• Jablonski, D., Roy, K. & Valentine, J. W. Out of the tropics: evolutionary dynamics of the latitudinal diversity gradient. Science 314, 102–106 (2006).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Weir, J. T. & Schluter, D. The latitudinal gradient in recent speciation and extinction rates of birds and mammals. Science 315, 1574–1576 (2007).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Mittelbach, G. G. et al. Evolution and the latitudinal diversity gradient: speciation, extinction and biogeography. Ecol. Lett. 10, 315–331 (2007).

    Article 
    PubMed 

    Google Scholar
     

  • Simpson, G. G. The Major Features of Evolution (Columbia Univ. Press, 1953).

  • Simpson, G. G. Tempo and Mode in Evolution (Columbia Univ. Press, 1944).

  • Schluter, D. The Ecology of Adaptive Radiation (OUP Oxford, 2000).

  • Schluter, D. Speciation, ecological opportunity, and latitude. Am. Nat. 187, 1–18 (2016).

    Article 
    PubMed 

    Google Scholar
     

  • Quintero, I. The diffused evolutionary dynamics of morphological novelty. Proc. Natl Acad. Sci. USA 122, e2425573122 (2025).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Saupe, E. E. et al. Extinction intensity during Ordovician and Cenozoic glaciations explained by cooling and palaeogeography. Nat. Geosci. 13, 65–70 (2020).

    Article 

    Google Scholar
     

  • Eiserhardt, W. L., Borchsenius, F., Plum, C. M., Ordonez, A. & Svenning, J.-C. Climate-driven extinctions shape the phylogenetic structure of temperate tree floras. Ecol. Lett. 18, 263–272 (2015).

    Article 
    PubMed 

    Google Scholar
     

  • Quintero, I. & Wiens, J. J. Rates of projected climate change dramatically exceed past rates of climatic niche evolution among vertebrate species. Ecol. Lett. 16, 1095–1103 (2013).

    Article 
    PubMed 

    Google Scholar
     

  • Weeks, B. C. et al. Skeletal trait measurements for thousands of bird species. Sci. Data 12, 884 (2025).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Weeks, B. C. et al. A deep neural network for high-throughput measurement of functional traits on museum skeletal specimens. Methods Ecol. Evol. 14, 347–359 (2023).

    Article 

    Google Scholar
     

  • Oliveros, C. H. et al. Earth history and the passerine superradiation. Proc. Natl Acad. Sci. USA 116, 7916–7925 (2019).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Mayr, G. Paleogene Fossil Birds 2nd edn (Springer Cham, 2022).

  • Lowi-Merri, T. M., Gjevori, M., Bochenski, Z. M., Wertz, K. & Claramunt, S. Total-evidence dating and the phylogenetic affinities of early fossil passerines. J. Syst. Paleontol. 22, 2356086 (2024).

    Article 

    Google Scholar
     

  • Steell, E. M., Nguyen, J. M. T., Benson, R. B. J. & Field, D. J. Comparative anatomy of the passerine carpometacarpus helps illuminate the early fossil record of crown Passeriformes. J. Anat. 242, 495–509 (2023).

    Article 
    PubMed 

    Google Scholar
     

  • Claramunt, S. et al. A new time tree of birds reveals the interplay between dispersal, geographic range size, and diversification. Curr. Biol. 35, 1–13 (2025).

    Article 

    Google Scholar
     

  • Stiller, J. et al. Complexity of avian evolution revealed by family-level genomes. Nature 629, 851–860 (2024).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Grant, P. R. & Grant, B. R. Evolution of character displacement in Darwin’s finches. Science 313, 224–226 (2006).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Prum, R. O. The Evolution of Beauty: How Darwin’s Forgotten Theory of Mate Choice Shapes the Animal World-and Us (Anchor, 2018).

  • Ricklefs, R. E. Species richness and morphological diversity of passerine birds. Proc. Natl Acad. Sci. USA 109, 14482–14487 (2012).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Price, T. D. et al. Niche filling slows the diversification of Himalayan songbirds. Nature 509, 222–225 (2014).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Claramunt, S. & Cracraft, J. A new time tree reveals Earth history’s imprint on the evolution of modern birds. Sci. Adv. 1, e1501005 (2015).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Navalón, G., Bjarnason, A., Griffiths, E. & Benson, R. B. J. Environmental signal in the evolutionary diversification of bird skeletons. Nature 611, 306–311 (2022).

