Butterfly Papilio Research Articles

A model for population dynamics of the mimetic butterfly Papilio polytes in the Sakishima Islands, Japan

We present a mathematical model for population dynamics of the mimetic swallowtail butterfly Papilio polytes in the Sakishima Islands, Japan. The model includes four major variables, that is, population densities of three kinds of butterflies (two female forms f. cyrus, f. polytes and the unpalatable butterfly Pachliopta aristolochiae) and their predator. It is well-known that the non-mimic f. cyrus resembles and attracts the male most, and the mimic f. polytes mimics the model butterfly P. aristolochiae. Based on experimental evidence, we assume that two forms f. cyrus and f. polytes interact under intraspecific competition for resources including the male, and the growth rate of f. cyrus is higher than that of f. polytes. We further assume that both the benefit of mimicry for the mimic f. polytes and the cost for the model are dependent on their relative frequencies, i.e. the motality of the mimic by predation decreases with increase in frequency of the model, while the motality of the model increases as the frequency of the mimic increases. Taking the density-dependent effect through carrying capacity into account, we set up a model system consisting of three ordinary differential equations (ODEs), analyze it mathematically and provide computer simulations that confirm the analytical results. Our results reproduce field records on population dynamics of P. polytes in the Miyako-jima Island. They also explain the positive dependence of the relative abundance (RA) of the mimic on the advantage index (AI) of the mimicry in the Sakishima Islands defined in Section 2.

Geographical Diversity of Japanese Papilio Butterflies Inferred from the Number of Contact Chemosensillum on the Fifth Foretarsal Segments

Geographical Diversity of Japanese Papilio Butterflies Inferred from the Number of Contact Chemosensillum on the Fifth Foretarsal Segments

Iridescence and nano-structure differences in Papilio butterflies

The nano-photonic structures on the wings of three Papilionidae butterflies, Papilio blumei, Papilio ulysses and Papilio peranthus, were investigated. It was observed that the photonic structure is multi-layer with alternate air and cuticle layers forming one-dimensional photonic crystal. The multi-layer structures of the three butterflies differ subtly but are sufficient to account for the differences in their iridescence. The subtleness is more obvious in their polarized reflection results. We performed the simulation of polarized reflection using characteristic matrix method with parameters obtained from SEM images of butterfly wing scales’ cross-section. The simulated reflection spectra are matched with the experimental spectra to derive the effective refractive index of the air lamina in the butterfly wing scales. It shows that through varying the optical thickness and periodicities in air/cuticle bilayer stacks, the iridescent color of these three Papilionidae butterflies appear different. The result lays the foundation for mimicking the photonic structures of these butterflies.

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

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

Neurons innervating the lamina in the butterfly, Papilio xuthus

The butterfly Papilio xuthus has compound eyes with three types of ommatidia. Each type houses nine spectrally heterogeneous photoreceptors (R1-R9) that are divided into six spectral classes: ultraviolet, violet, blue, green, red, and broad-band. Analysis of color discrimination has shown that P. xuthus uses the ultraviolet, blue, green, and red receptors for foraging. The ultraviolet and blue receptors are long visual fibers terminating in the medulla, whereas the green and red receptors are short visual fibers terminating in the lamina. This suggests that processing of wavelength information begins in the lamina in P. xuthus, unlike in flies. To establish the anatomical basis of color discrimination mechanisms, we examined neurons innervating the lamina by injecting neurobiotin into this neuropil. We found that in addition to photoreceptors and lamina monopolar cells, three distinct groups of cells project fibers into the lamina. Their cell bodies are located (1) at the anterior rim of the medulla, (2) between the proximal surface of the medulla and lobula plate, and (3) in the medulla cell body rind. Neurobiotin injection also labeled distinct terminals in medulla layers 1, 2, 3, 4 and 5. Terminals in layer 4 belong to the long visual fibers (R1, 2 and 9), while arbors in layers 1, 2 and 3 probably correspond to terminals of three subtypes of lamina monopolar cells, respectively. Immunocytochemistry coupled with neurobiotin injection revealed their transmitter candidates; neurons in (1) and a subset of neurons in (2) are immunoreactive to anti-serotonin and anti-γ-aminobutyric acid, respectively.

