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What drives the evolution of plumage color in birds? Bird color is likely to be under both natural and sexual selection where natural selection may favor evolution toward crypsis or camouflage whereas sexual selection may favor evolution toward conspicuousness. The responses to selection are predicted to relate to species’ ecology, behavior, and life history. Key hypotheses have focused on habitat and light environment, breeding strategy, territoriality, and hunting behavior. We tested these potential causes of color variation in the Coraciiformes, a colorful clade of non-passerine birds, using phylogenetic comparative methods and data on chromatic and achromatic properties of plumage coloration measured from museum specimens. We found that correlates of color evolution in Coraciiformes vary across body regions and depend on the focal color property (chromatic or achromatic properties of plumage coloration). While the light environment showed widespread effects on coloration in multiple body regions for both color properties, selection pressures related to behavioral characteristics had more spatially localized effects (e.g. territoriality on achromatic properties of wing feathers and hunting strategy on chromatic properties of belly feathers). Our results reveal both general patterns that may hold across other bird clades and more nuanced effects of selection that are likely to be mediated through the visual ecology of the signaler and receiver and the behavioral characteristics of Coraciiform species.

Tobias, Joseph A., Sheard, Catherine, Pigot, Alex L., Devenish, Adam J. M., Yang, Jingyi, Sayol, Ferran, Neate‐Clegg, Montague H. C., Alioravainen, Nico, Weeks, Thomas L., Barber, Robert A., Walkden, Patrick A., MacGregor, Hannah E. A., Jones, Samuel E. I., Vincent, Claire, Phillips, Anna G., Marples, Nicola M., Montaño‐Centellas, Flavia A., Leandro‐Silva, Victor, Claramunt, Santiago, Darski, Bianca, Freeman, Benjamin G., Bregman, Tom P., Cooney, Christopher R., Hughes, Emma C., Capp, Elliot J. R., Varley, Zoë K., Friedman, Nicholas R., Korntheuer, Heiko, Corrales‐Vargas, Andrea, Trisos, Christopher H., Weeks, Brian C., Hanz, Dagmar M., Töpfer, Till, Bravo, Gustavo A., Remeš, Vladimír, Nowak, Larissa, Carneiro, Lincoln S., Moncada R., Amilkar J., Matysioková, Beata, Baldassarre, Daniel T., Martínez‐Salinas, Alejandra, Wolfe, Jared D., Chapman, Philip M., Daly, Benjamin G., Sorensen, Marjorie C., Neu, Alexander, Ford, Michael A., Mayhew, Rebekah J., Fabio Silveira, Luis, Kelly, David J., Annorbah, Nathaniel N. D., Pollock, Henry S., Grabowska‐Zhang, Ada M., McEntee, Jay P., Carlos T. Gonzalez, Juan, Meneses, Camila G., Muñoz, Marcia C., Powell, Luke L., Jamie, Gabriel A., Matthews, Thomas J., Johnson, Oscar, Brito, Guilherme R. R., Zyskowski, Kristof, Crates, Ross, Harvey, Michael G., Jurado Zevallos, Maura, Hosner, Peter A., Bradfer‐Lawrence, Tom, Maley, James M., Stiles, F. Gary, Lima, Hevana S., Provost, Kaiya L., Chibesa, Moses, Mashao, Mmatjie, Howard, Jeffrey T., Mlamba, Edson, Chua, Marcus A. H., Li, Bicheng, Gómez, M. Isabel, García, Natalia C., Päckert, Martin, Fuchs, Jérôme, Ali, Jarome R., Derryberry, Elizabeth P., Carlson, Monica L., Urriza, Rolly C., Brzeski, Kristin E., Prawiradilaga, Dewi M., Rayner, Matt J., Miller, Eliot T., Bowie, Rauri C. K., Lafontaine, René‐Marie, Scofield, R. Paul, Lou, Yingqiang, Somarathna, Lankani, Lepage, Denis, Illif, Marshall, Neuschulz, Eike Lena, Templin, Mathias, Dehling, D. Matthias, Cooper, Jacob C., Pauwels, Olivier S. G., Analuddin, Kangkuso, Fjeldså, Jon, Seddon, Nathalie, Sweet, Paul R., DeClerck, Fabrice A. J., Naka, Luciano N., Brawn, Jeffrey D., Aleixo, Alexandre, Böhning‐Gaese, Katrin, Rahbek, Carsten, Fritz, Susanne A., Thomas, Gavin H., Schleuning, Matthias (2022): AVONET: morphological, ecological and geographical data for all birds. Ecology Letters 25 (3): 581-597, DOI: 10.1111/ele.13898, URL: http://dx.doi.org/10.1111/ele.13898

