taxa and provide first insight into the phylogeny of the genus. Unexpectedly we found conflicting patterns with deep divergences between presumably conspecific taxa and lack of divergence among well-defined species. We propose separate species status for Pseudochazara tisiphone, Pseudochazara amalthea, Pseudochazara amymone, and Pseudochazara kermana all of which have separate well supported clades, with the majority of them becoming local endemics. Lack of resolution in the ‘Mamurra’ species group with well-defined species (in terms of wing pattern and coloration) such as Pseudochazara geyeri, Pseudochazara daghestana and Pseudochazara alpina should be further explored using nuclear molecular markers with higher genetic resolution. PMID:27408604

  13. Cultural Nuances, Assumptions, and the Butterfly Effect: Addressing the Unpredictability Caused by Unconscious Values Structures in Cross-Cultural Interactions

    ERIC Educational Resources Information Center

    Remer, Rory

    2007-01-01

    Cultural values, cross-cultural interaction patterns that are produced by dynamical (chaotic) systems, have a significant impact on interaction, particularly among and between people from different cultures. The butterfly effect, which states that small differences in initial conditions may have severe consequences for patterns in the long run,…

  14. A fossil biting midge (Diptera: Ceratopogonidae) from early Eocene Indian amber with a complex pheromone evaporator

    NASA Astrophysics Data System (ADS)

    Stebner, Frauke; Szadziewski, Ryszard; Rühr, Peter T.; Singh, Hukam; Hammel, Jörg U.; Kvifte, Gunnar Mikalsen; Rust, Jes

    2016-10-01

    The life-like fidelity of organisms captured in amber is unique among all kinds of fossilization and represents an invaluable source for different fields of palaeontological and biological research. One of the most challenging aspects in amber research is the study of traits related to behaviour. Here, indirect evidence for pheromone-mediated mating behaviour is recorded from a biting midge (Ceratopogonidae) in 54 million-year-old Indian amber. Camptopterohelea odora n. sp. exhibits a complex, pocket shaped structure on the wings, which resembles the wing folds of certain moth flies (Diptera: Psychodidae) and scent organs that are only known from butterflies and moths (Lepidoptera) so far. Our studies suggests that pheromone releasing structures on the wings have evolved independently in biting midges and might be much more widespread in fossil as well as modern insects than known so far.

  15. A fossil biting midge (Diptera: Ceratopogonidae) from early Eocene Indian amber with a complex pheromone evaporator.

    PubMed

    Stebner, Frauke; Szadziewski, Ryszard; Rühr, Peter T; Singh, Hukam; Hammel, Jörg U; Kvifte, Gunnar Mikalsen; Rust, Jes

    2016-10-04

    The life-like fidelity of organisms captured in amber is unique among all kinds of fossilization and represents an invaluable source for different fields of palaeontological and biological research. One of the most challenging aspects in amber research is the study of traits related to behaviour. Here, indirect evidence for pheromone-mediated mating behaviour is recorded from a biting midge (Ceratopogonidae) in 54 million-year-old Indian amber. Camptopterohelea odora n. sp. exhibits a complex, pocket shaped structure on the wings, which resembles the wing folds of certain moth flies (Diptera: Psychodidae) and scent organs that are only known from butterflies and moths (Lepidoptera) so far. Our studies suggests that pheromone releasing structures on the wings have evolved independently in biting midges and might be much more widespread in fossil as well as modern insects than known so far.

  16. A fossil biting midge (Diptera: Ceratopogonidae) from early Eocene Indian amber with a complex pheromone evaporator

    PubMed Central

    Stebner, Frauke; Szadziewski, Ryszard; Rühr, Peter T.; Singh, Hukam; Hammel, Jörg U.; Kvifte, Gunnar Mikalsen; Rust, Jes

    2016-01-01

    The life-like fidelity of organisms captured in amber is unique among all kinds of fossilization and represents an invaluable source for different fields of palaeontological and biological research. One of the most challenging aspects in amber research is the study of traits related to behaviour. Here, indirect evidence for pheromone-mediated mating behaviour is recorded from a biting midge (Ceratopogonidae) in 54 million-year-old Indian amber. Camptopterohelea odora n. sp. exhibits a complex, pocket shaped structure on the wings, which resembles the wing folds of certain moth flies (Diptera: Psychodidae) and scent organs that are only known from butterflies and moths (Lepidoptera) so far. Our studies suggests that pheromone releasing structures on the wings have evolved independently in biting midges and might be much more widespread in fossil as well as modern insects than known so far. PMID:27698490

  17. Comparing the response of birds and butterflies to vegetation-based mountain ecotones using boundary detection approaches.

    PubMed

    Kent, Rafi; Levanoni, Oded; Banker, Eran; Pe'er, Guy; Kark, Salit

    2013-01-01

    Mountains provide an opportunity to examine changes in biodiversity across environmental gradients and areas of transition (ecotones). Mountain ecotones separate vegetation belts. Here, we aimed to examine whether transition areas for birds and butterflies spatially correspond with ecotones between three previously described altitudinal vegetation belts on Mt. Hermon, northern Israel. These include the Mediterranean Maquis, xero-montane open forest and Tragacanthic mountain steppe vegetation belts. We sampled the abundance of bird and butterfly species in 34 sampling locations along an elevational gradient between 500 and 2200 m. We applied wombling, a boundary-detection technique, which detects rapid changes in a continuous variable, in order to locate the transition areas for bird and butterfly communities and compare the location of these areas with the location of vegetation belts as described in earlier studies of Mt. Hermon. We found some correspondence between the areas of transition of both bird and butterfly communities and the ecotones between vegetation belts. For birds and butterflies, important transitions occurred at the lower vegetation ecotone between Mediterranean maquis and the xero-montane open forest vegetation belts, and between the xero-montane open forest and the mountain steppe Tragacanthic belts. While patterns of species turnover with elevation were similar for birds and butterflies, the change in species richness and diversity with elevation differed substantially between the two taxa. Birds and butterflies responded quite similarly to the elevational gradient and to the shift between vegetation belts in terms of species turnover rates. While the mechanisms generating these patterns may differ, the resulting areas of peak turnover in species show correspondence among three different taxa (plants, birds and butterflies).

  18. Evolutionary history of the recruitment of conserved developmental genes in association to the formation and diversification of a novel trait.

    PubMed

    Shirai, Leila T; Saenko, Suzanne V; Keller, Roberto A; Jerónimo, Maria A; Brakefield, Paul M; Descimon, Henri; Wahlberg, Niklas; Beldade, Patrícia

    2012-02-15

    The origin and modification of novel traits are important aspects of biological diversification. Studies combining concepts and approaches of developmental genetics and evolutionary biology have uncovered many examples of the recruitment, or co-option, of genes conserved across lineages for the formation of novel, lineage-restricted traits. However, little is known about the evolutionary history of the recruitment of those genes, and of the relationship between them -for example, whether the co-option involves whole or parts of existing networks, or whether it occurs by redeployment of individual genes with de novo rewiring. We use a model novel trait, color pattern elements on butterfly wings called eyespots, to explore these questions. Eyespots have greatly diversified under natural and sexual selection, and their formation involves genetic circuitries shared across insects. We investigated the evolutionary history of the recruitment and co-recruitment of four conserved transcription regulators to the larval wing disc region where circular pattern elements develop. The co-localization of Antennapedia, Notch, Distal-less, and Spalt with presumptive (eye)spot organizers was examined in 13 butterfly species, providing the largest comparative dataset available for the system. We found variation between families, between subfamilies, and between tribes. Phylogenetic reconstructions by parsimony and maximum likelihood methods revealed an unambiguous evolutionary history only for Antennapedia, with a resolved single origin of eyespot-associated expression, and many homoplastic events for Notch, Distal-less, and Spalt. The flexibility in the (co-)recruitment of the targeted genes includes cases where different gene combinations are associated with morphologically similar eyespots, as well as cases where identical protein combinations are associated with very different phenotypes. The evolutionary history of gene (co-)recruitment is consistent with both divergence from a recruited putative ancestral network, and with independent co-option of individual genes. The diversity in the combinations of genes expressed in association with eyespot formation does not parallel diversity in characteristics of the adult phenotype. We discuss these results in the context of inferring homology. Our study underscores the importance of widening the representation of phylogenetic, morphological, and genetic diversity in order to establish general principles about the mechanisms behind the evolution of novel traits.

  19. The wing pattern of Moerarchis Durrant, 1914 (Lepidoptera: Tineidae) clarifies transitions between predictive models

    PubMed Central

    2017-01-01

    The evolution of wing pattern in Lepidoptera is a popular area of inquiry but few studies have examined microlepidoptera, with fewer still focusing on intraspecific variation. The tineid genus Moerarchis Durrant, 1914 includes two species with high intraspecific variation of wing pattern. A subset of the specimens examined here provide, to my knowledge, the first examples of wing patterns that follow both the ‘alternating wing-margin’ and ‘uniform wing-margin’ models in different regions along the costa. These models can also be evaluated along the dorsum of Moerarchis, where a similar transition between the two models can be seen. Fusion of veins is shown not to effect wing pattern, in agreement with previous inferences that the plesiomorphic location of wing veins constrains the development of colour pattern. The significant correlation between wing length and number of wing pattern elements in Moerarchis australasiella shows that wing size can act as a major determinant of wing pattern complexity. Lastly, some M. australasiella specimens have wing patterns that conform entirely to the ‘uniform wing-margin’ model and contain more than six bands, providing new empirical insight into the century-old question of how wing venation constrains wing patterns with seven or more bands. PMID:28405390

  20. Wing scale ultrastructure underlying convergent and divergent iridescent colours in mimetic Heliconius butterflies.

    PubMed

    Parnell, Andrew J; Bradford, James E; Curran, Emma V; Washington, Adam L; Adams, Gracie; Brien, Melanie N; Burg, Stephanie L; Morochz, Carlos; Fairclough, J Patrick A; Vukusic, Pete; Martin, Simon J; Doak, Scott; Nadeau, Nicola J

    2018-04-01

    Iridescence is an optical phenomenon whereby colour changes with the illumination and viewing angle. It can be produced by thin film interference or diffraction. Iridescent optical structures are fairly common in nature, but relatively little is known about their production or evolution. Here we describe the structures responsible for producing blue-green iridescent colour in Heliconius butterflies. Overall the wing scale structures of iridescent and non-iridescent Heliconius species are very similar, both having longitudinal ridges joined by cross-ribs. However, iridescent scales have ridges composed of layered lamellae, which act as multilayer reflectors. Differences in brightness between species can be explained by the extent of overlap of the lamellae and their curvature as well as the density of ridges on the scale. Heliconius are well known for their Müllerian mimicry. We find that iridescent structural colour is not closely matched between co-mimetic species. Differences appear less pronounced in models of Heliconius vision than models of avian vision, suggesting that they are not driven by selection to avoid heterospecific courtship by co-mimics. Ridge profiles appear to evolve relatively slowly, being similar between closely related taxa, while ridge density evolves faster and is similar between distantly related co-mimics. © 2018 The Authors.

  1. Wing scale ultrastructure underlying convergent and divergent iridescent colours in mimetic Heliconius butterflies

    PubMed Central

    Bradford, James E.; Curran, Emma V.; Washington, Adam L.; Adams, Gracie; Brien, Melanie N.; Burg, Stephanie L.; Morochz, Carlos; Fairclough, J. Patrick A.; Vukusic, Pete; Martin, Simon J.; Doak, Scott

    2018-01-01

    Iridescence is an optical phenomenon whereby colour changes with the illumination and viewing angle. It can be produced by thin film interference or diffraction. Iridescent optical structures are fairly common in nature, but relatively little is known about their production or evolution. Here we describe the structures responsible for producing blue-green iridescent colour in Heliconius butterflies. Overall the wing scale structures of iridescent and non-iridescent Heliconius species are very similar, both having longitudinal ridges joined by cross-ribs. However, iridescent scales have ridges composed of layered lamellae, which act as multilayer reflectors. Differences in brightness between species can be explained by the extent of overlap of the lamellae and their curvature as well as the density of ridges on the scale. Heliconius are well known for their Müllerian mimicry. We find that iridescent structural colour is not closely matched between co-mimetic species. Differences appear less pronounced in models of Heliconius vision than models of avian vision, suggesting that they are not driven by selection to avoid heterospecific courtship by co-mimics. Ridge profiles appear to evolve relatively slowly, being similar between closely related taxa, while ridge density evolves faster and is similar between distantly related co-mimics. PMID:29669892

  2. Bioinspired fabrication of hierarchically structured, pH-tunable photonic crystals with unique transition.

    PubMed

    Yang, Qingqing; Zhu, Shenmin; Peng, Wenhong; Yin, Chao; Wang, Wanlin; Gu, Jiajun; Zhang, Wang; Ma, Jun; Deng, Tao; Feng, Chuanliang; Zhang, Di

    2013-06-25

    We herein report a new class of photonic crystals with hierarchical structures, which are of color tunability over pH. The materials were fabricated through the deposition of polymethylacrylic acid (PMAA) onto a Morpho butterfly wing template by using a surface bonding and polymerization route. The amine groups of chitosan in Morpho butterfly wings provide reaction sites for the MAA monomer, resulting in hydrogen bonding between the template and MAA. Subsequent polymerization results in PMAA layers coating homogenously on the hierarchical photonic structures of the biotemplate. The pH-induced color change was detected by reflectance spectra as well as optical observation. A distinct U transition with pH was observed, demonstrating PMAA content-dependent properties. The appearance of the unique U transition results from electrostatic interaction between the -NH3(+) of chitosan and the -COO(-) groups of PMAA formed, leading to a special blue-shifted point at the pH value of the U transition, and the ionization of the two functional groups in the alkali and acid environment separately, resulting in a red shift. This work sets up a strategy for the design and fabrication of tunable photonic crystals with hierarchical structures, which provides a route for combining functional polymers with biotemplates for wide potential use in many fields.

  3. Cool habitats support darker and bigger butterflies in Australian tropical forests.

    PubMed

    Xing, Shuang; Bonebrake, Timothy C; Tang, Chin Cheung; Pickett, Evan J; Cheng, Wenda; Greenspan, Sasha E; Williams, Stephen E; Scheffers, Brett R

    2016-11-01

    Morphology mediates the relationship between an organism's body temperature and its environment. Dark organisms, for example, tend to absorb heat more quickly than lighter individuals, which could influence their responses to temperature. Therefore, temperature-related traits such as morphology may affect patterns of species abundance, richness, and community assembly across a broad range of spatial scales. In this study, we examined variation in color lightness and body size within butterfly communities across hot and cool habitats in the tropical woodland-rainforest ecosystems of northeast Queensland, Australia. Using thermal imaging, we documented the absorption of solar radiation relative to color lightness and wingspan and then built a phylogenetic tree based on available sequences to analyze the effects of habitat on these traits within a phylogenetic framework. In general, darker and larger individuals were more prevalent in cool, closed-canopy rainforests than in immediately adjacent and hotter open woodlands. In addition, darker and larger butterflies preferred to be active in the shade and during crepuscular hours, while lighter and smaller butterflies were more active in the sun and midday hours-a pattern that held after correcting for phylogeny. Our ex situ experiment supported field observations that dark and large butterflies heated up faster than light and small butterflies under standardized environmental conditions. Our results show a thermal consequence of butterfly morphology across habitats and how environmental factors at a microhabitat scale may affect the distribution of species based on these traits. Furthermore, this study highlights how butterfly species might differentially respond to warming based on ecophysiological traits and how thermal refuges might emerge at microclimatic and habitat scales.

