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Structural color is poorly known in plants relative to animals. In fruits, only a handful of cases have been described, including in Viburnum tinus where the blue color results from a disordered multilayered reflector made of lipid droplets. Here, we examine the broader evolutionary context of fruit structural color across the genus Viburnum. We obtained fresh and herbarium fruit material from 30 Viburnum species spanning the phylogeny and used transmission electron microscopy, optical simulations, and ancestral state reconstruction to identify the presence/absence of photonic structures in each species, understand the mechanism producing structural color in newly identified species, relate the development of cell wall structure to reflectance in Viburnum dentatum, and describe the evolution of cell wall architecture across Viburnum. We identify at least two (possibly three) origins of blue fruit color in Viburnum in species which produce large photonic structures made of lipid droplets embedded in the cell wall and which reflect blue light. Examining the full spectrum of mechanisms producing color in pl, including structural color as well as pigments, will yield further insights into the diversity, ecology, and evolution of fruit color.

The majority of plant colours are produced by anthocyanin and carotenoid pigments, but colouration obtained by nanostructured materials (i.e. structural colours) is increasingly reported in plants. Here, we identify a multilayer photonic structure in the fruits of Lantana strigocamara and compare it with a similar structure in Viburnum tinus fruits. We used a combination of transmission electron microscopy (EM), serial EM tomography, scanning force microscopy and optical simulations to characterise the photonic structure in L. strigocamara. We also examine the development of the structure during maturation. We found that the structural colour derives from a disordered, multilayered reflector consisting of lipid droplets of c.105 nm that form a plate-like structure in 3D. This structure begins to form early in development and reflects blue wavelengths of light with increasing intensity over time as the structure develops. The materials used are likely to be lipid polymers. Lantana strigocamara is the second origin of a lipid-based photonic structure, convergently evolved with the structure in Viburnum tinus. Chemical differences between the lipids in L. strigocamara and those of V. tinus suggest a distinct evolutionary trajectory with implications for the signalling function of structural colours in fruits.

Funder: National Park Foundation; Id: http://dx.doi.org/10.13039/100001261, Funder: National Council for Scientific Research and Techniques, Funder: Mt Cuba Center, Syndromes, wherein multiple traits evolve convergently in response to a shared selective driver, form a central concept in ecology and evolution. Recent work has questioned the existence of some classic syndromes, such as pollination and seed dispersal syndromes. Here, we discuss some of the major issues that have afflicted research into syndromes in macroevolution and ecology. First, correlated evolution of traits and hypothesized selective drivers is often relied on as the only evidence for adaptation of those traits to those hypothesized drivers, without supporting evidence. Second, the selective driver is often inferred from a combination of traits without explicit testing. Third, researchers often measure traits that are easy for humans to observe rather than measuring traits that are suited to testing the hypothesis of adaptation. Finally, species are often chosen for study because of their striking phenotypes, which leads to the illusion of syndromes and divergence. We argue that these issues can be avoided by combining studies of trait variation across entire clades or communities with explicit tests of adaptive hypotheses and that taking this approach will lead to a better understanding of syndrome-like evolution and its drivers.

Additional file 1: Table S1. Nutritional, morphological and (categorical) color data for Viburnum species, as well as accession numbers and sample sizes for each measurement. Table S2. Step cost matrix implemented in Mesquite to represent the paedomorphy model. Table S3. Transition rate matrices used in the ancestral state reconstruction by maximum likelihood. Table S4. Transition rates between each fruit color state as estimated by make.simmap in phytools using a symmetrical model of evolution.. Figure. S1. Species’ colors plotted in tetrahedral color space according to a visual model of a UV-sensitive bird. Figure S2. Volume overlap between each fruit color category in tetrahedral color space.