Trapliners are pollinators that visit widely dispersed flowers along circuitous foraging routes. The evolution of traplining in hummingbirds is thought to entail morphological specialization through the reciprocal coevolution of longer bills with the long-tubed flowers of widely dispersed plant species. Specialization, such as that exhibited by traplining hummingbirds, is often viewed as both irreversible and an evolutionary dead end. We tested these predictions in a macroevolutionary framework. Specifically, we assessed the relationship between beak morphology and foraging and tested whether transitions to traplining are irreversible and lead to lower rates of diversification as predicted by the hypothesis that specialization is an evolutionary dead end. We find that there have been multiple independent transitions to traplining across the hummingbird phylogeny, but reversals have been rare or incomplete at best. Multiple independent lineages of trapliners have become morphologically specialized, convergently evolving relatively large bills for their body size. Traplining is not an evolutionary dead end however, since trapliners continue to give rise to new traplining species at a rate comparable to non-trapliners.

Evolution can involve periods of rapid divergent adaptation and expansion in the range of diversity, but evolution can also be relatively conservative over certain timescales due to functional, genetic-developmental, and ecological constraints. One way in which evolution may be conservative is in terms of allometry, the scaling relationship between the traits of organisms and body size. Here, we investigate patterns of allometric conservatism in the evolution of bird beaks with beak size and body size data for a representative sample of over 5000 extant bird species within a phylogenetic framework. We identify clades in which the allometric relationship between beak size and body size has remained relatively conserved across species over millions to tens of millions of years. We find that allometric conservatism is nonetheless punctuated by occasional shifts in the slopes and intercepts of allometric relationships. A steady accumulation of such shifts through time has given rise to the tremendous diversity of beak size relative to body size across birds today. Our findings are consistent with the Simpsonian vision of macroevolution, with evolutionary conservatism being the rule but with occasional shifts to new adaptive zones.

Understanding the biogeographical patterns, and evolutionary and environmental drivers, underpinning morphological diversity are key for determining its origins and conservation. Using a comprehensive set of continuous morphological traits extracted from museum collections of 8353 bird species, including geometric morphometric beak shape data, we find that avian morphological diversity is unevenly distributed globally, even after controlling for species richness, with exceptionally dense packing of species in hyper-diverse tropical hotspots. At the regional level, these areas also have high morphological variance, with species exhibiting high phenotypic diversity. Evolutionary history likely plays a key role in shaping these patterns, with evolutionarily old species contributing to niche expansion, and young species contributing to niche packing. Taken together, these results imply that the tropics are both 'cradles' and 'museums' of phenotypic diversity.

It has long been suggested that tropical species are generally more colourful than temperate species, but whether latitudinal gradients in organismal colourfulness exist remains controversial. Here we quantify global latitudinal trends in colourfulness (within-individual colour diversity) by collating and analysing a photographic dataset of whole-body plumage reflectance information for4,500 species of passerine birds. We show that male and female birds of tropical passerine species are generally more colourful than their temperate counterparts, both on average and in the extreme. We also show that these geographic gradients can be explained in part by the effects of several latitude-related factors related to classic hypotheses for climatic and ecological determinants of organismal colourfulness. Taken together, our results reveal that species' colourfulness peaks in the tropics for passerine birds, confirming the existence of a long-suspected yet hitherto elusive trend in the distribution of global biodiversity.

Ultraviolet (UV) colouration is thought to be an important signalling mechanism in many bird species, yet broad insights regarding the prevalence of UV plumage colouration and the factors promoting its evolution are currently lacking. Here, we develop a novel image segmentation pipeline based on deep learning that considerably outperforms classical (i.e. non-deep learning) segmentation methods, and use this to extract accurate information on whole-body plumage colouration from photographs of >24,000 museum specimens covering >4,500 species of passerine birds. Our results demonstrate that UV reflectance, particularly as a component of other colours, is widespread across the passerine radiation but is strongly phylogenetically conserved. We also find clear evidence in support of the role of light environment in promoting the evolution of UV plumage colouration, and a weak trend towards higher UV plumage reflectance among bird species with ultraviolet rather than violet-sensitive visual systems. Overall, our study provides important broad-scale insight into an enigmatic component of avian colouration, as well as demonstrating that deep learning has considerable promise for allowing new data to be bought to bear on long-standing questions in ecology and evolution.