  4. Extensive range overlap between heliconiine sister species: evidence for sympatric speciation in butterflies?

    PubMed

    Rosser, Neil; Kozak, Krzysztof M; Phillimore, Albert B; Mallet, James

    2015-06-30

    Sympatric speciation is today generally viewed as plausible, and some well-supported examples exist, but its relative contribution to biodiversity remains to be established. We here quantify geographic overlap of sister species of heliconiine butterflies, and use age-range correlations and spatial simulations of the geography of speciation to infer the frequency of sympatric speciation. We also test whether shifts in mimetic wing colour pattern, host plant use and climate niche play a role in speciation, and whether such shifts are associated with sympatry. Approximately a third of all heliconiine sister species pairs exhibit near complete range overlap, and analyses of the observed patterns of range overlap suggest that sympatric speciation contributes 32%-95% of speciation events. Müllerian mimicry colour patterns and host plant choice are highly labile traits that seem to be associated with speciation, but we find no association between shifts in these traits and range overlap. In contrast, climatic niches of sister species are more conserved. Unlike birds and mammals, sister species of heliconiines are often sympatric and our inferences using the most recent comparative methods suggest that sympatric speciation is common. However, if sister species spread rapidly into sympatry (e.g. due to their similar climatic niches), then assumptions underlying our methods would be violated. Furthermore, although we find some evidence for the role of ecology in speciation, ecological shifts did not show the associations with range overlap expected under sympatric speciation. We delimit species of heliconiines in three different ways, based on "strict and " "relaxed" biological species concepts (BSC), as well as on a surrogate for the widely-used "diagnostic" version of the phylogenetic species concept (PSC). We show that one reason why more sympatric speciation is inferred in heliconiines than in birds may be due to a different culture of species delimitation in the two groups. To establish whether heliconiines are exceptional will require biogeographic comparative studies for a wider range of animal taxa including many more invertebrates.

  5. Peptide toxin glacontryphan-M is present in the wings of the butterfly Hebomoia glaucippe (Linnaeus, 1758) (Lepidoptera: Pieridae).

    PubMed

    Bae, Narkhyun; Li, Lin; Lödl, Martin; Lubec, Gert

    2012-10-30

    Protein profiling has revealed the presence of glacontryphan-M, a peptide toxin identified only in the sea snail genus Conus, in the wings of Hebomoia glaucippe (HG). The wings and body of HG were homogenized and the proteins were extracted and analyzed by 2D gel electrophoresis with subsequent in-gel digestion. Posttranslational protein modifications were detected and analyzed by nano-LC-MS/MS. An antibody was generated against glacontryphan-M, and protein extracts from the wings of HG samples from Malaysia, Indonesia, and the Philippines were tested by immunoblotting. Glacontryphan-M was unambiguously identified in the wings of HG containing the following posttranslational protein modifications: monoglutamylation at E55, methylation at E53, quinone modification at W61, cyanylation at C56, and amidation of the C terminus at G63. Immunoblotting revealed the presence of the toxin in the wings of HG from all origins, showing a single band for glacontryphan-M in HG samples from Malaysia and Philippines and a double band in HG samples from Indonesia. Intriguingly, sequence analysis indicated that the Conus glacontryphan is identical to that of HG. The toxin may function as a defense against diverse predators, including ants, mantes, spiders, lizards, green frogs, and birds.

  6. Fluid Dynamics of Underwater Flight in Sea Butterflies: Insights from Computational Modeling

    NASA Astrophysics Data System (ADS)

    Zhou, Zhuoyu; Mittal, Rajat; Yen, Jeannette; Webster, Donald

    2014-11-01

    Sea butterflies such as Limacine helicina swim by flapping their wing-like parapodia, in a stroke that exhibits a clap-and-fling type kinematics as well as a strong interaction between the parapodia and the body of the animal at the end of downstroke. We used numerical simulations based on videogrammetric data to examine the fluid dynamics and force generation associated with this swimming motion. The unsteady lift-generating mechanism of clap-and-fling results in a sawtooth trajectory with a characteristic ``wobble'' in pitch. We employ coupled flow-body-dynamics simulations to model the free-swimming motion of the organism and explore the efficiency of propulsion as well the factors such as shell weight, that affect its sawtooth swimming trajectory. This work is funded by NSF Grant 1246317 from the Division of Polar Programs.

  7. Macromoths of northwest forests and woodlands

    USGS Publications Warehouse

    Miller, Jeffrey C.; Hammond, Paul C.

    2000-01-01

    The macromoths are a group of families within the order Lepidoptera. The macromoths in the woodlands and forests of the Pacific Northwest are represented by 1,200 species in 12 families: Arctiidae, Dioptidae, Drepanidae, Epiplemidae, Geometridae, Lasiocampidae, Lymantriidae, Noctuidae, Notodontidae, Saturniidae, Sphingidae, and Thyatiridae. In addition to the macromoths, the Lepidoptera are represented by the butterflies and skippers, and the micromoths. Butterflies possess a knob at the tip of the antennae while the tip of the antennae in skippers is typically hooked. The tip of the antennae in macromoths and micromoths is tapered. The differences between macromoths and micromoths is not literally based on size as the names suggest but rather in details of the female reproductive tract and wing venation. These details are discussed and illustrated in most texts on general entomology (Borror et al. 1989) and in books about Lepidoptera (Covell 1984).

  8. Movement patterns and population characteristics of the Karner blue butterfly (Lycaeides melissa samuelis) at Indiana Dunes National Lakeshore

    USGS Publications Warehouse

    Knutson, Randy L.; Kwilosz, John R.; Grundel, Ralph

    1999-01-01

    We conducted a three-year mark-release-recapture study of the endangered Karner blue butterfly (Lycaeides melissa samuelis Nabokov) at Indiana Dunes National Lakeshore to describe the butterfly's movement patterns and to assess seasonal changes in the Karner blue's population structure. Estimated mean Karner blue adult life span was less than 3.5 days. Populations exhibited protandry and about a 2:1 male:female sex ratio at population peak within a brood. Ranges, or maximum distances moved by individual butterflies, were typically less than 100 m. Maximum ranges were less than 1 km. These distances are similar to those reported for other lycaenid butterflies and from other studies of the Karner blue in the midwestern United States. At two sites, fewer than 2% of adults had ranges greater than 300 m, while at a third site 4.3% of adults had ranges greater than 300 m. Given typical subpopulation sizes these movement percentages suggest that few adults per generation will move between subpopulations separated by more than 300 m. Movement of individuals between subpopulation sites is important for maintaining genetic diversity within a metapopulation and for recolonizing areas following local extinctions. Therefore, prudent conservation planning should aim for a landscape with habitat patches suitable for Karner blue butterfly occupancy separated by less than 300 m.

  9. Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea).

    PubMed

    Jong, Rienk DE

    2017-05-25

    Fossil butterflies are extremely rare. Yet, they are the only direct evidence of the first appearance of particular characters and as such, they are crucial for calibrating a molecular clock, from which divergence ages are estimated. In turn, these estimates, in combination with paleogeographic information, are most important in paleobiogeographic considerations. The key issue here is the correct allocation of fossils on the phylogenetic tree from which the molecular clock is calibrated.The allocation of a fossil on a tree should be based on an apomorphic character found in a tree based on extant species, similar to the allocation of a new extant species. In practice, the latter is not done, at least not explicitly, on the basis of apomorphy, but rather on overall similarity or on a phylogenetic analysis, which is not possible for most butterfly fossils since they usually are very fragmentary. Characters most often preserved are in the venation of the wings. Therefore, special attention is given to possible apomorphies in venational characters in extant butterflies. For estimation of divergence times, not only the correct allocation of the fossil on the tree is important, but also the tree itself influences the outcome as well as the correct determination of the age of the fossil. These three aspects are discussed.        All known butterfly fossils, consisting of 49 taxa, are critically reviewed and their relationship to extant taxa is discussed as an aid for correctly calibrating a molecular clock for papilionoid Lepidoptera. In this context some aspects of age estimation and biogeographic conclusions are briefly mentioned in review. Specific information has been summarized in four appendices.

  10. Reflectance of topologically disordered photonic-crystal films

    NASA Astrophysics Data System (ADS)

    Vigneron, Jean-Pol; Lousse, Virginie M.; Biro, Laszlo P.; Vertesy, Zofia; Balint, Zolt

    2005-04-01

    Periodicity implies the creation of discretely diffracted beams while various departures from periodicity lead to broadened scattering angles. This effect is investigated for disturbed lattices exhibiting randomly varying periods. In the Born approximation, the diffused reflection is shown to be related to a pair correlation function constructed from the distribution of the film scattering power. The technique is first applied to a natural photonic crystal found on the ventral side of the wings of the butterfly Cyanophrys remus, where scanning electron microscopy reveals the formation of polycrystalline photonic structures. Second, the disorder in the distribution of the cross-ribs on the scales another butterfly, Lycaena virgaureae, is investigated. The irregular arrangement of scatterers found in chitin structure of this insect produces light reflection in the long-wavelength part of the visible range, with a quite unusual broad directionality. The use of the pair correlation function allows to propose estimates of the diffusive spreading in these very different systems.

  11. Various Types of Coexisting Attractors in a New 4D Autonomous Chaotic System

    NASA Astrophysics Data System (ADS)

    Lai, Qiang; Akgul, Akif; Zhao, Xiao-Wen; Pei, Huiqin

    An unique 4D autonomous chaotic system with signum function term is proposed in this paper. The system has four unstable equilibria and various types of coexisting attractors appear. Four-wing and four-scroll strange attractors are observed in the system and they will be broken into two coexisting butterfly attractors and two coexisting double-scroll attractors with the variation of the parameters. Numerical simulation shows that the system has various types of multiple coexisting attractors including two butterfly attractors with four limit cycles, two double-scroll attractors with a limit cycle, four single-scroll strange attractors, four limit cycles with regard to different parameters and initial values. The coexistence of the attractors is determined by the bifurcation diagrams. The chaotic and hyperchaotic properties of the attractors are verified by the Lyapunov exponents. Moreover, we present an electronic circuit to experimentally realize the dynamic behavior of the system.

  12. Structural Color Model Based on Surface Morphology of MORPHO Butterfly Wing Scale

    NASA Astrophysics Data System (ADS)

    Huang, Zhongjia; Cai, Congcong; Wang, Gang; Zhang, Hui; Huttula, Marko; Cao, Wei

    2016-05-01

    Color production through structural coloration is created by micrometer and sub-micrometer surface textures which interfere with visible light. The shiny blue of morpho menelaus is a typical example of structural coloring. Modified from morphology of the morpho scale, a structure of regular windows with two side offsets was constructed on glass substrates. Optical properties of the bioinspired structure were studied through numerical simulations of light scattering. Results show that the structure can generate monochromatic light scattering. Wavelength of scattered light is tunable via changing the spacing between window shelves. Compared to original butterfly model, the modified one possesses larger illumination scopes in azimuthal distributions despite being less in polar directions. Present bionic structure is periodically repeated and is easy to fabricate. It is hoped that the computational materials design work can inspire future experimental realizations of such a structure in photonics applications.

  13. Changes in butterfly abundance in response to global warming and reforestation.

    PubMed

    Kwon, Tae-Sung; Kim, Sung-Soo; Chun, Jung Hwa; Byun, Bong-Kyu; Lim, Jong-Hwan; Shin, Joon Hwan

    2010-04-01

    In the Republic of Korea, most denuded forest lands have been restored since the 1960s. In addition, the annual mean temperature in the Republic of Korea has increased approximately 1.0 degrees C during the last century, which is higher than the global mean increase of 0.74 degrees C. Such rapid environmental changes may have resulted in changes in the local butterfly fauna. For example, the number of butterflies inhabiting forests may have increased because of reforestation, whereas the number of butterflies inhabiting grasslands may have declined. Furthermore, the number of northern butterflies may have declined, whereas the number of southern butterflies may have increased in response to global warming. Therefore, we compared current data (2002 approximately 2007) regarding the abundance of butterfly species at two sites in the central portion of the Korean Peninsula to data from the late 1950s and early 1970s for the same sites. Changes in the abundance rank of each species between the two periods were evaluated to determine whether any patterns corresponded to the predicted temporal changes. The predicted changes in butterfly abundance were confirmed in this study. In addition, the results showed a different response to habitat change between northern and southern species. In northern butterfly species, butterflies inhabiting forests increased, whereas those inhabiting grasslands declined. However, the opposite was true when southern butterfly species were evaluated. Changes in the abundance indicate that habitat change may be one of the key factors related to the survival of populations that remain around the southern boundary of butterfly species.

  14. Dual Imprinted Polymer Thin Films via Pattern Directed Self-Organization.

    PubMed

    Grolman, Danielle; Bandyopadhyay, Diya; Al-Enizi, Abdullah; Elzatahry, Ahmed; Karim, Alamgir

    2017-06-21

    Synthetic topographically patterned films and coatings are typically contoured on one side, yet many of nature's surfaces have distinct textures on different surfaces of the same object. Common examples are the top and bottom sides of the butterfly wing or lotus leaf, onion shells, and the inside versus outside of the stem of a flower. Inspired by nature, we create dual (top and bottom) channel patterned polymer films. To this end, we first develop a novel fabrication method to create ceramic line channel relief structures by converting the oligomeric residue of stamped poly(dimethylsiloxane) (PDMS) nanopatterns on silicon substrates to glass (SiOx, silica) by ultraviolet-ozone (UVO) exposure. These silica patterned substrates are flow coated with polystyrene (PS) films and confined within an identically patterned top confining soft PDMS elastomer film. Annealing of the sandwich structures drives the PS to rapidly mold fill the top PDMS pattern in conjunction with a dewetting tendency of the PS on the silica pattern. Varying the film thickness h, from less than to greater than the pattern height, and varying the relative angle between the top-down and bottom-up patterned confinement surfaces create interesting uniform and nonuniform digitized defects in PS channel patterns, as also a defect-free channel regime. Our dual patterned polymer channels provide a novel fabrication route to topographically imprinted Moiré patterns (whose applications range from security encrypting holograms to sensitive strain gauges), and their basic laser light diffractions properties are illustrated and compared to graphical simulations and 2D-FFT of real-space AFM channel patterns. While traditional "geometrical" and "fringe" Moiré patterns function by superposition of two misaligned optical patterned transmittance gratings, our topographic pattern gratings are quite distinct and may allow for more unique holographic optical characteristics with further development.

  15. Do Healthy Monarchs Migrate Farther? Tracking Natal Origins of Parasitized vs. Uninfected Monarch Butterflies Overwintering in Mexico

    PubMed Central

    Altizer, Sonia; Hobson, Keith A.; Davis, Andrew K.; De Roode, Jacobus C.; Wassenaar, Leonard I.

    2015-01-01

    Long-distance migration can lower parasite prevalence if strenuous journeys remove infected animals from wild populations. We examined wild monarch butterflies (Danaus plexippus) to investigate the potential costs of the protozoan Ophryocystis elektroscirrha on migratory success. We collected monarchs from two wintering sites in central Mexico to compare infection status with hydrogen isotope (δ 2H) measurements as an indicator of latitude of origin at the start of fall migration. On average, uninfected monarchs had lower δ 2H values than parasitized butterflies, indicating that uninfected butterflies originated from more northerly latitudes and travelled farther distances to reach Mexico. Within the infected class, monarchs with higher quantitative spore loads originated from more southerly latitudes, indicating that heavily infected monarchs originating from farther north are less likely to reach Mexico. We ruled out the alternative explanation that lower latitudes give rise to more infected monarchs prior to the onset of migration using citizen science data to examine regional differences in parasite prevalence during the summer breeding season. We also found a positive association between monarch wing area and estimated distance flown. Collectively, these results emphasize that seasonal migrations can help lower infection levels in wild animal populations. Our findings, combined with recent declines in the numbers of migratory monarchs wintering in Mexico and observations of sedentary (winter breeding) monarch populations in the southern U.S., suggest that shifts from migratory to sedentary behavior will likely lead to greater infection prevalence for North American monarchs. PMID:26606389

  16. The interaction of the halo around the butterfly planetary nebula NGC 650-1 with the interstellar medium

    NASA Astrophysics Data System (ADS)

    Ramos-Larios, G.; Guerrero, M. A.; Nigoche-Netro, A.; Olguín, L.; Gómez-Muñoz, M. A.; Sabin, L.; Vázquez, R.; Akras, S.; Ramírez Vélez, J. C.; Chávez, M.

