Blue reflectance in tarantulas is evolutionarily conserved despite nanostructural diversity

Slight shifts in arrangement within biological photonic nanostructures can produce large color differences, and sexual selection often leads to high color diversity in clades with structural colors. We use phylogenetic reconstruction, electron microscopy, spectrophotometry, and opticalmodeling to showan opposing pattern of nanostructural diversification accompanied by unusual conservation of blue color in tarantulas (Araneae: Theraphosidae). In contrast to other clades, blue coloration in phylogenetically distant tarantulas peaks within a narrow 20-nm region around 450 nm. Both quasi-ordered and multilayer nanostructures found in different tarantulas produce this blue color. Thus, even within monophyletic lineages, tarantulas have evolved strikingly similar blue coloration through divergent mechanisms. The poor color perception and lack of conspicuous display during courtship of tarantulas argue that these colors are not sexually selected. Therefore, our data contrast with sexual selection that typically produces a diverse array of colors with a single structuralmechanismby showing that natural selection on structural color in tarantulas resulted in convergence on similar color through diverse structural mechanisms

Spiders have rich pigmentary and structural colour palettes

Elucidating the mechanisms of colour production in organisms is important for understanding how selection acts upon a variety of behaviours. Spiders provide many spectacular examples of colours used in courtship, predation, defence and thermoregulation, but are thought to lack many types of pigments common in other animals. Ommochromes, bilins and eumelanin have been identified in spiders, but not carotenoids or melanosomes. Here, we combined optical microscopy, refractive index matching, confocal Raman microspectroscopy and electron microscopy to investigate the basis of several types of colourful patches in spiders. We obtained four major results. First, we show that spiders use carotenoids to produce yellow, suggesting that such colours may be used for condition-dependent courtship signalling. Second, we established the Raman signature spectrum for ommochromes, facilitating the identification of ommochromes in a variety of organisms in the future. Third, we describe a potential new pigmentary-structural colour interaction that is unusual because of the use of long wavelength structural colour in combination with a slightly shorter wavelength pigment in the production of red. Finally, we present the first evidence for the presence of melanosomes in arthropods, using both scanning and transmission electron microscopy, overturning the assumption that melanosomes are a synapomorphy of vertebrates. Our research shows that spiders have a much richer colour production palette than previously thought, and this has implications for colour diversification and function in spiders and other arthropods

Rainbow peacock spiders inspire miniature super-iridescent optics

Colour produced by wavelength-dependent light scattering is a key component of visual communication in nature and acts particularly strongly in visual signalling by structurally-coloured animals during courtship. Two miniature peacock spiders (Maratus robinsoni and M. chrysomelas) court females using tiny structured scales (similar to 40 x 10 mu m(2)) that reflect the full visual spectrum. Using TEM and optical modelling, we show that the spiders' scales have 2D nanogratings on microscale 3D convex surfaces with at least twice the resolving power of a conventional 2D diffraction grating of the same period. Whereas the long optical path lengths required for light-dispersive components to resolve individual wavelengths constrain current spectrometers to bulky sizes, our nano-3D printed prototypes demonstrate that the design principle of the peacock spiders' scales could inspire novel, miniature light-dispersive components

Rainbow peacock spiders inspire miniature superiridescent optics

Colour produced by wavelength-dependent light scattering is a key component of visual communication in nature and acts particularly strongly in visual signalling by structurallycoloured animals during courtship. Two miniature peacock spiders (Maratus robinsoni and M. chrysomelas) court females using tiny structured scales (~ 40 × 10 μm2) that reflect the full visual spectrum. Using TEM and optical modelling, we show that the spiders’ scales have 2D nanogratings on microscale 3D convex surfaces with at least twice the resolving power of a conventional 2D diffraction grating of the same period. Whereas the long optical path lengths required for light-dispersive components to resolve individual wavelengths constrain current spectrometers to bulky sizes, our nano-3D printed prototypes demonstrate that the design principle of the peacock spiders’ scales could inspire novel, miniature light-dispersive components

Rainbow peacock spiders inspire miniature super-iridescent optics

Colour produced by wavelength-dependent light scattering is a key component of visual communication in nature and acts particularly strongly in visual signalling by structurally-coloured animals during courtship. Two miniature peacock spiders (Maratus robinsoni and M. chrysomelas) court females using tiny structured scales (~ 40 × 10 μm^2) that reflect the full visual spectrum. Using TEM and optical modelling, we show that the spiders’ scales have 2D nanogratings on microscale 3D convex surfaces with at least twice the resolving power of a conventional 2D diffraction grating of the same period. Whereas the long optical path lengths required for light-dispersive components to resolve individual wavelengths constrain current spectrometers to bulky sizes, our nano-3D printed prototypes demonstrate that the design principle of the peacock spiders’ scales could inspire novel, miniature light-dispersive components

Spiders do have melanin after all

Melanin pigments are broadly distributed in nature - from bacteria to fungi to plants and animals. However, many previous attempts to identify melanins in spiders were unsuccessful, suggesting that these otherwise ubiquitous pigments were lost during spider evolution. Yet, spiders exhibit many dark colours similar to those produced by melanins in other organisms, and the low solubility of melanins makes isolation and characterization difficult. Therefore, whether melanins are truly absent or have simply not yet been detected is an open question. Raman spectroscopy provides a reliable way to detect melanins in situ, without the need for isolation. In this study, we document the presence of eumelanin in diverse species of spiders using confocal Raman microspectroscopy. Comparisons of spectra with theoretically calculated data falsify the previous hypothesis that dark colours are produced solely by ommochromes in spiders. Our data indicate that melanins are present in spiders and further supporting that they are present in most living organisms

Color production mechanisms in spiders

Spiders were once thought to have limited color production palettes, and even to lack melanin - one of the most ubiquitous biological pigments. Recent discoveries upend that view and show that the color production mechanisms in spiders are as elaborate as some of the more classically colorful groups of animals, such as birds, butterflies, and beetles. Here we summarize how colors are produced by spiders, identify gaps in our knowledge, show how researchers investigating color in different groups of animals can learn from each other, and suggest future opportunities for spider color research. Our understanding of color production mechanisms in other colorful groups of animals can be used as guidelines for discovering existing mechanisms previously unknown in spiders and vice versa. For example, spider species with colors potentially produced by the same kind of photonic structures previously described in white beetles, and in the blue/green scales of fishes and lizards are suggested here. In addition, novel principles first found in spiders that modify the iridescence of structural colors via the interaction of structural features across different length scales (i.e., micro-nano) may also be found in other colorful groups in the future. This review summarizes the state-of-the-art understanding regarding the proximate color production mechanisms in spiders, suggests a few future research directions that are likely to be fruitful, and facilitates the advancements in related fields, including the ecology, evolution, and functions of spider coloration

Structural color and its interaction with other color-producing elements : perspectives from spiders

Structural color is produced when nanostructures alter light in contrast with pigment-based colors that are produced by selective absorption of certain wavelengths of light. Research on biogenic photonic nanostructures has primarily focused on bird feathers, butterfly wings and beetle elytra, and not 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 some spiders 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, a hugely diverse set of colors is produced using the same materials. Using spiders, we can study how colors evolve to serve different functions under a variety of selective pressures, 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? Addressing any of these questions in the future will bring spiders to the forefront of the study of structural colors in nature