Sexual selection is proposed to be a powerful driver of phenotypic evolution in animal systems. At macroevolutionary scales, sexual selection can theoretically drive both the rate and direction of phenotypic evolution, but this hypothesis remains contentious. Here, we find that differences in the rate and direction of plumage colour evolution are predicted by a proxy for sexual selection intensity (plumage dichromatism) in a large radiation of suboscine passerine birds (Tyrannida). We show that rates of plumage evolution are correlated between the sexes, but that sexual selection has a strong positive effect on male, but not female, interspecific divergence rates. Furthermore, we demonstrate that rapid male plumage divergence is biased towards carotenoid-based (red/yellow) colours widely assumed to represent honest sexual signals. Our results highlight the central role of sexual selection in driving avian colour divergence, and reveal the existence of convergent evolutionary responses of animal signalling traits under sexual selection., What factors explain variation in the pace and trajectory of evolutionary divergence between lineages? Here, the authors show that a proxy measure for sexual selection intensity predicts both the rate and direction of plumage colour evolution in a diverse radiation of New World passerine birds.

Competition for shared resources represents a fundamental driver of biological diversity. However, the tempo and mode of phenotypic evolution in deep-time has been predominantly investigated using trait evolutionary models which assume that lineages evolve independently from each other. Consequently, the role of species interactions in driving macroevolutionary dynamics remains poorly understood. Here, we quantify the prevalence for signatures of competition between related species in the evolution of ecomorphological traits across the bird radiation. We find that mechanistic trait models accounting for the effect of species interactions on phenotypic divergence provide the best fit for the data on at least one trait axis in 27 out of 59 clades ranging between 21 and 195 species. Where it occurs, the signature of competition generally coincides with positive species diversity-dependence, driven by the accumulation of lineages with similar ecologies, and we find scarce evidence for trait-dependent or negative diversity-dependent phenotypic evolution. Overall, our results suggest that the footprint of interspecific competition is often eroded in long-term patterns of phenotypic diversification, and that other selection pressures may predominantly shape ecomorphological diversity among extant species at macroevolutionary scales.

Heterogeneity in rates of trait evolution is widespread, but it remains unclear which processes drive fast and slow character divergence across global radiations. Here, we test multiple hypotheses for explaining rate variation in an ecomorphological trait (beak shape) across a globally distributed group (birds). We find low support that variation in evolutionary rates of species is correlated with life history, environmental mutagenic factors, range size, number of competitors, or living on islands. Indeed, after controlling for the negative effect of species' age, 80% of variation in species‐specific evolutionary rates remains unexplained. At the clade level, high evolutionary rates are associated with unusual phenotypes or high species richness. Taken together, these results imply that macroevolutionary rates of ecomorphological traits are governed by both ecological opportunity in distinct adaptive zones and niche differentiation among closely related species.

The origin and expansion of biological diversity is regulated by both developmental trajectories and limits on available ecological niches. As lineages diversify, an early and often rapid phase of species and trait proliferation gives way to evolutionary slow-downs as new species pack into ever more densely occupied regions of ecological niche space. Small clades such as Darwin's finches demonstrate that natural selection is the driving force of adaptive radiations, but how microevolutionary processes scale up to shape the expansion of phenotypic diversity over much longer evolutionary timescales is unclear. Here we address this problem on a global scale by analysing a crowdsourced dataset of three-dimensional scanned bill morphology from more than 2,000 species. We find that bill diversity expanded early in extant avian evolutionary history, before transitioning to a phase dominated by packing of morphological space. However, this early phenotypic diversification is decoupled from temporal variation in evolutionary rate: rates of bill evolution vary among lineages but are comparatively stable through time. We find that rare, but major, discontinuities in phenotype emerge from rapid increases in rate along single branches, sometimes leading to depauperate clades with unusual bill morphologies. Despite these jumps between groups, the major axes of within-group bill-shape evolution are remarkably consistent across birds. We reveal that macroevolutionary processes underlying global-scale adaptive radiations support Darwinian and Simpsonian ideas of microevolution within adaptive zones and accelerated evolution between distinct adaptive peaks.

Nature 542, 344–347 (2017); doi:10.1038/nature21074 In this Letter, two files were inadvertently omitted from the Supplementary Information. The two files (‘Prum_merge_taxonomy_CRC_v2.csv’ and ‘PrumTreeMerge_CRC_v1.csv’) are needed to replicate the alternative phylogeny of Prum et al. (ref. 43) presented in Extended Data Figure 4.