    2018-03-01

    With its bright and wide equatorial waist seen almost edge-on (`the butterfly body') and the faint and broad bipolar extensions (`the butterfly wings'), NGC 650-1 is the archetypical example of bipolar planetary nebula (PN) with butterfly morphology. We present here deep high-resolution broad- and narrow-band optical images that expose the rich and intricate fine structure of this bipolar PN, with small-scale bubble-like features and collimated outflows. A SHAPE spatio-kinematic model indicates that NGC 650-1 has a broad central torus with an inclination angle of 75° with respect to the line of sight, whereas that of the bipolar lobes, which are clearly seen in the position-velocity maps, is 85°. Large field of view deep images show, for first time, an arc-like diffuse envelope in low- and high-excitation emission lines located up to 180 arcsec towards the east-south-east of the central star, well outside the main nebula. This morphological component is confirmed by Spitzer MIPS and WISE infrared imaging, as well as by long-slit low- and high-dispersion optical spectroscopic observations. Hubble Space Telescope images of NGC 650-1 obtained at two different epochs ˜14 yr apart reveal the proper motion of the central star along this direction. We propose that this motion of the star through the interstellar medium compresses the remnant material of a slow asymptotic giant branch wind, producing this bow-shock-like feature.

  17. Do Healthy Monarchs Migrate Farther? Tracking Natal Origins of Parasitized vs. Uninfected Monarch Butterflies Overwintering in Mexico.

    PubMed

    Altizer, Sonia; Hobson, Keith A; Davis, Andrew K; De Roode, Jacobus C; Wassenaar, Leonard I

    2015-01-01

    Long-distance migration can lower parasite prevalence if strenuous journeys remove infected animals from wild populations. We examined wild monarch butterflies (Danaus plexippus) to investigate the potential costs of the protozoan Ophryocystis elektroscirrha on migratory success. We collected monarchs from two wintering sites in central Mexico to compare infection status with hydrogen isotope (δ2H) measurements as an indicator of latitude of origin at the start of fall migration. On average, uninfected monarchs had lower δ2H values than parasitized butterflies, indicating that uninfected butterflies originated from more northerly latitudes and travelled farther distances to reach Mexico. Within the infected class, monarchs with higher quantitative spore loads originated from more southerly latitudes, indicating that heavily infected monarchs originating from farther north are less likely to reach Mexico. We ruled out the alternative explanation that lower latitudes give rise to more infected monarchs prior to the onset of migration using citizen science data to examine regional differences in parasite prevalence during the summer breeding season. We also found a positive association between monarch wing area and estimated distance flown. Collectively, these results emphasize that seasonal migrations can help lower infection levels in wild animal populations. Our findings, combined with recent declines in the numbers of migratory monarchs wintering in Mexico and observations of sedentary (winter breeding) monarch populations in the southern U.S., suggest that shifts from migratory to sedentary behavior will likely lead to greater infection prevalence for North American monarchs.

  18. Alignment of crystal orientations of the multi-domain photonic crystals in Parides sesostris wing scales

    PubMed Central

    Yoshioka, S.; Fujita, H.; Kinoshita, S.; Matsuhana, B.

    2014-01-01

    It is known that the wing scales of the emerald-patched cattleheart butterfly, Parides sesostris, contain gyroid-type photonic crystals, which produce a green structural colour. However, the photonic crystal is not a single crystal that spreads over the entire scale, but it is separated into many small domains with different crystal orientations. As a photonic crystal generally has band gaps at different frequencies depending on the direction of light propagation, it seems mysterious that the scale is observed to be uniformly green under an optical microscope despite the multi-domain structure. In this study, we have carefully investigated the structure of the wing scale and discovered that the crystal orientations of different domains are not perfectly random, but there is a preferred crystal orientation that is aligned along the surface normal of the scale. This finding suggests that there is an additional factor during the developmental process of the microstructure that regulates the crystal orientation. PMID:24352678

  19. Direct writing of micro/nano-scale patterns by means of particle lens arrays scanned by a focused diode pumped Nd:YVO4 laser

    NASA Astrophysics Data System (ADS)

    Pena, Ana; Wang, Zengbo; Whitehead, David; Li, Lin

    2010-11-01

    A practical approach to a well-known technique of laser micro/nano-patterning by optical near fields is presented. It is based on surface patterning by scanning a Gaussian laser beam through a self-assembled monolayer of silica micro-spheres on a single-crystalline silicon (Si) substrate. So far, the outcome of this kind of near-field patterning has been related to the simultaneous, parallel surface-structuring of large areas either by top hat or Gaussian laser intensity distributions. We attempt to explore the possibility of using the same technique in order to produce single, direct writing of features. This could be of advantage for applications in which only some areas need to be patterned (i.e. local area selective patterning) or single lines are required (e.g. a particular micro/nano-fluidic channel). A diode pumped Nd:YVO4 laser system (wavelength of 532 nm, pulse duration of 8 ns, repetition rate of 30 kHz) with a computer-controlled 3 axis galvanometer beam scanner was employed to write user-defined patterns through the particle lens array on the Si substrate. After laser irradiation, the obtained patterns which are in the micro-scale were composed of sub-micro/micro-holes or bumps. The micro-pattern resolution depends on the dimension of both the micro-sphere’s diameter and the beam’s spot size. The developed technique could potentially be employed to fabricate photonic crystal structures mimicking nature’s butterfly wings and anti-reflective “moth eye” arrays for photovoltaic cells.

  20. Warning color changes in response to food deprivation in the pipevine swallowtail butterfly, Battus philenor.

    PubMed

    Pegram, Kimberly V; Nahm, Alexandra C; Rutowski, Ronald L

    2013-01-01

    Predation on distasteful animals should favor warning coloration that is relatively conspicuous and phenotypically invariable. However, even among similarly colored individuals there can be variation in their warning signals. In butterflies, individual differences in larval feeding history could cause this variation. The warning signal of the pipevine swallowtail butterfly, Battus philenor L. (Lepidoptera: Papilionidae) consists of both a blue iridescent patch and pigmentbased orange spots on the ventral hindwing. B. philenor males also display a dorsal surface iridescent patch that functions as a sexual indicator signal. A previous study of iridescence in B. philenor found that the iridescent blue on both the dorsal and ventral hind wings is variable and significantly different between lab-reared and field-caught individuals. These differences could be the result of larval food deprivation in the field. Through experimental manipulation of larval diet, larval food deprivation was evaluated as a potential cause of the differences observed between lab and field individuals, and if food deprivation is a source of inter-individual variation in warning signals. B. philenor larvae were food restricted starting at two points in the last larval instar, and one group was fed through pupation. Adult coloration was then compared. Food deprivation led to poorer adult condition, as indicated by lower adult body mass, forewing length, and fat content of stressed individuals. As the level of food deprivation increased, the hue of the iridescent patches on both the dorsal and ventral hind wing shifted to shorter wavelengths, and the chroma of the orange spots decreased. The shifts in iridescent color did not match the differences previously found between lab and field individuals. However, the treatment differences indicate that food deprivation may be a significant source of warning color variation. The differences between the treatment groups are likely detectable by predators, but the effect of the variation on signal effectiveness and function remains to be empirically explored.

  1. Metamorphosis of a butterfly-associated bacterial community.

    PubMed

    Hammer, Tobin J; McMillan, W Owen; Fierer, Noah

    2014-01-01

    Butterflies are charismatic insects that have long been a focus of biological research. They are also habitats for microorganisms, yet these microbial symbionts are little-studied, despite their likely importance to butterfly ecology and evolution. In particular, the diversity and composition of the microbial communities inhabiting adult butterflies remain uncharacterized, and it is unknown how the larval (caterpillar) and adult microbiota compare. To address these knowledge gaps, we used Illumina sequencing of 16S rRNA genes from internal bacterial communities associated with multiple life stages of the neotropical butterfly Heliconius erato. We found that the leaf-chewing larvae and nectar- and pollen-feeding adults of H. erato contain markedly distinct bacterial communities, a pattern presumably rooted in their distinct diets. Larvae and adult butterflies host relatively small and similar numbers of bacterial phylotypes, but few are common to both stages. The larval microbiota clearly simplifies and reorganizes during metamorphosis; thus, structural changes in a butterfly's bacterial community parallel those in its own morphology. We furthermore identify specific bacterial taxa that may mediate larval and adult feeding biology in Heliconius and other butterflies. Although male and female Heliconius adults differ in reproductive physiology and degree of pollen feeding, bacterial communities associated with H. erato are not sexually dimorphic. Lastly, we show that captive and wild individuals host different microbiota, a finding that may have important implications for the relevance of experimental studies using captive butterflies.

  2. Metamorphosis of a Butterfly-Associated Bacterial Community

    PubMed Central

    Hammer, Tobin J.; McMillan, W. Owen; Fierer, Noah

    2014-01-01

    Butterflies are charismatic insects that have long been a focus of biological research. They are also habitats for microorganisms, yet these microbial symbionts are little-studied, despite their likely importance to butterfly ecology and evolution. In particular, the diversity and composition of the microbial communities inhabiting adult butterflies remain uncharacterized, and it is unknown how the larval (caterpillar) and adult microbiota compare. To address these knowledge gaps, we used Illumina sequencing of 16S rRNA genes from internal bacterial communities associated with multiple life stages of the neotropical butterfly Heliconius erato. We found that the leaf-chewing larvae and nectar- and pollen-feeding adults of H. erato contain markedly distinct bacterial communities, a pattern presumably rooted in their distinct diets. Larvae and adult butterflies host relatively small and similar numbers of bacterial phylotypes, but few are common to both stages. The larval microbiota clearly simplifies and reorganizes during metamorphosis; thus, structural changes in a butterfly's bacterial community parallel those in its own morphology. We furthermore identify specific bacterial taxa that may mediate larval and adult feeding biology in Heliconius and other butterflies. Although male and female Heliconius adults differ in reproductive physiology and degree of pollen feeding, bacterial communities associated with H. erato are not sexually dimorphic. Lastly, we show that captive and wild individuals host different microbiota, a finding that may have important implications for the relevance of experimental studies using captive butterflies. PMID:24466308

  3. Barcoding the butterflies of southern South America: Species delimitation efficacy, cryptic diversity and geographic patterns of divergence.

    PubMed

    Lavinia, Pablo D; Núñez Bustos, Ezequiel O; Kopuchian, Cecilia; Lijtmaer, Darío A; García, Natalia C; Hebert, Paul D N; Tubaro, Pablo L

    2017-01-01

    Because the tropical regions of America harbor the highest concentration of butterfly species, its fauna has attracted considerable attention. Much less is known about the butterflies of southern South America, particularly Argentina, where over 1,200 species occur. To advance understanding of this fauna, we assembled a DNA barcode reference library for 417 butterfly species of Argentina, focusing on the Atlantic Forest, a biodiversity hotspot. We tested the efficacy of this library for specimen identification, used it to assess the frequency of cryptic species, and examined geographic patterns of genetic variation, making this study the first large-scale genetic assessment of the butterflies of southern South America. The average sequence divergence to the nearest neighbor (i.e. minimum interspecific distance) was 6.91%, ten times larger than the mean distance to the furthest conspecific (0.69%), with a clear barcode gap present in all but four of the species represented by two or more specimens. As a consequence, the DNA barcode library was extremely effective in the discrimination of these species, allowing a correct identification in more than 95% of the cases. Singletons (i.e. species represented by a single sequence) were also distinguishable in the gene trees since they all had unique DNA barcodes, divergent from those of the closest non-conspecific. The clustering algorithms implemented recognized from 416 to 444 barcode clusters, suggesting that the actual diversity of butterflies in Argentina is 3%-9% higher than currently recognized. Furthermore, our survey added three new records of butterflies for the country (Eurema agave, Mithras hannelore, Melanis hillapana). In summary, this study not only supported the utility of DNA barcoding for the identification of the butterfly species of Argentina, but also highlighted several cases of both deep intraspecific and shallow interspecific divergence that should be studied in more detail.

  4. Barcoding the butterflies of southern South America: Species delimitation efficacy, cryptic diversity and geographic patterns of divergence

    PubMed Central

    Núñez Bustos, Ezequiel O.; Kopuchian, Cecilia; Lijtmaer, Darío A.; García, Natalia C.; Hebert, Paul D. N.; Tubaro, Pablo L.

    2017-01-01

    Because the tropical regions of America harbor the highest concentration of butterfly species, its fauna has attracted considerable attention. Much less is known about the butterflies of southern South America, particularly Argentina, where over 1,200 species occur. To advance understanding of this fauna, we assembled a DNA barcode reference library for 417 butterfly species of Argentina, focusing on the Atlantic Forest, a biodiversity hotspot. We tested the efficacy of this library for specimen identification, used it to assess the frequency of cryptic species, and examined geographic patterns of genetic variation, making this study the first large-scale genetic assessment of the butterflies of southern South America. The average sequence divergence to the nearest neighbor (i.e. minimum interspecific distance) was 6.91%, ten times larger than the mean distance to the furthest conspecific (0.69%), with a clear barcode gap present in all but four of the species represented by two or more specimens. As a consequence, the DNA barcode library was extremely effective in the discrimination of these species, allowing a correct identification in more than 95% of the cases. Singletons (i.e. species represented by a single sequence) were also distinguishable in the gene trees since they all had unique DNA barcodes, divergent from those of the closest non-conspecific. The clustering algorithms implemented recognized from 416 to 444 barcode clusters, suggesting that the actual diversity of butterflies in Argentina is 3%–9% higher than currently recognized. Furthermore, our survey added three new records of butterflies for the country (Eurema agave, Mithras hannelore, Melanis hillapana). In summary, this study not only supported the utility of DNA barcoding for the identification of the butterfly species of Argentina, but also highlighted several cases of both deep intraspecific and shallow interspecific divergence that should be studied in more detail. PMID:29049373

  5. A statistical study of ion pitch-angle distributions

    NASA Technical Reports Server (NTRS)

    Sibeck, D. G.; Mcentire, R. W.; Lui, A. T. Y.; Krimigis, S. M.

    1987-01-01

    Preliminary results of a statistical study of energetic (34-50 keV) ion pitch-angle distributions (PADs) within 9 Re of earth provide evidence for an orderly pattern consistent with both drift-shell splitting and magnetopause shadowing. Normal ion PADs dominate the dayside and inner magnetosphere. Butterfly PADs typically occur in a narrow belt stretching from dusk to dawn through midnight, where they approach within 6 Re of earth. While those ion butterfly PADs that typically occur on closed drift paths are mainly caused by drift-shell splitting, there is also evidence for magnetopause shadowing in observations of more frequent butterfly PAD occurrence in the outer magnetosphere near dawn than dusk. Isotropic and gradient boundary PADs terminate the tailward extent of the butterfly ion PAD belt.

  6. Flapping, wobbling, and zig-zagging: Tomographic PIV measurements of Antarctic sea butterfly ``flying'' underwater

    NASA Astrophysics Data System (ADS)

    Adhikari, D.; Webster, D. R.; Yen, J.

    2015-11-01

    A portable tomographic PIV technique was used to study the fluid dynamics and kinematics of sea butterflies in Antarctica. Antarctic pteropods (or sea butterflies), which are currently threatened by ocean acidification, swim in seawater with a pair of gelatinous parapodia (or ``wings'') via a unique propulsion mechanism. Both power and recovery strokes propel the organism (1.5 - 5 mm in size) upward in a sawtooth-like trajectory with average speed of 14 - 30 mm/s and pitch the shell forwards-and-backwards at 1.9 - 3 Hz. The pitching motion effectively positions the parapodia such that they stroke downward during both the power and recovery strokes. Reynolds numbers defined for flapping, translating, and pitching (i.e. Ref, ReU, and ReΩ) characterize the motion of the pteropod. For Ref <50, the shell does not pitch and the pteropod swims abnormally. We present a detailed comparison of the volumetric fluid velocity fields induced by pteropods swimming upwards with Ref = 80 and 180. The pteropod at the lower Ref creates an attached shear flow along the parapodia and pushes fluid in a method analogous to a paddle. In contrast, at higher Ref, the flow along the parapodia separates and generates complex vortex structures.

  7. Sexual dimorphism in the compound eye of Heliconius erato: a nymphalid butterfly with at least five spectral classes of photoreceptor.