    Article 
    PubMed 

    Google Scholar
     

  • Brinkworth, A. et al. Bird clades with less complex appendicular skeletons tend to have higher species richness. Nat. Commun. 14, 5817 (2023).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Chitwood, D. H. et al. Latent developmental and evolutionary shapes embedded within the grapevine leaf. New Phytol. 210, 343–355 (2016).

    Article 
    PubMed 

    Google Scholar
     

  • Klingenberg, C. P. Size, shape, and form: concepts of allometry in geometric morphometrics. Dev. Genes Evol. 226, 113–137 (2016).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Wagner, G. P. Homology, Genes, and Evolutionary Innovation (Princeton Univ. Press, 2014).

  • Pavlicev, M. & Hansen, T. F. Genotype-phenotype maps maximizing evolvability: modularity revisited. Evol. Biol. 38, 371–389 (2011).

    Article 

    Google Scholar
     

  • Cavalli-Sforza, L. L. & Edwards, A. W. F. Phylogenetic analysis: models and estimation procedures. Evolution 21, 550–570 (1967).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Lande, R. Natural selection and random genetic drift in phenotypic evolution. Evolution 30, 314–334 (1976).

    Article 
    PubMed 

    Google Scholar
     

  • Felsenstein, J. Phylogenies and quantitative characters. Annu. Rev. Ecol. Syst. 19, 445–471 (1988).

    Article 

    Google Scholar
     

  • Clavel, J., Aristide, L. & Morlon, H. A penalized likelihood framework for high-dimensional phylogenetic comparative methods and an application to new-world monkeys brain evolution. Syst. Biol. 68, 93–116 (2019).

    Article 
    PubMed 

    Google Scholar
     

  • Clavel, J., Escarguel, G. & Merceron, G. mvMORPH: an R package for fitting multivariate evolutionary models to morphometric data. Methods Ecol. Evol. 6, 1311–1319 (2015).

    Article 

    Google Scholar
     

  • Berv, J. S. et al. Genome and life-history evolution link bird diversification to the end-Cretaceous mass extinction. Sci. Adv. 10, eadp0114 (2024).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Smith, S. A., Walker-Hale, N. & Parins-Fukuchi, C. T. Compositional shifts associated with major evolutionary transitions in plants. New Phytol. 239, 2404–2415 (2023).

    Article 
    PubMed 

    Google Scholar
     

  • Mitov, V., Bartoszek, K. & Stadler, T. Automatic generation of evolutionary hypotheses using mixed Gaussian phylogenetic models. Proc. Natl Acad. Sci. USA 116, 16921–16926 (2019).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Coles, S. An Introduction to Statistical Modeling of Extreme Values (Springer, 2001).

  • Gould, S. J. & Eldredge, N. Punctuated equilibrium comes of age. Nature 366, 223–227 (1993).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Uyeda, J. C., Hansen, T. F., Arnold, S. J. & Pienaar, J. The million-year wait for macroevolutionary bursts. Proc. Natl Acad. Sci. USA 108, 15908–15913 (2011).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Landis, M. J. & Schraiber, J. G. Pulsed evolution shaped modern vertebrate body sizes. Proc. Natl Acad. Sci. USA 114, 13224–13229 (2017).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Wagner, G. P. & Altenberg, L. Perspective: complex adaptations and the evolution of evolvability. Evolution 50, 967–976 (1996).

    Article 
    PubMed 

    Google Scholar
     

  • Wagner, P. J. Early bursts of disparity and the reorganization of character integration. Proc. R. Soc. B 285, 20181604 (2018).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Gatesy, S. M. & Dial, K. P. Locomotor modules and the evolution of avian flight. Evolution 50, 331–340 (1996).

    Article 
    PubMed 

    Google Scholar
     

  • Losos, J. B. Adaptive radiation, ecological opportunity, and evolutionary determinism. Am. Nat. 175, 623–639 (2010).

    Article 
    PubMed 

    Google Scholar
     

  • Eldredge, N. & Gould, S. J. in Models in Paleobiology (ed. Schopf, T. J. M.) 82–115 (Freeman, Cooper and Company, 1972).