Structural integration design for enhanced photoluminescence in butterfly scales

Here we present a structural integration design in butterfly scales that helps to enhance the photoluminescence of the scale pigment. The optical mechanism lying behind blue scales of the butterfly Papilio oribazus and green ones from Papilio nireus is illustrated through respective detailed investigation. Microscopic characterization reveals a nano-scaled optical architecture comprised of a network on top of a two-dimensional (2D) photonic crystal slab (PCS) and a multilayer structure located under the PCS. The top side of the PCS is infused with pigments. Combining the experimental reflection spectra and computational simulation through finite difference time domain (FDTD) and plane wave expansion (PWE) methods, we find that the network acts as an antireflection layer harvesting UV excitation light for the pigment, and the 2D PCS works as a light filter and waveguide, functioning together with the multilayer structure to extract needed light from the pigment emission and guide it out of the scales. This finding provides a working prototype to enrich the horizon for the design of sophisticated optical devices with high functionality.

Geographical Diversity of Japanese Papilio Butterflies Inferred from the Number of Contact Chemosensillum on the Fifth Foretarsal Segments

The numbers of contact chemosensillum on ventral side of fifth foretarsal segments of Japanese Papilio butterflies were determined by scanning electron microscope observation. P. bianor, P. helenus, P. protenor and P. memnon exhibited geographical diversity for this character among mainland Japan, Amami Islands, Okinawa Islands and Yaeyama Islands populations. This characteristic was useful for reconfirming the geographical variations proposed by the wing color patterns and genetic characteristics of each these species. This characteristic also uncovered hidden geographical variations and phylogenetic structures of P. machaon, P. xuthus, P. maackii, P. macilentus and that of P. memnon on mainland Japan and that of P. protenor on Okinawa Islands. Also discussed are possible causes for the apparent phylogeny disparities between this report and other previous studies including DNA analyses.

Replication of Papilio maackii Ménétriés butterfly scale structural color using a magnetron sputtering method

The natural butterfly Papilio maackii Ménétriés scales were found, through scanning and transmission electron microscopy, to comprise stacks of thin layers. Replicated scales were then fabricated on a silicon substrate using a magnetron sputtering (MS) method with ITO (In2O3(90%)+SnO2(10%)) and polymethyl methacrylate (PMMA). Finally, the optical effects of the replicated scale were examined by a spectrometer. The results showed that the replicated and natural scales have the similar characteristics in structural colors. As a result, it can be concluded that the MS method reported in this paper is a feasible method for fabricating an artificial biological structural color or other optical elements.

Japanese Papilio butterflies puddle using Na+ detected by contact chemosensilla in the proboscis

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.Electronic supplementary materialThe online version of this article (doi:10.1007/s00114-012-0976-3) contains supplementary material, which is available to authorized users.

Sex chromosome mosaicism and hybrid speciation among tiger swallowtail butterflies.

Hybrid speciation, or the formation of a daughter species due to interbreeding between two parental species, is a potentially important means of diversification, because it generates new forms from existing variation. However, factors responsible for the origin and maintenance of hybrid species are largely unknown. Here we show that the North American butterfly Papilio appalachiensis is a hybrid species, with genomic admixture from Papilio glaucus and Papilio canadensis. Papilio appalachiensis has a mosaic phenotype, which is hypothesized to be the result of combining sex-linked traits from P. glaucus and P. canadensis. We show that P. appalachiensis' Z-linked genes associated with a cooler thermal habitat were inherited from P. canadensis, whereas its W-linked mimicry and mitochondrial DNA were inherited from P. glaucus. Furthermore, genome-wide AFLP markers showed nearly equal contributions from each parental species in the origin of P. appalachiensis, indicating that it formed from a burst of hybridization between the parental species, with little subsequent backcrossing. However, analyses of genetic differentiation, clustering, and polymorphism based on molecular data also showed that P. appalachiensis is genetically distinct from both parental species. Population genetic simulations revealed P. appalachiensis to be much younger than the parental species, with unidirectional gene flow from P. glaucus and P. canadensis into P. appalachiensis. Finally, phylogenetic analyses, combined with ancestral state reconstruction, showed that the two traits that define P. appalachiensis' mosaic phenotype, obligatory pupal diapause and mimicry, evolved uniquely in P. canadensis and P. glaucus, respectively, and were then recombined through hybridization to form P. appalachiensis. These results suggest that natural selection and sex-linked traits may have played an important role in the origin and maintenance of P. appalachiensis as a hybrid species. In particular, ecological barriers associated with a steep thermal cline appear to maintain the distinct, mosaic genome of P. appalachiensis despite contact and occasional hybridization with both parental species.