    PubMed

    McCulloch, Kyle J; Osorio, Daniel; Briscoe, Adriana D

    2016-08-01

    Most butterfly families expand the number of spectrally distinct photoreceptors in their compound eye by opsin gene duplications together with lateral filter pigments; however, most nymphalid genera have limited diversity, with only three or four spectral types of photoreceptor. Here, we examined the spatial pattern of opsin expression and photoreceptor spectral sensitivities in Heliconius erato, a nymphalid with duplicate ultraviolet opsin genes, UVRh1 and UVRh2 We found that the H. erato compound eye is sexually dimorphic. Females express the two UV opsin proteins in separate photoreceptors, but males do not express UVRh1. Intracellular recordings confirmed that females have three short wavelength-sensitive photoreceptors (λmax=356, ∼390 and 470 nm), while males have two (λmax=390 and ∼470 nm). We also found two long wavelength-sensitive photoreceptors (green, λmax∼555 nm, and red, λmax∼600 nm), which express the same LW opsin. The red cell's shifted sensitivity is probably due to perirhabdomal filtering pigments. Sexual dimorphism of the UV-absorbing rhodopsins may reflect the females' need to discriminate conspecifics from co-mimics. Red-green color vision may be used to detect differences in red coloration on Heliconius wings, or for host-plant identification. Among nymphalids so far investigated, only H. erato is known to possess five spectral classes of photoreceptor; sexual dimorphism of the eye via suppression of one class of opsin (here UVRh1 in males) has not - to our knowledge - been reported in any animal. © 2016. Published by The Company of Biologists Ltd.

  8. Japanese Papilio butterflies puddle using Na+ detected by contact chemosensilla in the proboscis

    NASA Astrophysics Data System (ADS)

    Inoue, Takashi A.; Hata, Tamako; Asaoka, Kiyoshi; Ito, Tetsuo; Niihara, Kinuko; Hagiya, Hiroshi; Yokohari, Fumio

    2012-12-01

    Many butterflies acquire nutrients from non-nectar sources such as puddles. To better understand how male Papilio butterflies identify suitable sites for puddling, we used behavioral and electrophysiological methods to examine the responses of Japanese Papilio butterflies to Na+, K+, Ca2+, and Mg2+. Based on behavioral analyses, these butterflies preferred a 10-mM Na+ solution to K+, Ca2+, and Mg2+ solutions of the same concentration and among a tested range of 1 mM to 1 M NaCl. We also measured the ion concentrations of solutions sampled from puddling sites in the field. Na+ concentrations of the samples were up to 6 mM, slightly lower than that preferred by butterflies in the behavioral experiments. Butterflies that sipped the 10 mM Na+ solution from the experimental trays did not continue to puddle on the ground. Additionally, butterflies puddled at sites where the concentrations of K+, Ca2+, and/or Mg2+ were higher than that of Na+. This suggests that K+, Ca2+, and Mg2+ do not interfere with the detection of Na+ by the Papilio butterfly. Using an electrophysiological method, tip recordings, receptor neurons in contact chemosensilla inside the proboscis evoked regularly firing impulses to 1, 10, and 100 mM NaCl solutions but not to CaCl2 or MgCl2. The dose-response patterns to the NaCl solutions were different among the neurons, which were classified into three types. These results showed that Japanese Papilio butterflies puddle using Na+ detected by the contact chemosensilla in the proboscis, which measure its concentration.

  9. Parallel evolution of Batesian mimicry supergene in two Papilio butterflies, P. polytes and P. memnon

    PubMed Central

    Itoh, Takehiko

    2018-01-01

    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. PMID:29675466

  10. The anti-bat strategy of ultrasound absorption: the wings of nocturnal moths (Bombycoidea: Saturniidae) absorb more ultrasound than the wings of diurnal moths (Chalcosiinae: Zygaenoidea: Zygaenidae).

    PubMed

    Ntelezos, Athanasios; Guarato, Francesco; Windmill, James F C

    2017-01-15

    The selection pressure from echolocating bats has driven the development of a diverse range of anti-bat strategies in insects. For instance, several studies have proposed that the wings of some moths absorb a large portion of the sound energy contained in a bat's ultrasonic cry; as a result, the bat receives a dampened echo, and the moth becomes invisible to the bat. To test the hypothesis that greater exposure to bat predation drives the development of higher ultrasound absorbance, we used a small reverberation chamber to measure the ultrasound absorbance of the wings of nocturnal (Bombycoidea: Saturniidae) and diurnal moths (Chalcosiinae: Zygaenoidea: Zygaenidae). The absorption factor of the nocturnal saturniids peaks significantly higher than the absorption factor of the diurnal chalcosiines. However, the wings of the chalcosiines absorb more ultrasound than the wings of some diurnal butterflies. Following a phylogenetic analysis on the character state of diurnality/ nocturnality in the Zygaenidae, we propose that diurnality in the Chalcosiinae is plesiomorphic (retained); hence, the absorbance of their wings is probably not a vestigial trait from an ancestral, nocturnal form but an adaptation to bat activity that overlaps their own. On a within-species level, females of the saturniids Argema mittrei and Samia cynthia ricini have significantly higher absorption factors than the males. In the female S. c. ricini, the higher absorption factor corresponds to a detection distance by bats that is at best 20-30% shorter than that of the male. © 2017. Published by The Company of Biologists Ltd.

  11. The anti-bat strategy of ultrasound absorption: the wings of nocturnal moths (Bombycoidea: Saturniidae) absorb more ultrasound than the wings of diurnal moths (Chalcosiinae: Zygaenoidea: Zygaenidae)

    PubMed Central

    Guarato, Francesco; Windmill, James F. C.

    2017-01-01

    ABSTRACT The selection pressure from echolocating bats has driven the development of a diverse range of anti-bat strategies in insects. For instance, several studies have proposed that the wings of some moths absorb a large portion of the sound energy contained in a bat's ultrasonic cry; as a result, the bat receives a dampened echo, and the moth becomes invisible to the bat. To test the hypothesis that greater exposure to bat predation drives the development of higher ultrasound absorbance, we used a small reverberation chamber to measure the ultrasound absorbance of the wings of nocturnal (Bombycoidea: Saturniidae) and diurnal moths (Chalcosiinae: Zygaenoidea: Zygaenidae). The absorption factor of the nocturnal saturniids peaks significantly higher than the absorption factor of the diurnal chalcosiines. However, the wings of the chalcosiines absorb more ultrasound than the wings of some diurnal butterflies. Following a phylogenetic analysis on the character state of diurnality/ nocturnality in the Zygaenidae, we propose that diurnality in the Chalcosiinae is plesiomorphic (retained); hence, the absorbance of their wings is probably not a vestigial trait from an ancestral, nocturnal form but an adaptation to bat activity that overlaps their own. On a within-species level, females of the saturniids Argema mittrei and Samia cynthia ricini have significantly higher absorption factors than the males. In the female S. c. ricini, the higher absorption factor corresponds to a detection distance by bats that is at best 20-30% shorter than that of the male. PMID:27913454

  12. Dynamic Pattern Formation for Wings of Pterygota in an Eclosion ---Pattern Analysis for Wings with the Imago---

    NASA Astrophysics Data System (ADS)

    Seino, M.; Kakazu, Y.

    The vein and cell patterns for the fore and hind wing of Lepidoptera, Hemiptera, Orthoptera and Odonata are analyzed and discussed. For vein patterns of them, the fractal properties are shown and the inequality between four orders is obtained. The nature of wings observed by mass distributions for fractal dimensions of the vein pattern is presented.

  13. Stable structural color patterns displayed on transparent insect wings.

    PubMed

    Shevtsova, Ekaterina; Hansson, Christer; Janzen, Daniel H; Kjærandsen, Jostein

    2011-01-11

    Color patterns play central roles in the behavior of insects, and are important traits for taxonomic studies. Here we report striking and stable structural color patterns--wing interference patterns (WIPs)--in the transparent wings of small Hymenoptera and Diptera, patterns that have been largely overlooked by biologists. These extremely thin wings reflect vivid color patterns caused by thin film interference. The visibility of these patterns is affected by the way the insects display their wings against various backgrounds with different light properties. The specific color sequence displayed lacks pure red and matches the color vision of most insects, strongly suggesting that the biological significance of WIPs lies in visual signaling. Taxon-specific color patterns are formed by uneven membrane thickness, pigmentation, venation, and hair placement. The optically refracted pattern is also stabilized by microstructures of the wing such as membrane corrugations and spherical cell structures that reinforce the pattern and make it essentially noniridescent over a large range of light incidences. WIPs can be applied to map the micromorphology of wings through direct observation and are useful in several fields of biology. We demonstrate their usefulness as identification patterns to solve cases of cryptic species complexes in tiny parasitic wasps, and indicate their potentials for research on the genetic control of wing development through direct links between the transregulatory wing landscape and interference patterns we observe in Drosophila model species. Some species display sexually dimorphic WIPs, suggesting sexual selection as one of the driving forces for their evolution.

  14. Butterfly Larval Host Plant use in a Tropical Urban Context: Life History Associations, Herbivory, and Landscape Factors

    PubMed Central

    Tiple, Ashish D.; Khurad, Arun M.; Dennis, Roger L. H.

    2011-01-01

    This study examines butterfly larval host plants, herbivory and related life history attributes within Nagpur City, India. The larval host plants of 120 butterfly species are identified and their host specificity, life form, biotope, abundance and perennation recorded; of the 126 larval host plants, most are trees (49), with fewer herbs (43), shrubs (22), climbers (7) and stem parasites (2). They include 89 wild, 23 cultivated, 11 wild/cultivated and 3 exotic plant species; 78 are perennials, 43 annuals and 5 biannuals. Plants belonging to Poaceae and Fabaceae are most widely used by butterfly larvae. In addition to distinctions in host plant family affiliation, a number of significant differences between butterfly families have been identified in host use patterns: for life forms, biotopes, landforms, perennation, host specificity, egg batch size and ant associations. These differences arising from the development of a butterfly resource database have important implications for conserving butterfly species within the city area. Differences in overall butterfly population sizes within the city relate mainly to the number of host plants used, but other influences, including egg batch size and host specificity are identified. Much of the variation in population size is unaccounted for and points to the need to investigate larval host plant life history and strategies as population size is not simply dependent on host plant abundance. PMID:21864159

  15. Stable structural color patterns displayed on transparent insect wings

    PubMed Central

    Shevtsova, Ekaterina; Hansson, Christer; Janzen, Daniel H.; Kjærandsen, Jostein

    2011-01-01

    Color patterns play central roles in the behavior of insects, and are important traits for taxonomic studies. Here we report striking and stable structural color patterns—wing interference patterns (WIPs)—in the transparent wings of small Hymenoptera and Diptera, patterns that have been largely overlooked by biologists. These extremely thin wings reflect vivid color patterns caused by thin film interference. The visibility of these patterns is affected by the way the insects display their wings against various backgrounds with different light properties. The specific color sequence displayed lacks pure red and matches the color vision of most insects, strongly suggesting that the biological significance of WIPs lies in visual signaling. Taxon-specific color patterns are formed by uneven membrane thickness, pigmentation, venation, and hair placement. The optically refracted pattern is also stabilized by microstructures of the wing such as membrane corrugations and spherical cell structures that reinforce the pattern and make it essentially noniridescent over a large range of light incidences. WIPs can be applied to map the micromorphology of wings through direct observation and are useful in several fields of biology. We demonstrate their usefulness as identification patterns to solve cases of cryptic species complexes in tiny parasitic wasps, and indicate their potentials for research on the genetic control of wing development through direct links between the transregulatory wing landscape and interference patterns we observe in Drosophila model species. Some species display sexually dimorphic WIPs, suggesting sexual selection as one of the driving forces for their evolution. PMID:21199954

  16. Monitoring Temporal Variation to Assess Changes in the Structure of Subtropical Atlantic Forest Butterfly Communities.

    PubMed

    Iserhard, Cristiano Agra; Romanowski, Helena Piccoli; Richter, Aline; Mendonça, Milton de Souza

    2017-08-01

    The study of fauna through long-term surveys is important in unveiling how temporal patterns shape the structure of communities in tropical habitats. The butterfly assemblage of the subtropical Atlantic Forest may be considered highly diverse and shows changes in diversity and composition over time, highlighting the importance of long-term inventories. This work assessed temporal diversity patterns in the distribution and composition of butterfly assemblages in an Atlantic Forest site in southern Brazil using combined data from three years of standardized sampling with entomological nets, increasing the knowledge on this group in the Neotropics for monitoring and conservation. The butterfly fauna was analyzed in terms of richness, abundance, and composition. The inventories reached 401 species, with 14,442 butterfly individuals sampled. All the diversity parameters evaluated show significant differences between the first year of sampling compared to the second and third years. The latter had higher values of richness and abundance, followed by the first and second years. Hesperiidae was the richest family, followed by Nymphalidae and Lycaenidae, indicating a good representation of the assemblage as a whole. The results of this work are important for developing conservation programs in the Atlantic Forest and other forested environments in the neotropics, especially concerning reliable diversity assessments for the monitoring and management of protected areas. Decision making and public policy might also benefit from knowledge on temporal patterns of diversity regarding the maintenance of native habitats and integrity of biomes and their associated fauna. © The Authors 2017. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For Permissions, please email: journals.permissions@oup.com.

  17. Unique Temporal Expression of Triplicated Long-Wavelength Opsins in Developing Butterfly Eyes

    PubMed Central

    Arikawa, Kentaro; Iwanaga, Tomoyuki; Wakakuwa, Motohiro; Kinoshita, Michiyo

    2017-01-01

    Following gene duplication events, the expression patterns of the resulting gene copies can often diverge both spatially and temporally. Here we report on gene duplicates that are expressed in distinct but overlapping patterns, and which exhibit temporally divergent expression. Butterflies have sophisticated color vision and spectrally complex eyes, typically with three types of heterogeneous ommatidia. The eyes of the butterfly Papilio xuthus express two green- and one red-absorbing visual pigment, which came about via gene duplication events, in addition to one ultraviolet (UV)- and one blue-absorbing visual pigment. We localized mRNAs encoding opsins of these visual pigments in developing eye disks throughout the pupal stage. The mRNAs of the UV and blue opsin are expressed early in pupal development (pd), specifying the type of the ommatidium in which they appear. Red sensitive photoreceptors first express a green opsin mRNA, which is replaced later by the red opsin mRNA. Broadband photoreceptors (that coexpress the green and red opsins) first express the green opsin mRNA, later change to red opsin mRNA and finally re-express the green opsin mRNA in addition to the red mRNA. Such a unique temporal and spatial expression pattern of opsin mRNAs may reflect the evolution of visual pigments and provide clues toward understanding how the spectrally complex eyes of butterflies evolved. PMID:29238294

  18. Rhabdom evolution in butterflies: insights from the uniquely tiered and heterogeneous ommatidia of the Glacial Apollo butterfly, Parnassius glacialis.

    PubMed

    Matsushita, Atsuko; Awata, Hiroko; Wakakuwa, Motohiro; Takemura, Shin-ya; Arikawa, Kentaro

    2012-09-07

    The eye of the Glacial Apollo butterfly, Parnassius glacialis, a 'living fossil' species of the family Papilionidae, contains three types of spectrally heterogeneous ommatidia. Electron microscopy reveals that the Apollo rhabdom is tiered. The distal tier is composed exclusively of photoreceptors expressing opsins of ultraviolet or blue-absorbing visual pigments, and the proximal tier consists of photoreceptors expressing opsins of green or red-absorbing visual pigments. This organization is unique because the distal tier of other known butterflies contains two green-sensitive photoreceptors, which probably function in improving spatial and/or motion vision. Interspecific comparison suggests that the Apollo rhabdom retains an ancestral tiered pattern with some modification to enhance its colour vision towards the long-wavelength region of the spectrum.

  19. Divergent RNA Localisation Patterns of Maternal Genes Regulating Embryonic Patterning in the Butterfly Pararge aegeria

    PubMed Central

    Carter, Jean-Michel; Gibbs, Melanie; Breuker, Casper J.