  • Malmgren, B. A., Berggren, W. A. & Lohmann, G. P. Evidence for punctuated gradualism in the Late Neogene Globorotalia tumida lineage of planktonic foraminifera. Paleobiology 9, 377–389 (1983).

    Article 

    Google Scholar
     

  • Brennan, I. G., Chapple, D. G., Keogh, J. S. & Donnellan, S. Evolutionary bursts drive morphological novelty in the world’s largest skinks. Curr. Biol. 34, 3905–3916.e3905 (2024).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Dufour, P., Sayol, F., Steinbauer, M. J., Cooke, R. & Faurby, S. Niche filling predicts evolutionary trajectories in insular bird communities. Funct. Ecol. 38, 2636–2647 (2024).

    Article 
    CAS 

    Google Scholar
     

  • Rabosky, D. L. & Lovette, I. J. Density-dependent diversification in North American wood warblers. Proc. R. Soc. B 275, 2363–2371 (2008).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Cooney, C. R. et al. Mega-evolutionary dynamics of the adaptive radiation of birds. Nature 542, 344–347 (2017).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Jønsson, K. A. et al. Ecological and evolutionary determinants for the adaptive radiation of the Madagascan vangas. Proc. Natl Acad. Sci. USA 109, 6620–6625 (2012).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Vinciguerra, N. T. & Burns, K. J. Species diversification and ecomorphological evolution in the radiation of tanagers (Passeriformes: Thraupidae). Biol. J. Linn. Soc. 133, 920–930 (2021).

    Article 

    Google Scholar
     

  • Harmon, L. J. et al. Early bursts of body size and shape evolution are rare in comparative data. Evolution 64, 2385–2396 (2010).

    PubMed 

    Google Scholar
     

  • Jetz, W., Thomas, G. H., Joy, J. B., Hartmann, K. & Mooers, A. O. The global diversity of birds in space and time. Nature 491, 444–448 (2012).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Phillimore, A. B. & Price, T. D. Density-dependent cladogenesis in birds. PLoS Biol. 6, e71 (2008).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Derryberry, E. P. et al. Lineage diversification and morphological evolution in a large-scale continental radiation: the neotropical ovenbirds and woodcreepers (Aves: Furnariidae). Evolution 65, 2973–2986 (2011).

    Article 
    PubMed 

    Google Scholar
     

  • Pennell, M. W., FitzJohn, R. G., Cornwell, W. K. & Harmon, L. J. Model adequacy and the macroevolution of angiosperm functional traits. Am. Nat. 186, E33–E50 (2015).

    Article 
    PubMed 

    Google Scholar
     

  • Rabosky, D. L. et al. Rates of speciation and morphological evolution are correlated across the largest vertebrate radiation. Nat. Commun. 4, 1958 (2013).

    Article 
    PubMed 

    Google Scholar
     

  • Slater, G. J. Phylogenetic evidence for a shift in the mode of mammalian body size evolution at the Cretaceous-Palaeogene boundary. Methods Ecol. Evol. 4, 734–744 (2013).

    Article 

    Google Scholar
     

  • Uyeda, J. C., Caetano, D. S. & Pennell, M. W. Comparative analysis of principal components can be misleading. Syst. Biol. 64, 677–689 (2015).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Westerhold, T. et al. An astronomically dated record of Earth’s climate and its predictability over the last 66 million years. Science 369, 1383–1387 (2020).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Zachos, J., Pagani, M., Sloan, L., Thomas, E. & Billups, K. Trends, rhythms, and aberrations in global climate 65 Ma to present. Science 292, 686–693 (2001).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Prothero, D. R. & Berggren, W. A. in Princeton Series in Geology and Paleontology 582 (Princeton Univ. Press, 1992).

  • Steinthorsdottir, M. et al. The Miocene: the future of the past. Paleoceanogr. Paleoclimatol. 36, e2020PA004037 (2021).

    Article 

    Google Scholar
     

  • Clavel, J. & Morlon, H. Accelerated body size evolution during cold climatic periods in the Cenozoic. Proc. Natl Acad. Sci. USA 114, 4183–4188 (2017).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Selvatti, A. P., Gonzaga, L. P. & Russo, C. A.dM. A Paleogene origin for crown passerines and the diversification of the Oscines in the New World. Mol. Phylogenet. Evol. 88, 1–15 (2015).