Intensity contrast as a crucial cue for butterfly landing

Papilio butterflies use a tetrachromatic color vision to discriminate a rewarding flower, approach, land and take nectar from the flower. In the course of further analyzing their foraging behavior in a laboratory condition, we found that some butterflies could not land on the target flower even they discriminated and tried to land on it, especially when the target was dark. This phenomenon, which we call "landing suppression", indicates that the cue for landing differs from the cue for visually locating a flower. We hypothesized that a possible cue for landing was intensity contrast between the target and background, and have initiated to test this hypothesis. We tested the butterflies' landing behavior to targets of various colors and intensities presented on background of black or various densities of gray. As a result, the landing was most strongly suppressed when the intensity contrast was close to zero irrespective of the target colors, suggesting that the butterflies used the target-background intensity contrast when landing.

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

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.

Structural colouration and optical effects in the wings of Papilio peranthus

The butterfly Papilio peranthus displays an iridescent green colour. Through opticalmeasurements, structural characterizations and theoretical analyses, we reveal that thecolour is actually a mixing effect of green and blue which originates from the interiormultilayer structures of scales imbricated in the wings. The chromatic difference betweenthe produced green and blue colour is attributed to the modulations in the butterfly wings.Reflected light by the inclined sides of pits changes its polarization to a perpendiculardirection. Besides, elongated pits lead to anisotropic polarization conversion. A widerangle spread reflection caused by the morphology of pits and the nearly ‘ideal’multilayer structures in scales may be advantageous to conspecific recognition.

Mimicking the colourful wing scale structure of the Papilio blumei butterfly

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.

The mitochondrial genome of the butterfly Papilio xuthus (Lepidoptera: Papilionidae) and related phylogenetic analyses

The nearly complete mitochondrial genome of the butterfly Papilio xuthus (Lepidoptera: Papilionidae) was sequenced for its nucleotide sequence of 13,964 bp. The genome has a typical gene order identical to other lepidopteran species. All tRNAs showed same stable canonical clover-leaf structure as those of other insects, except for tRNA(Ser) (AGN), in which the dihydrouracil arm (DHU arm) could not form stable stem-loop structure. Anomalous initiation codons have been observed for the cox1 gene, where the ATTACG hexa-nucleotide was believed to be involved in the initiation signaling. Twelve mitochondrial protein-coding gene sequence data were used to infer the phylogenetic relationships among the insect orders. Even though the number of insect orders represented by complete mitochondrial genomes is still limited, several well-established relationships are evident in the phylogenetic analysis of the complete sequences. Monophyly of the Homometabola was not supported in this paper. Phylogenetic analyses of the available species of Bombycoidea, Pyraloidea, Papilionoidea and Tortricidea bolstered the current morphology-based hypothesis that Bombycoidea, Pyraloidea and Papilionoidea are monophyletic (Obtectomera). Bombycoidea (Bombyx mandarina and Antheraea pernyi) and Papilionoidea (P. xuthus and Coreana raphaelis) formed a sister group.

Composite organic-inorganic butterfly scales: Production of photonic structures with atomic layer deposition

Recent advances in the photonics and optics industries have produced great demand for ever more sophisticated optical devices, such as photonic crystals. However, photonic crystals are notoriously difficult to manufacture. Increasingly, therefore, researchers have turned towards naturally occurring photonic structures for inspiration and a wide variety of elaborate techniques have been attempted to copy and harness biological processes to manufacture artificial photonic structures. Here, we describe a simple, direct process for producing an artificial photonic device by using a naturally occurring structure from the wings of the butterfly Papilio blumei as a template and low-temperature atomic layer deposition of TiO2 to create a faithful cast of the structure. The optical properties of the organic-inorganic diffraction structures produced are assessed by normal-incidence specular reflectance and found to be well described by multilayer computation method using a two-dimensional photonic crystal model. Depending on the structural integrity of the initially sealed scale, it was found possible not only to replicate the outer but also the inner and more complex surfaces of the structure, each resulting in distinct multicolor optical behavior as revealed by experimental and theoretical data. In this paper, we also explore tailoring the process to design composite skeleton architectures with desired optical properties and integrated multifunctional (mechanical, thermal, optical, fluidic) properties.