    2015-01-01

    The maternal effect genes responsible for patterning the embryo along the antero-posterior (AP) axis are broadly conserved in insects. The precise function of these maternal effect genes is the result of the localisation of their mRNA in the oocyte. The main developmental mechanisms involved have been elucidated in Drosophila melanogaster, but recent studies have shown that other insect orders often diverge in RNA localisation patterns. A recent study has shown that in the butterfly Pararge aegeria the distinction between blastodermal embryonic (i.e. germ band) and extra-embryonic tissue (i.e. serosa) is already specified in the oocyte during oogenesis in the ovariole, long before blastoderm cellularisation. To examine the extent by which a female butterfly specifies and patterns the AP axis within the region fated to be the germ band, and whether she specifies a germ plasm, we performed in situ hybridisation experiments on oocytes in P. aegeria ovarioles and on early embryos. RNA localisation of the following key maternal effect genes were investigated: caudal (cad), orthodenticle (otd), hunchback (hb) and four nanos (nos) paralogs, as well as TDRD7 a gene containing a key functional domain (OST-HTH/LOTUS) shared with oskar. TDRD7 was mainly confined to the follicle cells, whilst hb was exclusively zygotically transcribed. RNA of some of the nos paralogs, otd and cad revealed complex localisation patterns within the cortical region prefiguring the germ band (i.e. germ cortex). Rather interestingly, otd was localised within and outside the anterior of the germ cortex. Transcripts of nos-O formed a distinct granular ring in the middle of the germ cortex possibly prefiguring the region where germline stem cells form. These butterfly RNA localisation patterns are highly divergent with respect to other insects, highlighting the diverse ways in which different insect orders maternally regulate early embryogenesis of their offspring. PMID:26633019

  20. Butterfly genome reveals promiscuous exchange of mimicry adaptations among species

    PubMed Central

    Dasmahapatra, Kanchon K; Walters, James R.; Briscoe, Adriana D.; Davey, John W.; Whibley, Annabel; Nadeau, Nicola J.; Zimin, Aleksey V.; Hughes, Daniel S. T.; Ferguson, Laura C.; Martin, Simon H.; Salazar, Camilo; Lewis, James J.; Adler, Sebastian; Ahn, Seung-Joon; Baker, Dean A.; Baxter, Simon W.; Chamberlain, Nicola L.; Chauhan, Ritika; Counterman, Brian A.; Dalmay, Tamas; Gilbert, Lawrence E.; Gordon, Karl; Heckel, David G.; Hines, Heather M.; Hoff, Katharina J.; Holland, Peter W.H.; Jacquin-Joly, Emmanuelle; Jiggins, Francis M.; Jones, Robert T.; Kapan, Durrell D.; Kersey, Paul; Lamas, Gerardo; Lawson, Daniel; Mapleson, Daniel; Maroja, Luana S.; Martin, Arnaud; Moxon, Simon; Palmer, William J.; Papa, Riccardo; Papanicolaou, Alexie; Pauchet, Yannick; Ray, David A.; Rosser, Neil; Salzberg, Steven L.; Supple, Megan A.; Surridge, Alison; Tenger-Trolander, Ayse; Vogel, Heiko; Wilkinson, Paul A.; Wilson, Derek; Yorke, James A.; Yuan, Furong; Balmuth, Alexi L.; Eland, Cathlene; Gharbi, Karim; Thomson, Marian; Gibbs, Richard A.; Han, Yi; Jayaseelan, Joy C.; Kovar, Christie; Mathew, Tittu; Muzny, Donna M.; Ongeri, Fiona; Pu, Ling-Ling; Qu, Jiaxin; Thornton, Rebecca L.; Worley, Kim C.; Wu, Yuan-Qing; Linares, Mauricio; Blaxter, Mark L.; Constant, Richard H. ffrench; Joron, Mathieu; Kronforst, Marcus R.; Mullen, Sean P.; Reed, Robert D.; Scherer, Steven E.; Richards, Stephen; Mallet, James; McMillan, W. Owen; Jiggins, Chris D.

    2012-01-01

    The evolutionary importance of hybridization and introgression has long been debated1. We used genomic tools to investigate introgression in Heliconius, a rapidly radiating genus of neotropical butterflies widely used in studies of ecology, behaviour, mimicry and speciation2-5 . We sequenced the genome of Heliconius melpomene and compared it with other taxa to investigate chromosomal evolution in Lepidoptera and gene flow among multiple Heliconius species and races. Among 12,657 predicted genes for Heliconius, biologically important expansions of families of chemosensory and Hox genes are particularly noteworthy. Chromosomal organisation has remained broadly conserved since the Cretaceous, when butterflies split from the silkmoth lineage. Using genomic resequencing, we show hybrid exchange of genes between three co-mimics, H. melpomene, H. timareta, and H. elevatus, especially at two genomic regions that control mimicry pattern. Closely related Heliconius species clearly exchange protective colour pattern genes promiscuously, implying a major role for hybridization in adaptive radiation. PMID:22722851

  1. Japanese Papilio butterflies puddle using Na+ detected by contact chemosensilla in the proboscis.

    PubMed

    Inoue, Takashi A; Hata, Tamako; Asaoka, Kiyoshi; Ito, Tetsuo; Niihara, Kinuko; Hagiya, Hiroshi; Yokohari, Fumio

    2012-12-01

    Many butterflies acquire nutrients from non-nectar sources such as puddles. To better understand how male Papilio butterflies identify suitable sites for puddling, we used behavioral and electrophysiological methods to examine the responses of Japanese Papilio butterflies to Na(+), K(+), Ca(2+), and Mg(2+). Based on behavioral analyses, these butterflies preferred a 10-mM Na(+) solution to K(+), Ca(2+), and Mg(2+) solutions of the same concentration and among a tested range of 1 mM to 1 M NaCl. We also measured the ion concentrations of solutions sampled from puddling sites in the field. Na(+) concentrations of the samples were up to 6 mM, slightly lower than that preferred by butterflies in the behavioral experiments. Butterflies that sipped the 10 mM Na(+) solution from the experimental trays did not continue to puddle on the ground. Additionally, butterflies puddled at sites where the concentrations of K(+), Ca(2+), and/or Mg(2+) were higher than that of Na(+). This suggests that K(+), Ca(2+), and Mg(2+) do not interfere with the detection of Na(+) by the Papilio butterfly. Using an electrophysiological method, tip recordings, receptor neurons in contact chemosensilla inside the proboscis evoked regularly firing impulses to 1, 10, and 100 mM NaCl solutions but not to CaCl(2) or MgCl(2). The dose-response patterns to the NaCl solutions were different among the neurons, which were classified into three types. These results showed that Japanese Papilio butterflies puddle using Na(+) detected by the contact chemosensilla in the proboscis, which measure its concentration.

  2. Some Possible Cases of Escape Mimicry in Neotropical Butterflies.

    PubMed

    Pinheiro, C E G; Freitas, A V L

    2014-10-01

    The possibility that escape or evasive mimicry evolved in butterflies and other prey insects in a similar fashion to classical Batesian and Müllerian mimicry has long been advanced in the literature. However, there is a general disagreement among lepidopterists and evolutionary biologists on whether or not escape mimicry exists, as well as in which mimicry rings this form of mimicry has evolved. Here, we review some purported cases of escape mimicry in Neotropical butterflies and suggest new mimicry rings involving several species of Archaeoprepona, Prepona, and Doxocopa (the "bright blue bands" ring) and species of Colobura and Hypna (the "creamy bands" ring) where the palatability of butterflies, their ability to escape predator attacks, geographic distribution, relative abundance, and co-occurrence in the same habitats strongly suggest that escape mimicry is involved. In addition, we also indicate other butterfly taxa whose similarities of coloration patterns could be due to escape mimicry and would constitute important case studies for future investigation.

  3. Butterfly Floquet Spectrum in Driven SU(2) Systems

    DOE Office of Scientific and Technical Information (OSTI.GOV)

    Wang Jiao; Department of Physics, Institute of Theoretical Physics and Astrophysics, Xiamen University, Xiamen 361005; Gong Jiangbin

    2009-06-19

    The Floquet spectrum of a class of driven SU(2) systems is shown to display a butterfly pattern with multifractal properties. The level crossing between Floquet states of the same parity or different parities is studied. The results are relevant to studies of fractal statistics, quantum chaos, coherent destruction of tunneling, and the validity of mean-field descriptions of Bose-Einstein condensates.

  4. Pollen Processing Behavior of Heliconius Butterflies: A Derived Grooming Behavior

    PubMed Central

    Hikl, Anna-Laetitia; Krenn, Harald W.

    2011-01-01

    Pollen feeding behaviors Heliconius and Laparus (Lepidoptera: Nymphalidae) represent a key innovation that has shaped other life history traits of these neotropical butterflies. Although all flower visiting Lepidoptera regularly come in contact with pollen, only Heliconius and Laparus butterflies actively collect pollen with the proboscis and subsequently take up nutrients from the pollen grains. This study focused on the behavior of pollen processing and compared the movement patterns with proboscis grooming behavior in various nymphalid butterflies using video analysis. The proboscis movements of pollen processing behavior consisted of a lengthy series of repeated coiling and uncoiling movements in a loosely coiled proboscis position combined with up and down movements and the release of saliva. The proboscis-grooming behavior was triggered by contamination of the proboscis in both pollen feeding and non-pollen feeding nymphalid butterflies. Proboscis grooming movements included interrupted series of coiling and uncoiling movements, characteristic sideways movements, proboscis lifting, and occasionally full extension of the proboscis. Discharge of saliva was more pronounced in pollen feeding species than in non-pollen feeding butterfly species. We conclude that the pollen processing behavior of Heliconius and Laparus is a modified proboscis grooming behavior that originally served to clean the proboscis after contamination with particles. PMID:22208893

  5. Compensation for fluctuations in crosswind drift without stationary landmarks in butterflies migrating over seas.

    PubMed

    Srygley, Robert B.

    2001-01-01

    Migrating insects may fly over large bodies of water that lack landmarks, but little is known about their ability to navigate in such a fluid environment. Using boat navigation instruments to measure compensation for fluctuations in crosswind drift, I investigated the ability of butterflies (Lepidoptera: Hesperiidae, Nymphalidae and Pieridae) to orient with and without landmarks as they migrated naturally over the Caribbean Sea. I used the presence or absence of landmarks or clouds to evaluate their use by the butterflies as guides for compensation. Forty-one per cent of the butterflies compensated for crosswind drift, whereas only 16% did not compensate. No conclusion could be drawn for the remainder. Without landmarks or clouds, butterflies were significantly less likely to compensate for drift than when these local cues were present. Butterflies were more likely to compensate fully in the presence of a landmark than when only clouds were present. Phoebis sennae butterflies drifted in the morning and overcompensated for drift in the afternoon, a pattern found both within and between individuals independent of landmarks. Although I cannot exclude the use of clouds, this would probably result in undercompensation. Hence, a ground reference in conjunction with a sun or magnetic compass is the most likely orientation cue. In the absence of clouds, one butterfly compensated, at least in part, indicating that it was using ripples on the sea surface as a ground reference in conjunction with a sun or magnetic compass. Copyright 2001 The Association for the Study of Animal Behaviour.

  6. Eyespot display in the peacock butterfly triggers antipredator behaviors in naïve adult fowl

    PubMed Central

    2013-01-01

    Large conspicuous eyespots have evolved in multiple taxa and presumably function to thwart predator attacks. Traditionally, large eyespots were thought to discourage predator attacks because they mimicked eyes of the predators’ own predators. However, this idea is controversial and the intimidating properties of eyespots have recently been suggested to simply be a consequence of their conspicuousness. Some lepidopteran species include large eyespots in their antipredation repertoire. In the peacock butterfly, Inachis io, eyespots are typically hidden during rest and suddenly exposed by the butterfly when disturbed. Previous experiments have shown that small wild passerines are intimidated by this display. Here, we test whether eyespots also intimidate a considerably larger bird, domestic fowl, Gallus gallus domesticus, by staging interactions between birds and peacock butterflies that were sham-painted or had their eyespots painted over. Our results show that birds typically fled when peacock butterflies performed their display regardless of whether eyespots were visible or painted over. However, birds confronting butterflies with visible eyespots delayed their return to the butterfly, were more vigilant, and more likely to utter alarm calls associated with detection of ground-based predators, compared with birds confronting butterflies with eyespots painted over. Because production of alarm calls and increased vigilance are antipredation behaviors in the fowl, their reaction suggests that eyespots may elicit fear rather than just an aversion to conspicuous patterns. Our results, therefore, suggest that predators perceive large lepidopteran eyespots as belonging to the eyes of a potential predator. PMID:23243378

  7. Bioinspired Thermoresponsive Photonic Polymers with Hierarchical Structures and Their Unique Properties.

    PubMed

    Lu, Tao; Zhu, Shenmin; Ma, Jun; Lin, Jinyou; Wang, Wanlin; Pan, Hui; Tian, Feng; Zhang, Wang; Zhang, Di

    2015-10-01

    Thermoresponsive photonic materials having hierarchical structures are created by combining a template of Morpho butterfly wings with poly(N-isopropylacrylamide) (PNIPAM) through a chemical bonding and polymerization route. These materials show temperature-induced color tunability. Through reacting with both NIPAM monomers and the amino groups of chitosan in wing scales, glutaraldehyde workes as a bridge by creating chemical bonding between the biotemplate and the PNIPAM. The corresponding reflection peaks red-shift with increase in temperature-an opposite phenomenon to previous studies, demonstrating a thermoresponsive photonic property. This unique phenomenon is caused by the refractive index change due to the volume change of PNIPAM during the temperature rising. This work sets up an efficient strategy for the fabrication of stimuli-responsive photonic materials with hierarchical structures toward extensive applications in science and technology. © 2015 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim.

  8. Do territorial butterflies show a macroecological fighting pattern in response to environmental stability?

    PubMed

    Peixoto, Paulo Enrique Cardoso; Medina, Anderson Matos; Mendoza-Cuenca, Luis

    2014-11-01

    The territorial defense of mating sites by males should be favored when female monopolization is possible. Such monopolization should occur in species in which females emerge asynchronously, since males may have time to copulate with one female before the arrival of other. However, regions with smaller reproductive windows should promote higher synchronicity of female emergence, generating a predictable macroecological pattern associated to the rewards from territorial defense. In this study we evaluated the hypothesis that territorial male butterflies should invest more in fighting in species that occur in areas with stable climatic conditions that should present longer reproductive windows. We compiled studies reporting mean butterfly fighting times, mean trait differences among winners and losers and local Köppen climatic classification (a surrogate for climatic stability). We found that males from butterfly species located in areas with stable climatic conditions fight for longer times. However, although winners were stronger than intruders only in areas with intermediate climatic conditions, there was a marked variation among winner-loser comparisons in species in areas with stable climatic conditions. We conclude that males from butterfly species that occur in areas with stable climatic regimes invest more in fighting due to the higher payoffs accrued with territorial defense, but that such investment does not change the effect of trait asymmetries in settling territorial conflicts. This article is part of a Special Issue entitled: Special Issue:Neotropical Behaviour. Copyright © 2014 Elsevier B.V. All rights reserved.

  9. Effects of Increased Flight on the Energetics and Life History of the Butterfly Speyeria mormonia

    PubMed Central

    Niitepõld, Kristjan; Boggs, Carol L.

    2015-01-01

    Movement uses resources that may otherwise be allocated to somatic maintenance or reproduction. How does increased energy expenditure affect resource allocation? Using the butterfly Speyeria mormonia, we tested whether experimentally increased flight affects fecundity, lifespan or flight capacity. We measured body mass (storage), resting metabolic rate and lifespan (repair and maintenance), flight metabolic rate (flight capacity), egg number and composition (reproduction), and food intake across the adult lifespan. The flight treatment did not affect body mass or lifespan. Food intake increased sufficiently to offset the increased energy expenditure. Total egg number did not change, but flown females had higher early-life fecundity and higher egg dry mass than control females. Egg dry mass decreased with age in both treatments. Egg protein, triglyceride or glycogen content did not change with flight or age, but some components tracked egg dry mass. Flight elevated resting metabolic rate, indicating increased maintenance costs. Flight metabolism decreased with age, with a steeper slope for flown females. This may reflect accelerated metabolic senescence from detrimental effects of flight. These effects of a drawdown of nutrients via flight contrast with studies restricting adult nutrient input. There, fecundity was reduced, but flight capacity and lifespan were unchanged. The current study showed that when food resources were abundant, wing-monomorphic butterflies living in a continuous meadow landscape resisted flight-induced stress, exhibiting no evidence of a flight-fecundity or flight-longevity trade-off. Instead, flight changed the dynamics of energy use and reproduction as butterflies adopted a faster lifestyle in early life. High investment in early reproduction may have positive fitness effects in the wild, as long as food is available. Our results help to predict the effect of stressful conditions on the life history of insects living in a changing world. PMID:26510164

  10. Small Impact Crater

    NASA Technical Reports Server (NTRS)

    2005-01-01

    22 June 2005 This Mars Global Surveyor (MGS) Mars Orbiter Camera (MOC) image shows a small impact crater with a 'butterfly' ejecta pattern. The butterfly pattern results from an oblique impact. Not all oblique impacts result in an elliptical crater, but they can result in a non-radial pattern of ejecta distribution. The two-toned nature of the ejecta -- with dark material near the crater and brighter material further away -- might indicate the nature of subsurface materials. Below the surface, there may be a layer of lighter-toned material, underlain by a layer of darker material. The impact throws these materials out in a pattern that reflects the nature of the underlying layers.