    Article 
    PubMed 

    Google Scholar
     

  • Rohde, K. Latitudinal gradients in species diversity: the search for the primary cause. Oikos 65, 514–527 (1992).

    Article 

    Google Scholar
     

  • Quintero, I., Landis, M. J., Jetz, W. & Morlon, H. The build-up of the present-day tropical diversity of tetrapods. Proc. Natl Acad. Sci. USA 120, e2220672120 (2023).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Pulido-Santacruz, P. & Weir, J. T. Extinction as a driver of avian latitudinal diversity gradients. Evolution 70, 860–872 (2016).

    Article 
    PubMed 

    Google Scholar
     

  • Jablonski, D. et al. Out of the tropics, but how? Fossils, bridge species, and thermal ranges in the dynamics of the marine latitudinal diversity gradient. Proc. Natl Acad. Sci. USA 110, 10487–10494 (2013).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Dimitrov, D. et al. Diversification of flowering plants in space and time. Nat. Commun. 14, 7609 (2023).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Rabosky, D. L. et al. An inverse latitudinal gradient in speciation rate for marine fishes. Nature 559, 392–395 (2018).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Harvey, M. G. et al. The evolution of a tropical biodiversity hotspot. Science 370, 1343–1348 (2020).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Upham, N. S., Esselstyn, J. A. & Jetz, W. Ecological causes of uneven mammal diversity. Preprint at bioRxiv https://doi.org/10.1101/504803 (2023).

  • Lawson, A. M. & Weir, J. T. Latitudinal gradients in climatic-niche evolution accelerate trait evolution at high latitudes. Ecol. Lett. 17, 1427–1436 (2014).

    Article 
    PubMed 

    Google Scholar
     

  • Weir, J. T. & Wheatcroft, D. A latitudinal gradient in rates of evolution of avian syllable diversity and song length. Proc. R. Soc. B 278, 1713–1720 (2011).

    Article 
    PubMed 

    Google Scholar
     

  • Burns, M. D. et al. High-latitude ocean habitats are a crucible of fish body shape diversification. Evol. Lett. 8, 669–679 (2024).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Tietje, M. et al. Global variation in diversification rate and species richness are unlinked in plants. Proc. Natl Acad. Sci. USA 119, e2120662119 (2022).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Adams, D. C., Berns, C. M., Kozak, K. H. & Wiens, J. J. Are rates of species diversification correlated with rates of morphological evolution? Proc. R. Soc. B. 276, 2729–2738 (2009).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Cracraft, J. Historical Biogeography and Patterns of Differentiation within the South American Avifauna: Areas of Endemism. Ornithol. Monogr. 36, 49–84 (1985).

    Article 

    Google Scholar
     

  • Knowles, L. L. Did the Pleistocene glaciations promote divergence? Tests of explicit refugial models in montane grasshoppers. Mol. Ecol. 10, 691–701 (2001).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Venditti, C., Meade, A. & Pagel, M. Multiple routes to mammalian diversity. Nature 479, 393–396 (2011).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Hansen, T. F. Is modularity necessary for evolvability?: Remarks on the relationship between pleiotropy and evolvability. Biosystems 69, 83–94 (2003).

    Article 
    PubMed 

    Google Scholar
     

  • Smith, B. T., Gehara, M. & Harvey, M. G. The demography of extinction in eastern North American birds. Proc. R. Soc. B 288, 20201945 (2021).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Santini, L., Tobias, J. A., Callaghan, C., Gallego-Zamorano, J. & Benítez-López, A. Global patterns and predictors of avian population density. Glob. Ecol. Biogeogr. 32, 1189–1204 (2023).

    Article 

    Google Scholar
     

  • Puttick, M. N., Guillerme, T. & Wills, M. A. The complex effects of mass extinctions on morphological disparity. Evolution 74, 2207–2220 (2020).

    Article 
    PubMed 

    Google Scholar
     

  • Hull, P. Life in the aftermath of mass extinctions. Curr. Biol. 25, R941–R952 (2015).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Cole, S. R. & Hopkins, M. J. Selectivity and the effect of mass extinctions on disparity and functional ecology. Sci. Adv. 7, eabf4072 (2021).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Clauset, A., Newman, M. E. & Moore, C. Finding community structure in very large networks. Phys. Rev. E 70, 066111 (2004).