Blue and Double-peaked Green Receptors Depend on Ommatidial Type in the Eye of the Japanese Yellow Swallowtail Papilio xuthus

The compound eye of the butterfly Papilio xuthus is composed of three spectrally distinct types of ommatidia. We investigated the blue and double-peaked green receptors that are encountered distally in type I and III ommatidia, by means of intracellular recordings, in vivo fluorescence microscopy, and histology. The blue receptors are R1 and/or R2 photoreceptors; they contain the same mRNA encoding the opsin of the blue-absorbing visual pigment. However, here we found that the sensitivity in the UV wavelength region strongly depends on the ommatidial type; the blue receptors in type I ommatidia have a distinctly depressed UV sensitivity, which is attributed to lateral filtering in the fused rhabdom. In the main, fronto-ventral part of the eye, the R3 and R4 photoreceptors of all ommatidia contain the same set of two mRNAs encoding the opsins of green-absorbing visual pigments, PxL1 and PxL2. The spectral sensitivities are double-peaked, but the UV sensitivity of the R3 and R4 photoreceptors in type I ommatidia appears to be reduced, similar to that of the co-localized blue receptors.

Morphology of Foretarsal Ventral Surfaces of Japanese Papilio Butterflies and Relations Between These Morphology, Phylogeny and Hostplant Preferring Hierarchy

Foretarsal ventral surface observation of Japanese Papilio butterflies showed that the shapes of fifth foretarsi and numbers and localization of contact chemosensilla and spines in these areas are closely related to both phylogeny and behavior in these species. My results basically supported the classification that Japanese Papilio divides into five subgenera -- Papilio (P. machaon), Princeps (P. xuthus and P. demoleus), Achillides (P. maackii and P. bianor), Menelaides (P. helenus, P. polytes, P. protenor and P. macilentus) and Iliades (P. memnon). Moreover, female foretarsal morphology also corresponded to the physical features of their preferring host plant leaves. The specificity of female P. machaon, female P. macilentus and female P. maackii seemed to also relate to the geographical diversities of these species and their hostplant, and their co-evolution.

How do foraging Papilio butterflies see polarized light

How do foraging Papilio butterflies see polarized light

Ommatidial type‐specific interphotoreceptor connections in the lamina of the swallowtail butterfly, Papilio xuthus

The eye of the butterfly Papilio xuthus contains a random array of three types of ommatidia (types I-III), each bearing nine photoreceptors, R1-R9. Of the six spectral classes of photoreceptors identified, types I, II, and III ommatidia contain four, three, and two classes, respectively: the ommatidia are thus spectrally heterogeneous. The photoreceptors send their axons to the lamina where, together with some large monopolar cells (LMCs), the nine from a single ommatidium contribute to a module called a lamina cartridge. We recently reported that among different photoreceptor axon terminals visualized by confocal microscopy, the number and length of axon collaterals differ for different spectral receptors, suggesting that lamina circuits are specific for each ommatidial type. Here we studied the distribution of synapse-like structures in the cartridges, first characterizing a photoreceptor by measuring its spectral sensitivity and then injecting Lucifer yellow (LY). We subsequently histologically identified the type of ommatidium to which the injected photoreceptor belonged, cut serial ultrathin sections of the entire lamina, labeled these with anti-LY immunocytochemistry, and then localized synapse-like structures. We found numerous interphotoreceptor contacts both within and between cartridges, the combination of which was again specific for the ommatidial type. R3 and R4, which are green-sensitive photoreceptors in all ommatidia, have thick axons lacking collaterals. We found that these cells exclusively make contacts with LMCs and not with photoreceptors. We therefore presume that R3 and R4 construct a system for motion vision, whereas other randomly distributed spectral types provide inputs for color vision.