    Location near: 3.7oN, 348.2oW Image width: 3 km (1.9 mi) Illumination from: lower left Season: Northern Autumn

  11. The Fukushima nuclear accident and the pale grass blue butterfly: evaluating biological effects of long-term low-dose exposures.

    PubMed

    Hiyama, Atsuki; Nohara, Chiyo; Taira, Wataru; Kinjo, Seira; Iwata, Masaki; Otaki, Joji M

    2013-08-12

    On August 9th 2012, we published an original research article in Scientific Reports, concluding that artificial radionuclides released from the Fukushima Dai-ichi Nuclear Power Plant exerted genetically and physiologically adverse effects on the pale grass blue butterfly Zizeeria maha in the Fukushima area. Immediately following publication, many questions and comments were generated from all over the world. Here, we have clarified points made in the original paper and answered questions posed by the readers. The following points were clarified. (1) There are many advantages to using the pale grass blue butterfly as an indicator species. (2) The forewings of the individuals collected in Fukushima were significantly smaller than in the northern and southern localities. (3) We observed growth retardation in the butterflies from the Fukushima area. (4) The aberrant colour patterns in the butterflies obtained in the Fukushima area were different from the colour patterns induced by temperature and sibling crosses but similar to those induced by external and internal exposures to the artificial radionuclides and by a chemical mutagen, suggesting that genetic mutations caused the aberrations. (5) This species of butterfly has been plentiful in Fukushima area for at least half a century. We here present specimens collected from Fukushima Prefecture before the accident. (6) Mutation accumulation was detected by the increase in the abnormality rates from May 2011 to September 2011. (7) The abnormal traits were heritable. (8) Our sampling localities were not affected by the tsunami. (9) We used a high enough number of samples to obtain statistically significant results. (10) The standard rearing method was followed, producing normal adults in the control groups. (11) The exposure experiments successfully reproduced the results of the field work. This species of butterfly is vulnerable to long-term low-dose internal and external exposures; however, insect cells are known to be resistant to short-term high-dose irradiation. This discrepancy is reconcilable based on the differences in the experimental conditions. We are just beginning to understand the biological effects of long-term low-dose exposures in animals. Further research is necessary to accurately assess the possible biological effects of the accident.

  12. Unscrambling butterfly oogenesis

    PubMed Central

    2013-01-01

    Background Butterflies are popular model organisms to study physiological mechanisms underlying variability in oogenesis and egg provisioning in response to environmental conditions. Nothing is known, however, about; the developmental mechanisms governing butterfly oogenesis, how polarity in the oocyte is established, or which particular maternal effect genes regulate early embryogenesis. To gain insights into these developmental mechanisms and to identify the conserved and divergent aspects of butterfly oogenesis, we analysed a de novo ovarian transcriptome of the Speckled Wood butterfly Pararge aegeria (L.), and compared the results with known model organisms such as Drosophila melanogaster and Bombyx mori. Results A total of 17306 contigs were annotated, with 30% possibly novel or highly divergent sequences observed. Pararge aegeria females expressed 74.5% of the genes that are known to be essential for D. melanogaster oogenesis. We discuss the genes involved in all aspects of oogenesis, including vitellogenesis and choriogenesis, plus those implicated in hormonal control of oogenesis and transgenerational hormonal effects in great detail. Compared to other insects, a number of significant differences were observed in; the genes involved in stem cell maintenance and differentiation in the germarium, establishment of oocyte polarity, and in several aspects of maternal regulation of zygotic development. Conclusions This study provides valuable resources to investigate a number of divergent aspects of butterfly oogenesis requiring further research. In order to fully unscramble butterfly oogenesis, we also now also have the resources to investigate expression patterns of oogenesis genes under a range of environmental conditions, and to establish their function. PMID:23622113

  13. Coherent random lasing from liquid waveguide gain channels with biological scatters

    NASA Astrophysics Data System (ADS)

    Zhang, Hong; Feng, Guoying; Wang, Shutong; Yang, Chao; Yin, Jiajia; Zhou, Shouhuan

    2014-12-01

    A unidirectional coherent random laser based on liquid waveguide gain channels with biological scatters is demonstrated. The optical feedback of the random laser is provided by both light scattering and waveguide confinement. This waveguide-scattering-feedback scheme not only reduces the pump threshold but also makes the output of random laser directional. The threshold of our random laser is about 11 μJ. The emission spectra can be sensitively tuned by changing pump position due to the micro/nano-scale randomness of butterfly wings. It shows the potential applications of optofluidic random lasers for bio-chemical sensors on-chip.

  14. Engineered biomimicry: polymeric replication of surface features found on insects

    NASA Astrophysics Data System (ADS)

    Pulsifer, Drew P.; Lakhtakia, Akhlesh; Martín-Palma, Raúl J.; Pantano, Carlo G.

    2011-04-01

    By combining the modified conformal-evaporated-film-by-rotation (M-CEFR) technique with nickel electroforming, we have produced master negatives of nonplanar biotemplates. An approximately 250-nm-thick conformal coating of nanocrystaline nickel is deposited on a surface structure of interest found in class Insecta, and the coating is then reinforced with a roughly 60-μm-thick structural layer of nickel by electroforming. This structural layer endows the M-CEFR coating with the mechanical robustness necessary for casting or stamping multiple polymer replicas of the biotemplate. We have made master negatives of blowfly corneas, beetle elytrons, and butterfly wings.

  15. Skipper Richness (Hesperiidae) Along Elevational Gradients in Brazilian Atlantic Forest.

    PubMed

    Carneiro, E; Mielke, O H H; Casagrande, M M; Fiedler, K

    2014-02-01

    Hesperiidae are claimed to be a group of elusive butterflies that need major effort for sampling, thus being frequently omitted from tropical butterfly surveys. As no studies have associated species richness patterns of butterflies with environmental gradients of high altitudes in Brazil, we surveyed Hesperiidae ensembles in Serra do Mar along elevational transects (900-1,800 m above sea level) on three mountains. Transects were sampled 11-12 times on each mountain to evaluate how local species richness is influenced by mountain region, vegetation type, and elevational zones. Patterns were also analyzed for the subfamilies, and after disregarding species that exhibit hilltopping behavior. Species richness was evaluated by the observed richness, Jacknife2 estimator and Chao 1 estimator standardized by sample coverage. Overall, 155 species were collected, but extrapolation algorithms suggest a regional richness of about 220 species. Species richness was far higher in forest than in early successional vegetation or grassland. Richness decreased with elevation, and was higher on Anhangava mountain compared with the two others. Patterns were similar between observed and extrapolated Jacknife2 richness, but vegetation type and mountain richness became altered using sample coverage standardization. Hilltopping species were more easily detected than species that do not show this behavior; however, their inclusion did neither affect estimated richness nor modify the shape of the species accumulation curve. This is the first contribution to systematically study highland butterflies in southern Brazil where all records above 1,200 m are altitudinal extensions of the known geographical ranges of skipper species in the region.

  16. Hubble Camera Resumes Science Operation With Picture Of 'Butterfly' In Space.

    NASA Technical Reports Server (NTRS)

    2002-01-01

    he Hubble Space Telescope's Wide Field and Planetary Camera 2 (WFPC2) is back at work, capturing this black-and-white image of the 'butterfly wing'-shaped nebula, NGC 2346. The nebula is about 2,000 light-years away from Earth in the direction of the constellation Monoceros. It represents the spectacular 'last gasp' of a binary star system at the nebula's center. The image was taken on March 6, as part of the recommissioning of the Hubble Space Telescope's previously installed scientific instruments following the successful servicing of the HST by NASA astronauts in February. WFPC2 was installed in HST during the servicing mission in 1993. At the center of the nebula lies a pair of stars that are so close together that they orbit around each other every 16 days. This is so close that, even with Hubble, the pair of stars cannot be resolved into its two components. One component of this binary is the hot core of a star that has ejected most of its outer layers, producing the surrounding nebula. Astronomers believe that this star, when it evolved and expanded to become a red giant, actually swallowed its companion star in an act of stellar cannibalism. The resulting interaction led to a spiraling together of the two stars, culminating in ejection of the outer layers of the red giant. Most of the outer layers were ejected into a dense disk, which can still be seen in the Hubble image, surrounding the central star. Later the hot star developed a fast stellar wind. This wind, blowing out into the surrounding disk, has inflated the large, wispy hourglass-shaped wings perpendicular to the disk. These wings produce the butterfly appearance when seen in projection. The total diameter of the nebula is about one-third of a light-year, or 2 trillion miles. Our own Sun will eject a nebula about 5 billion years from now. However, the Sun is not a double star, so its nebula may well be more spherical in shape. The image was taken through a filter that shows the light of glowing nitrogen atoms. Scientists are still testing and calibrating the newly installed instruments on Hubble , the Near Infrared Camera and Multi-Object Spectrometer (NICMOS) and the Space Telescope Imaging Spectrograph (STIS). These instruments will be ready to make observations in a few weeks. Credit: Massimo Stiavelli (STScI), and NASA other team member: Inge Heyer (STScI) Image files in GIF and JPEG format and captions may be accessed on the Internet via anonymous ftp from oposite.stsci.edu in /pubinfo.

  17. The Aerodynamics of Hovering Insect Flight. III. Kinematics

    NASA Astrophysics Data System (ADS)

    Ellington, C. P.

    1984-02-01

    Insects in free flight were filmed at 5000 frames per second to determine the motion of their wings and bodies. General comments are offered on flight behaviour and manoeuvrability. Changes in the tilt of the stroke plane with respect to the horizontal provides kinematic control of manoeuvres, analogous to the type of control used for helicopters. A projection analysis technique is described that solves for the orientation of the animal with respect to a camera-based coordinate system, giving full kinematic details for the longitudinal wing and body axes from single-view films. The technique can be applied to all types of flight where the wing motions are bilaterally symmetrical: forward, backward and hovering flight, as well as properly banked turns. An analysis of the errors of the technique is presented, and shows that the reconstructed angles for wing position should be accurate to within 1-2^circ in general. Although measurement of the angles of attack was not possible, visual estimations are given. Only 11 film sequences show flight velocities and accelerations that are small enough for the flight to be considered as `hovering'. Two sequences are presented for a hover-fly using an inclined stroke plane, and nine sequences of hovering with a horizontal stroke plane by another hover-fly, two crane-flies, a drone-fly, a ladybird beetle, a honey bee, and two bumble bees. In general, oscillations in the body position from its mean motion are within measurement error, about 1-2% of the wing length. The amplitudes of oscillation for the body angle are only a few degrees, but the phase relation of this oscillation to the wingbeat cycle could be determined for a few sequences. The phase indicates that the pitching moments governing the oscillations result from the wing lift at the ends of the wingbeat, and not from the wing drag or inertial forces. The mean pitching moment of the wings, which determines the mean body angle, is controlled by shifting the centre of lift over the cycle by changing the mean positional angle of the flapping wings. Deviations of the wing tip path from the stroke plane are never large, and no consistent pattern could be found for the wing paths of different insects; indeed, variations in the path were even observed for individual insects. The wing motion is not greatly different from simple harmonic motion, but does show a general trend towards higher accelerations and decelerations at either end of the wingbeat, with constant velocities during the middle of half-strokes. Root mean square and cube root mean cube angular velocities are on average about 4 and 9% lower than simple harmonic motion. Angles of attack are nearly constant during the middle of half-strokes, typically 35^circ at a position 70% along the wing length. The wing is twisted along its length, with angles of attack at the wing base some 10-20^circ greater than at the tip. The wings rotate through about 110^circ at either end of the wingbeat during 10-20% of the cycle period. The mean velocity of the wing edges during rotation is similar to the mean flapping velocity of the wing tip and greater than the flapping velocity for more proximal wing regions, which indicates that vortex shedding during rotation is comparable with that during flapping. The wings tend to rotate as a flat plate during the first half of rotation, which ends just before, or at, the end of the half-stroke. The hover-fly using an inclined stroke plane provides a notable exception to this general pattern: pronation is delayed and overlaps the beginning of the downstroke. The wing profile flexes along a more or less localized longitudinal axis during the second half of rotation, generating the `flip' profile postulated by Weis-Fogh for the hover-flies. This profile occurs to some extent for all of the insects, and is not exceptionally pronounced for the hover-fly. By the end of rotation the wings are nearly flat again, although a slight camber can sometimes be seen. Weis-Fogh showed that beneficial aerodynamic interference can result when the left and right wings come into contact during rotation at the end of the wingbeat. His `fling' mechanism creates the circulation required for wing lift on the subsequent half-stroke, and can be seen on my films of the Large Cabbage White butterfly, a plume moth, and the Mediterranean flour moth. However, their wings `peel' apart like two pieces of paper being separated, rather than fling open rigidly about the trailing edges. A `partial fling' was found for some insects, with the wings touching only along posterior wing areas. A `near fling' with the wings separated by a fraction of the chord was also observed for many insects. There is a continuous spectrum for the separation distance between the wings, in fact, and the separation can vary for a given insect during different manoeuvres. It is suggested that these variants on Weis-Fogh's fling mechanism also generate circulation for wing lift, although less effectively than a complete fling, and that changes in the separation distance may provide a fine control over the amount of lift produced.

  18. Expression of Genes Involved in Drosophila Wing Morphogenesis and Vein Patterning Are Altered by Spaceflight

    NASA Technical Reports Server (NTRS)

    Parsons-Wingerter, Patricia A.; Hosamani, Ravikumar; Bhattacharya, Sharmila

    2015-01-01

    Imaginal wing discs of Drosophila melanogaster (fruit fly) defined during embryogenesis ultimately result in mature wings of stereotyped (specific) venation patterning. Major regulators of wing disc development are the epidermal growth factor receptor (EGF), Notch, Hedgehog (Hh), Wingless (Wg), and Dpp signaling pathways. Highly stereotyped vascular patterning is also characteristic of tissues in other organisms flown in space such as the mouse retina and leaves of Arabidopsis thaliana. Genetic and other adaptations of vascular patterning to space environmental factors have not yet been systematically quantified, despite widespread recognition of their critical importance for terrestrial and microgravity applications. Here we report changes in gene expression with space flight related to Drosophila wing morphogenesis and vein patterning. In addition, genetically modified phenotypes of increasingly abnormal ectopic wing venation in the Drosophila wing1 were analyzed by NASA's VESsel GENeration Analysis (VESGEN) software2. Our goal is to further develop insightful vascular mappings associated with bioinformatic dimensions of genetic or other molecular phenotypes for correlation with genetic and other molecular profiling relevant to NASA's GeneLab and other Space Biology exploration initiatives.

  19. PARALLEL RACE FORMATION AND THE EVOLUTION OF MIMICRY IN HELICONIUS BUTTERFLIES: A PHYLOGENETIC HYPOTHESIS FROM MITOCHONDRIAL DNA SEQUENCES.

    PubMed

    Brower, Andrew V Z

    1996-02-01

    Mimicry has been a fundamental focus of research since the birth of evolutionary biology yet rarely has been studied from a phylogenetic perspective beyond the simple recognition that mimics are not similar due to common descent. The difficulty of finding characters to discern relationships among closely related and convergent taxa has challenged systematists for more than a century. The phenotypic diversity of wing pattens among mimetic Heliconius adds an additional twist to the problem, because single species contain more than a dozen radically different-looking geographical races even though the mimetic advantage is theoretically highest when all individuals within and between species appear the same. Mitochondrial DNA (mtDNA) offers an independent way to address these issues. In this study, Cytochrome Oxidase I and II sequences from multiple, parallel races of Heliconius erato and Heliconius melpomene are examined, to estimate intraspecific phylogeny and gauge sequence divergence and ages of clades among races within each species. Although phenotypes of sympatric races exhibit remarkable concordance between the two species, the mitochondrial cladograms show that the species have not shared a common evolutionary history. H. erato exhibits a basal split between trans- and cis-Andean groups of races, whereas H. melpomene originates in the Guiana Shield. Diverse races in either species appear to have evolved within the last 200,000 yr, and convergent phenotypes have evolved independently within as well as between species. These results contradict prior theories of the evolution of mimicry based on analysis of wing-pattern genetics. © 1996 The Society for the Study of Evolution.

  20. Sea Butterfly Swimming: Time-resolved Tomographic PIV measurements

    NASA Astrophysics Data System (ADS)

    Murphy, David; Zheng, Lingxiao; Mittal, Rajat; Webster, Donald; Yen, Jeannette

    2011-11-01

    The planktonic sea butterfly Limacina helicina swims by flapping its flexible, wing-like parapodia. The appendage stroke kinematics of this shell-bearing pteropod are three-dimensional and likely contain elements of both drag-based (rowing) and lift-based (flapping) propulsion. Unsteady lift-generating mechanisms such as clap-and-fling may also be present. Upstroke and downstroke motions both propel the animal upward and roll it forwards and backwards, resulting in a sawtooth trajectory. We present time-resolved, tomographic PIV measurements of flow generated by free-swimming pteropods (Limacina helicina) moving upwards with average swimming speeds of 5 - 17 mm/s. The pteropods beat their appendages with a stroke frequency of 4 - 5 Hz. With a size range of 1 - 2 mm, the animals filmed in this study operate in a viscous environment with a Reynolds number of 5 to 20. The volumetric flow measurements provide insight into the three dimensional nature of the flow and into the relative importance of drag- and lift-based propulsion at this low Reynolds number. Preliminary results from Navier-Stokes simulations of the flow associated with the swimming of this organism will also be presented.

  1. Experimental evidence shows no fractionation of strontium isotopes ((87)Sr/(86)Sr) among soil, plants, and herbivores: implications for tracking wildlife and forensic science.

    PubMed

    Flockhart, D T Tyler; Kyser, T Kurt; Chipley, Don; Miller, Nathan G; Norris, D Ryan

    2015-01-01

    Strontium isotopes ((87)Sr/(86)Sr) can be useful biological markers for a wide range of forensic science applications, including wildlife tracking. However, one of the main advantages of using (87)Sr/(86)Sr values, that there is no fractionation from geological bedrock sources through the food web, also happens to be a critical assumption that has never been tested experimentally. We test this assumption by measuring (87)Sr/(86)Sr values across three trophic levels in a controlled greenhouse experiment. Adult monarch butterflies were raised on obligate larval host milkweed plants that were, in turn, grown on seven different soil types collected across Canada. We found no significant differences between (87)Sr/(86)Sr values in leachable Sr from soil minerals, organic soil, milkweed leaves, and monarch butterfly wings. Our results suggest that strontium isoscapes developed from (87)Sr/(86)Sr values in bedrock or soil may serve as a reliable biological marker in forensic science for a range of taxa and across large geographic areas.

  2. Growth and patterning are evolutionarily dissociated in the vestigial wing discs of workers of the red imported fire ant, Solenopsis invicta.

    PubMed

    Bowsher, Julia H; Wray, Gregory A; Abouheif, Ehab

    2007-12-15

    Over the last decade, it has become clear that organismal form is largely determined by developmental and evolutionary changes in the growth and pattern formation of tissues. Yet, there is little known about how these two integrated processes respond to environmental cues or how they evolve relative to one another. Here, we present the discovery of vestigial wing imaginal discs in worker larvae of the red imported fire ant, Solenopsis invicta. These vestigial wing discs are present in all worker larvae, which is uncommon for a species with a large worker size distribution. Furthermore, the growth trajectory of these vestigial discs is distinct from all of the ant species examined to date because they grow at a rate slower than the leg discs. We predicted that the growth trajectory of the vestigial wing discs would be mirrored by evolutionary changes in their patterning. We tested this prediction by examining the expression of three patterning genes, extradenticle, ultrabithorax, and engrailed, known to underlie the wing polyphenism in ants. Surprisingly, the expression patterns of these three genes in the vestigial wing discs was the same as those found in ant species with different worker size distributions and wing disc growth than fire ants. We conclude that growth and patterning are evolutionarily dissociated in the vestigial wing discs of S. invicta because patterning in these discs is conserved, whereas their growth trajectories are not. The evolutionary dissociation of growth and patterning may be an important feature of gene networks that underlie polyphenic traits. 2007 Wiley-Liss, Inc

  3. Effects of patch size and type of coffee matrix on ithomiine butterfly diversity and dispersal in cloud-forest fragments.

    PubMed

    Muriel, Sandra B; Kattan, Gustavo H

    2009-08-01

    Determining the permeability of different types of landscape matrices to animal movement is essential for conserving populations in fragmented landscapes. We evaluated the effects of habitat patch size and matrix type on diversity, isolation, and dispersal of ithomiine butterflies in forest fragments surrounded by coffee agroecosystems in the Colombian Andes. Because ithomiines prefer a shaded understory, we expected the highest diversity and abundance in large fragments surrounded by shade coffee and the lowest in small fragments surrounded by sun coffee. We also thought shade coffee would favor butterfly dispersal and immigration into forest patches. We marked 9675 butterflies of 39 species in 12 forest patches over a year. Microclimate conditions were more similar to the forest interior in the shade-coffee matrix than in the sun-coffee matrix, but patch size and matrix type did not affect species richness and abundance in forest fragments. Furthermore, age structure and temporal recruitment patterns of the butterfly community were similar in all fragments, independent of patch size or matrix type. There were no differences in the numbers of butterflies flying in the matrices at two distances from the forest patch, but their behavior differed. Flight in the sun-coffee matrix was rapid and directional, whereas butterflies in shade-coffee matrix flew slowly. Seven out of 130 recaptured butterflies immigrated into patches in the shade-coffee matrix, and one immigrated into a patch surrounded by sun coffee. Although the shade-coffee matrix facilitated movement in the landscape, sun-coffee matrix was not impermeable to butterflies. Ithomiines exhibited behavioral plasticity in habitat use and high mobility. These traits favor their persistence in heterogeneous landscapes, opening opportunities for their conservation. Understanding the dynamics and resource requirements of different organisms in rural landscapes is critical for identifying management options that address both animals' and farmers' needs.

  4. Non-random dispersal in the butterfly Maniola jurtina: implications for metapopulation models.

    PubMed Central

    Conradt, L; Bodsworth, E J; Roper, T J; Thomas, C D

    2000-01-01

    The dispersal patterns of animals are important in metapopulation ecology because they affect the dynamics and survival of populations. Theoretical models assume random dispersal but little is known in practice about the dispersal behaviour of individual animals or the strategy by which dispersers locate distant habitat patches. In the present study, we released individual meadow brown butterflies (Maniola jurtina) in a non-habitat and investigated their ability to return to a suitable habitat. The results provided three reasons for supposing that meadow brown butterflies do not seek habitat by means of random flight. First, when released within the range of their normal dispersal distances, the butterflies orientated towards suitable habitat at a higher rate than expected at random. Second, when released at larger distances from their habitat, they used a non-random, systematic, search strategy in which they flew in loops around the release point and returned periodically to it. Third, butterflies returned to a familiar habitat patch rather than a non-familiar one when given a choice. If dispersers actively orientate towards or search systematically for distant habitat, this may be problematic for existing metapopulation models, including models of the evolution of dispersal rates in metapopulations. PMID:11007325

  5. Distributions of occupied and vacant butterfly habitats in fragmented landscapes.

    PubMed

    Thomas, C D; Thomas, J A; Warren, M S

    1992-12-01

    We found several rare UK butterflies to be restricted to relatively large and non-isolated habitat patches, while small patches and those that are isolated from population sources remain vacant. These patterns of occurrence are generated by the dynamic processes of local extinction and colonization. Habitat patches act as terrestrial archipelagos in which long-term population persistence, and hence effective long-term conservation, rely on networks of suitable habitats, sufficiently close to allow natural dispersal.

  6. Occupancy modeling of Parnassius clodius butterfly populations in Grand Teton National Park, Wyoming

    USGS Publications Warehouse

    Szcodronski, Kimberly E.; Debinski, Diane M.; Klaver, Robert W.

    2018-01-01

    Estimating occupancy patterns and identifying vegetation characteristics that influence the presence of butterfly species are essential approaches needed for determining how habitat changes may affect butterfly populations in the future. The montane butterfly species, Parnassius clodius, was investigated to identify patterns of occupancy relating to habitat variables in Grand Teton National Park and Bridger-Teton National Forest, Wyoming, United States. A series of presence–absence surveys were conducted in 2013 in 41 mesic to xeric montane meadows that were considered suitable habitat for P. clodius during their flight season (June–July) to estimate occupancy (ψ) and detection probability (p). According to the null constant parameter model, P. clodius had high occupancy of ψ = 0.78 ± 0.07 SE and detection probability of p = 0.75 ± 0.04 SE. In models testing covariates, the most important habitat indicator for the occupancy of P. clodius was a strong negative association with big sagebrush (Artemisia tridentata; β = − 21.39 ± 21.10 SE) and lupine (Lupinus spp.; β = − 20.03 ± 21.24 SE). While P. clodius was found at a high proportion of meadows surveyed, the presence of A. tridentata may limit their distribution within montane meadows at a landscape scale because A. tridentata dominates a large percentage of the montane meadows in our study area. Future climate scenarios predicted for high elevations globally could cause habitat shifts and put populations of P. clodius and similar non-migratory butterfly populations at risk.

  7. Outbred genome sequencing and CRISPR/Cas9 gene editing in butterflies

    PubMed Central

    Li, Xueyan; Fan, Dingding; Zhang, Wei; Liu, Guichun; Zhang, Lu; Zhao, Li; Fang, Xiaodong; Chen, Lei; Dong, Yang; Chen, Yuan; Ding, Yun; Zhao, Ruoping; Feng, Mingji; Zhu, Yabing; Feng, Yue; Jiang, Xuanting; Zhu, Deying; Xiang, Hui; Feng, Xikan; Li, Shuaicheng; Wang, Jun; Zhang, Guojie; Kronforst, Marcus R.; Wang, Wen

    2015-01-01

    Butterflies are exceptionally diverse but their potential as an experimental system has been limited by the difficulty of deciphering heterozygous genomes and a lack of genetic manipulation technology. Here we use a hybrid assembly approach to construct high-quality reference genomes for Papilio xuthus (contig and scaffold N50: 492 kb, 3.4 Mb) and Papilio machaon (contig and scaffold N50: 81 kb, 1.15 Mb), highly heterozygous species that differ in host plant affiliations, and adult and larval colour patterns. Integrating comparative genomics and analyses of gene expression yields multiple insights into butterfly evolution, including potential roles of specific genes in recent diversification. To functionally test gene function, we develop an efficient (up to 92.5%) CRISPR/Cas9 gene editing method that yields obvious phenotypes with three genes, Abdominal-B, ebony and frizzled. Our results provide valuable genomic and technological resources for butterflies and unlock their potential as a genetic model system. PMID:26354079

  8. Assessing Impact Direction in 3-point Bending of Human Femora: Incomplete Butterfly Fractures and Fracture Surfaces,.

    PubMed

    Isa, Mariyam I; Fenton, Todd W; Deland, Trevor; Haut, Roger C

    2018-01-01

    Current literature associates bending failure with butterfly fracture, in which fracture initiates transversely at the tensile surface of a bent bone and branches as it propagates toward the impact surface. The orientation of the resulting wedge fragment is often considered diagnostic of impact direction. However, experimental studies indicate bending does not always produce complete butterfly fractures or produces wedge fragments variably in tension or compression, precluding their use in interpreting directionality. This study reports results of experimental 3-point bending tests on thirteen unembalmed human femora. Complete fracture patterns varied following bending failure, but incomplete fractures and fracture surface characteristics were observed in all impacted specimens. A flat, billowy fracture surface was observed in tension, while jagged, angular peaks were observed in compression. Impact direction was accurately reconstructed using incomplete tension wedge butterfly fractures and tension and compression fracture surface criteria in all thirteen specimens. © 2017 American Academy of Forensic Sciences.

  9. Physiological differences between female limited, alternative life history strategies: The Alba phenotype in the butterfly Colias croceus.

    PubMed

    Woronik, Alyssa; Stefanescu, Constanti; Käkelä, Reijo; Wheat, Christopher W; Lehmann, Philipp

    Across a wide range of taxa, individuals within populations exhibit alternative life history strategies (ALHS) where their phenotypes dramatically differ due to divergent investments in growth, reproduction and survivorship, with the resulting trade-offs directly impacting Darwinian fitness. Though the maintenance of ALHS within populations is fairly well understood, little is known regarding the physiological mechanisms that underlie ALHS and how environmental conditions can affect the evolution and expression of these phenotypes. One such ALHS, known as Alba, exists within females of many species in the butterfly genus Colias. Previous works in New World species not only found that female morphs differ in their wing color due to a reallocation of resources away from the synthesis of wing pigments to other areas of development, but also that temperature played an important role in these trade-offs. Here we build on previous work conducted in New World species by measuring life history traits and conducting lipidomics on individuals reared at hot and cold temperatures in the Old World species Colias croceus. Results suggest that the fitness of Alba and orange morphs likely varies with rearing temperature, where Alba females have higher fitness in cold conditions and orange in warm. Additionally shared traits between Old and New World species suggest the Alba mechanism is likely conserved across the genus. Finally, in the cold treatment we observe an intermediate yellow morph that may have decreased fitness due to slower larval development. This cost may manifest as disruptive selection in the field, thereby favoring the maintenance of the two discrete morphs. Taken together these results add insights into the evolution of, and the selection on, the Alba ALHS. Copyright © 2018 Elsevier Ltd. All rights reserved.

  10. Isolation, identification, and quantification of potential defensive compounds in the viceroy butterfly and its larval host-plant, Carolina willow.

    PubMed

    Prudic, Kathleen L; Khera, Smriti; Sólyom, Anikó; Timmermann, Barbara N

    2007-06-01

    The viceroy-monarch and viceroy-queen butterfly associations are classic examples of mimicry. These relationships were originally classified as Batesian, or parasitic, but were later reclassified as Müllerian, or mutalistic, based on predator bioassays. The Müllerian reclassification implies that viceroy is unpalatable because it too is chemically defended like the queen and the monarch. However, unlike the queen and the monarch, the viceroy defensive chemistry has remained uncharacterized. We demonstrate that the viceroy butterfly (Limenitis archippus, Nymphalidae) not only sequesters nonvolatile defensive compounds from its larval host-plant, the Carolina willow (Salix caroliniana, Salicaceae), but also secretes volatile defensive compounds when disturbed. We developed liquid chromatography-mass spectrometry-mass spectrometry methods to identify a set of phenolic glycosides shared between the adult viceroy butterfly and the Carolina willow, and solid phase microextraction and gas chromatography-mass spectrometry methods to identify volatile phenolic compounds released from stressed viceroy butterflies. In both approaches, all structures were characterized based on their mass spectral fragmentation patterns and confirmed with authentic standards. The phenolics we found are known to deter predator attack in other prey systems, including other willow-feeding insect species. Because these compounds have a generalized defensive function at the concentrations we described, our results are consistent with the Müllerian reclassification put forth by other researchers based on bioassay results. It seems that the viceroy butterfly possesses chemical defenses different from its monarch and queen butterfly counterparts (phenolic glycosides vs. cardiac glycosides, respectively), an unusual phenomenon in mimicry warranting future study.

  11. Citizen Science: The First Peninsular Malaysia Butterfly Count

    PubMed Central

    Jisming-See, Shi-Wei; Brandon-Mong, Guo-Jie; Lim, Aik-Hean; Lim, Voon-Ching; Lee, Ping-Shin; Sing, Kong-Wah

    2015-01-01

    Abstract Background Over the past 50 years, Southeast Asia has suffered the greatest losses of biodiversity of any tropical region in the world. Malaysia is a biodiversity hotspot in the heart of Southeast Asia with roughly the same number of mammal species, three times the number of butterfly species, but only 4% of the land area of Australia. Consequently, in Malaysia, there is an urgent need for biodiversity monitoring and also public engagement with wildlife to raise awareness of biodiversity loss. Citizen science is “on the rise” globally and can make valuable contributions to long-term biodiversity monitoring, but perhaps more importantly, involving the general public in science projects can raise public awareness and promote engagement. Butterflies are often the focus of citizen science projects due to their charisma and familiarity and are particularly valuable “ambassadors” of biodiversity conservation for public outreach. New information Here we present the data from our citizen science project, the first “Peninsular Malaysia Butterfly Count”. Participants were asked to go outdoors on June 6, 2015, and (non-lethally) sample butterfly legs for species identification through DNA barcoding. Fifty-seven citizens responded to our adverts and registered to take part in the butterfly count with many registering on behalf of groups. Collectively the participants sampled 220 butterfly legs from 26 mostly urban and suburban sampling localities. These included our university campus, a highschool, several public parks and private residences. On the basis of 192 usable DNA barcodes, 43 species were sampled by the participants. The most sampled species was Appias olferna, followed by Junonia orithya and Zizina otis. Twenty-two species were only sampled once, five were only sampled twice, and four were only sampled three times. Three DNA barcodes could not be assigned species names. The sampled butterflies revealed that widely distributed, cosmopolitan species, often those recently arrived to the peninsula or with documented "invasive" potential, dominated the habitat types sampled by the participants. Data from this first Butterfly Count helps establish a baseline from which we can monitor the patterns and changes in butterfly communities in Peninsular Malaysia. PMID:26751033

  12. Citizen Science: The First Peninsular Malaysia Butterfly Count.

    PubMed

    Wilson, John-James; Jisming-See, Shi-Wei; Brandon-Mong, Guo-Jie; Lim, Aik-Hean; Lim, Voon-Ching; Lee, Ping-Shin; Sing, Kong-Wah

    2015-01-01

    Over the past 50 years, Southeast Asia has suffered the greatest losses of biodiversity of any tropical region in the world. Malaysia is a biodiversity hotspot in the heart of Southeast Asia with roughly the same number of mammal species, three times the number of butterfly species, but only 4% of the land area of Australia. Consequently, in Malaysia, there is an urgent need for biodiversity monitoring and also public engagement with wildlife to raise awareness of biodiversity loss. Citizen science is "on the rise" globally and can make valuable contributions to long-term biodiversity monitoring, but perhaps more importantly, involving the general public in science projects can raise public awareness and promote engagement. Butterflies are often the focus of citizen science projects due to their charisma and familiarity and are particularly valuable "ambassadors" of biodiversity conservation for public outreach. Here we present the data from our citizen science project, the first "Peninsular Malaysia Butterfly Count". Participants were asked to go outdoors on June 6, 2015, and (non-lethally) sample butterfly legs for species identification through DNA barcoding. Fifty-seven citizens responded to our adverts and registered to take part in the butterfly count with many registering on behalf of groups. Collectively the participants sampled 220 butterfly legs from 26 mostly urban and suburban sampling localities. These included our university campus, a highschool, several public parks and private residences. On the basis of 192 usable DNA barcodes, 43 species were sampled by the participants. The most sampled species was Appias olferna, followed by Junonia orithya and Zizina otis. Twenty-two species were only sampled once, five were only sampled twice, and four were only sampled three times. Three DNA barcodes could not be assigned species names. The sampled butterflies revealed that widely distributed, cosmopolitan species, often those recently arrived to the peninsula or with documented "invasive" potential, dominated the habitat types sampled by the participants. Data from this first Butterfly Count helps establish a baseline from which we can monitor the patterns and changes in butterfly communities in Peninsular Malaysia.

  13. Automatic processing of pragmatic information in the human brain: a mismatch negativity study.

    PubMed

    Zhao, Ming; Liu, Tao; Chen, Feiyan

    2018-05-23

    Language comprehension involves pragmatic information processing, which allows world knowledge to influence the interpretation of a sentence. This study explored whether pragmatic information can be automatically processed during spoken sentence comprehension. The experiment adopted the mismatch negativity (MMN) paradigm to capture the neurophysiological indicators of automatic processing of spoken sentences. Pragmatically incorrect ('Foxes have wings') and correct ('Butterflies have wings') sentences were used as the experimental stimuli. In condition 1, the pragmatically correct sentence was the deviant and the pragmatically incorrect sentence was the standard stimulus, whereas the opposite case was presented in condition 2. The experimental results showed that, compared with the condition that the pragmatically correct sentence is the deviant stimulus, when the condition that the pragmatically incorrect sentence is the deviant stimulus MMN effects were induced within 60-120 and 220-260 ms. The results indicated that the human brain can monitor for incorrect pragmatic information in the inattentive state and can automatically process pragmatic information at the beginning of spoken sentence comprehension.

  14. Asymmetric hindwing foldings in rove beetles.

    PubMed

    Saito, Kazuya; Yamamoto, Shuhei; Maruyama, Munetoshi; Okabe, Yoji

    2014-11-18

    Foldable wings of insects are the ultimate deployable structures and have attracted the interest of aerospace engineering scientists as well as entomologists. Rove beetles are known to fold their wings in the most sophisticated ways that have right-left asymmetric patterns. However, the specific folding process and the reason for this asymmetry remain unclear. This study reveals how these asymmetric patterns emerge as a result of the folding process of rove beetles. A high-speed camera was used to reveal the details of the wing-folding movement. The results show that these characteristic asymmetrical patterns emerge as a result of simultaneous folding of overlapped wings. The revealed folding mechanisms can achieve not only highly compact wing storage but also immediate deployment. In addition, the right and left crease patterns are interchangeable, and thus each wing internalizes two crease patterns and can be folded in two different ways. This two-way folding gives freedom of choice for the folding direction to a rove beetle. The use of asymmetric patterns and the capability of two-way folding are unique features not found in artificial structures. These features have great potential to extend the design possibilities for all deployable structures, from space structures to articles of daily use.

  15. Controlling light with freeform optics: recent progress in computational methods for optical design of freeform lenses with prescribed irradiance properties

    NASA Astrophysics Data System (ADS)

    Oliker, Vladimir I.; Cherkasskiy, Boris

    2014-09-01

    Structural color is produced when nanostructures called schemochromes alter light reflected from a surface through different optic principles, in contrast with other types of colors that are produced when pigments selectively absorb certain wavelengths of light. Research on biogenic photonic nanostructures has focused primarily on bird feathers, butterfly wings and beetle elytra, ignoring other diverse groups such as spiders. We argue that spiders are a good model system to study the functions and evolution of colors in nature for the following reasons. First, these colors clearly function in spiders such as the tarantulas outside of sexual selection, which is likely the dominant driver of the evolution of structural colors in birds and butterflies. Second, within more than 44,000 currently known spider species, colors are used in every possible way based on the same sets of relatively simple materials. Using spiders, we can study how colors evolve to serve different functions under a variety of combinations of driving forces, and how those colors are produced within a relatively simple system. Here, we first review the different color-producing materials and mechanisms (i.e., light absorbing, reflecting and emitting) in birds, butterflies and beetles, the interactions between these different elements, and the functions of colors in different organisms. We then summarize the current state of knowledge of spider colors and compare it with that of birds and insects. We then raise questions including: 1. Could spiders use fluorescence as a mechanism to protect themselves from UV radiation, if they do not have the biosynthetic pathways to produce melanins? 2. What functions could color serve for nearly blind tarantulas? 3. Why are only multilayer nanostructures (thus far) found in spiders, while birds and butterflies use many diverse nanostructures? And, does this limit the diversity of structural colors found in spiders? Answering any of these questions in the future will bring spiders to the forefront of the study of structural colors in nature.

  16. Structural color and its interaction with other color-producing elements: perspectives from spiders

    NASA Astrophysics Data System (ADS)

    Hsiung, Bor-Kai; Blackledge, Todd A.; Shawkey, Matthew D.

    2014-09-01

    Structural color is produced when nanostructures called schemochromes alter light reflected from a surface through different optic principles, in contrast with other types of colors that are produced when pigments selectively absorb certain wavelengths of light. Research on biogenic photonic nanostructures has focused primarily on bird feathers, butterfly wings and beetle elytra, ignoring other diverse groups such as spiders. We argue that spiders are a good model system to study the functions and evolution of colors in nature for the following reasons. First, these colors clearly function in spiders such as the tarantulas outside of sexual selection, which is likely the dominant driver of the evolution of structural colors in birds and butterflies. Second, within more than 44,000 currently known spider species, colors are used in every possible way based on the same sets of relatively simple materials. Using spiders, we can study how colors evolve to serve different functions under a variety of combinations of driving forces, and how those colors are produced within a relatively simple system. Here, we first review the different color-producing materials and mechanisms (i.e., light absorbing, reflecting and emitting) in birds, butterflies and beetles, the interactions between these different elements, and the functions of colors in different organisms. We then summarize the current state of knowledge of spider colors and compare it with that of birds and insects. We then raise questions including: 1. Could spiders use fluorescence as a mechanism to protect themselves from UV radiation, if they do not have the biosynthetic pathways to produce melanins? 2. What functions could color serve for nearly blind tarantulas? 3. Why are only multilayer nanostructures (thus far) found in spiders, while birds and butterflies use many diverse nanostructures? And, does this limit the diversity of structural colors found in spiders? Answering any of these questions in the future will bring spiders to the forefront of the study of structural colors in nature.

  17. Monitoring change in the abundance and distribution of insects using butterflies and other indicator groups

    PubMed Central

    Thomas, J.A

    2005-01-01

    Conservative estimates suggest that 50–90% of the existing insect species on Earth have still to be discovered, yet the named insects alone comprise more than half of all known species of organism. With such poor baseline knowledge, monitoring change in insect diversity poses a formidable challenge to scientists and most attempts to generalize involve large extrapolations from a few well-studied taxa. Butterflies are often the only group for which accurate measures of change can be obtained. Four schemes, used successfully to assess change in British butterflies, that are increasingly being applied across the world are described: Red Data Books (RDB) list the best judgements of experts of the conservation status of species in their field of expertise; mapping schemes plot the changing distributions of species at scales of 1–100 km2; transect monitoring schemes generate time series of changes in abundance in sample populations of species on fixed sites across the UK; and occasional surveys measure the number, boundaries and size of all populations of a (usually RDB) species at intervals of 10–30 years. All schemes describe consistent patterns of change, but if they are to be more generally useful, it is important to understand how well butterflies are representative of other taxa. Comparisons with similarly measured changes in native bird and plant species suggest that butterflies have declined more rapidly that these other groups in Britain; it should soon be possible to test whether this pattern exists elsewhere. It is also demonstrated that extinction rates in British butterflies are similar to those in a range of other insect groups over 100 years once recording bias is accounted for, although probably lower than in aquatic or parasitic taxa. It is concluded that butterflies represent adequate indicators of change for many terrestrial insect groups, but recommended that similar schemes be extended to other popular groups, especially dragonflies, bumblebees, hoverflies and ants. Given institutional backing, similar projects could be employed internationally and standardized. Finally, a range of schemes designed to monitor change in communities of aquatic macro-invertebrates is described. Although designed to use invertebrates as a bio-indicator of water quality for human use, these programmes could be extended to monitor the 2010 biodiversity targets of the World Summit on Sustainable Development. PMID:15814349

  18. Monitoring change in the abundance and distribution of insects using butterflies and other indicator groups.

    PubMed

    Thomas, J A

    2005-02-28

    Conservative estimates suggest that 50-90% of the existing insect species on Earth have still to be discovered, yet the named insects alone comprise more than half of all known species of organism. With such poor baseline knowledge, monitoring change in insect diversity poses a formidable challenge to scientists and most attempts to generalize involve large extrapolations from a few well-studied taxa. Butterflies are often the only group for which accurate measures of change can be obtained. Four schemes, used successfully to assess change in British butterflies, that are increasingly being applied across the world are described: Red Data Books (RDB) list the best judgements of experts of the conservation status of species in their field of expertise; mapping schemes plot the changing distributions of species at scales of 1-100 km2; transect monitoring schemes generate time series of changes in abundance in sample populations of species on fixed sites across the UK; and occasional surveys measure the number, boundaries and size of all populations of a (usually RDB) species at intervals of 10-30 years. All schemes describe consistent patterns of change, but if they are to be more generally useful, it is important to understand how well butterflies are representative of other taxa. Comparisons with similarly measured changes in native bird and plant species suggest that butterflies have declined more rapidly that these other groups in Britain; it should soon be possible to test whether this pattern exists elsewhere. It is also demonstrated that extinction rates in British butterflies are similar to those in a range of other insect groups over 100 years once recording bias is accounted for, although probably lower than in aquatic or parasitic taxa. It is concluded that butterflies represent adequate indicators of change for many terrestrial insect groups, but recommended that similar schemes be extended to other popular groups, especially dragonflies, bumblebees, hoverflies and ants. Given institutional backing, similar projects could be employed internationally and standardized. Finally, a range of schemes designed to monitor change in communities of aquatic macro-invertebrates is described. Although designed to use invertebrates as a bio-indicator of water quality for human use, these programmes could be extended to monitor the 2010 biodiversity targets of the World Summit on Sustainable Development.

  19. The Fukushima nuclear accident and the pale grass blue butterfly: evaluating biological effects of long-term low-dose exposures

    PubMed Central

    2013-01-01

    Background On August 9th 2012, we published an original research article in Scientific Reports, concluding that artificial radionuclides released from the Fukushima Dai-ichi Nuclear Power Plant exerted genetically and physiologically adverse effects on the pale grass blue butterfly Zizeeria maha in the Fukushima area. Immediately following publication, many questions and comments were generated from all over the world. Here, we have clarified points made in the original paper and answered questions posed by the readers. Results The following points were clarified. (1) There are many advantages to using the pale grass blue butterfly as an indicator species. (2) The forewings of the individuals collected in Fukushima were significantly smaller than in the northern and southern localities. (3) We observed growth retardation in the butterflies from the Fukushima area. (4) The aberrant colour patterns in the butterflies obtained in the Fukushima area were different from the colour patterns induced by temperature and sibling crosses but similar to those induced by external and internal exposures to the artificial radionuclides and by a chemical mutagen, suggesting that genetic mutations caused the aberrations. (5) This species of butterfly has been plentiful in Fukushima area for at least half a century. We here present specimens collected from Fukushima Prefecture before the accident. (6) Mutation accumulation was detected by the increase in the abnormality rates from May 2011 to September 2011. (7) The abnormal traits were heritable. (8) Our sampling localities were not affected by the tsunami. (9) We used a high enough number of samples to obtain statistically significant results. (10) The standard rearing method was followed, producing normal adults in the control groups. (11) The exposure experiments successfully reproduced the results of the field work. This species of butterfly is vulnerable to long-term low-dose internal and external exposures; however, insect cells are known to be resistant to short-term high-dose irradiation. This discrepancy is reconcilable based on the differences in the experimental conditions. Conclusions We are just beginning to understand the biological effects of long-term low-dose exposures in animals. Further research is necessary to accurately assess the possible biological effects of the accident. PMID:23937355

  20. The genetic pattern of population threat and loss: a case study of butterflies.

    PubMed

    Schmitt, T; Hewitt, G M

    2004-01-01

    Recent decreases in biodiversity in Europe are commonly thought to be due to land use and climate change. However, the genetic diversity of populations is also seen as one essential factor for their fitness. Genetic diversity in species across the continent of Europe has been recognized as being in part a consequence of ice age isolation in southern refugia and postglacial colonization northwards, and these phylogeographical patterns may themselves affect the adaptability of populations. Recent work on butterfly species with different refugia, colonization paths and genetic structures allows this idea to be examined. The 'chalk-hill blue' pattern is one of decreasing genetic diversity from south to north, whereas the 'woodland ringlet' pattern shows greater genetic diversity in eastern than in western lineages. Comparison of population demographic trends in species with these biogeographical patterns reveals higher rates of decrease with lower genetic diversity. This indicates reduced adaptability due to genetic impoverishment as a result of glacial and postglacial range changes. Analysis of phylogeographical pattern may be a useful guide to interpreting demographic trends and in conservation planning.

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