    Article 

    Google Scholar
     

  • Cantwell-Jones, A. et al. Mapping trait versus species turnover reveals spatiotemporal variation in functional redundancy and network robustness in a plant-pollinator community. Funct. Ecol. 37, 748–762 (2023).

    Article 
    CAS 

    Google Scholar
     

  • Kashtan, N., Parter, M., Dekel, E., Mayo, A. E. & Alon, U. Extinctions in heterogeneous environments and the evolution of modularity. Evolution 63, 1964–1975 (2009).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Melo, D. & Marroig, G. Directional selection can drive the evolution of modularity in complex traits. Proc. Natl Acad. Sci. USA 112, 470–475 (2015).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Wagner, G. P., Pavlicev, M. & Cheverud, J. M. The road to modularity. Nat. Rev. Genet. 8, 921–931 (2007).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Parter, M., Kashtan, N. & Alon, U. Environmental variability and modularity of bacterial metabolic networks. BMC Evol. Biol. 7, 169 (2007).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Kashtan, N. & Alon, U. Spontaneous evolution of modularity and network motifs. Proc. Natl Acad. Sci. USA 102, 13773–13778 (2005).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Ronneberger, O., Fischer, P. & Brox, T. U-Net: convolutional networks for biomedical image segmentation. in Medical Image Computing and Computer-Assisted Intervention (eds Navab N., Hornegger J., Wells W. M., & Frangi A. F.). 234–241 (Springer, 2015).

  • He, K., Gkioxari, G., Dollár, P. & Girshick, R. Mask R-CNN. in 2017 IEEE International Conference on Computer Vision (eds.). 2980–2988 (2017).

  • Goolsby, E. W., Bruggeman, J. & Ané, C. Rphylopars: fast multivariate phylogenetic comparative methods for missing data and within-species variation. Methods Ecol. Evol. 8, 22–27 (2017).

    Article 

    Google Scholar
     

  • Marki, P. Z. et al. Supermatrix phylogeny and biogeography of the Australasian Meliphagides radiation (Aves: Passeriformes). Mol. Phylogenet. Evol. 107, 516–529 (2017).

    Article 
    PubMed 

    Google Scholar
     

  • McCullough, J. M. et al. Wallacean and Melanesian Islands promote higher rates of diversification within the global passerine radiation Corvides. Syst. Biol. 71, 1423–1439 (2022).

    Article 
    PubMed 

    Google Scholar
     

  • Cai, T. et al. Near-complete phylogeny and taxonomic revision of the world’s babblers (Aves: Passeriformes). Mol. Phylogenet. Evol. 130, 346–356 (2019).

    Article 
    PubMed 

    Google Scholar
     

  • Alström, P. et al. Systematics of the avian family Alaudidae using multilocus and genomic data. Avian Res. 14, 100095 (2023).

    Article 

    Google Scholar
     

  • Shakya, S. B. & Sheldon, F. H. The phylogeny of the world’s bulbuls (Pycnonotidae) inferred using a supermatrix approach. Ibis 159, 498–509 (2017).

    Article 

    Google Scholar
     

  • De Silva, T. N., Peterson, A. T. & Perktas, U. An extensive molecular phylogeny of weaverbirds (Aves: Ploceidae) unveils broad nonmonophyly of traditional genera and new relationships. Auk 136, ukz041 (2019).

    Article 

    Google Scholar
     

  • Olsson, U. & Alström, P. A comprehensive phylogeny and taxonomic evaluation of the waxbills (Aves: Estrildidae). Mol. Phylogenet. Evol. 146, 106757 (2020).

    Article 
    PubMed 

    Google Scholar
     

  • Barker, F. K., Burns, K. J., Klicka, J., Lanyon, S. M. & Lovette, I. J. New insights into New World biogeography: an integrated view from the phylogeny of blackbirds, cardinals, sparrows, tanagers, warblers, and allies. Auk 132, 333–348 (2015).

    Article 

    Google Scholar
     

  • Baum, B. R. Combining trees as a way of combining data sets for phylogenetic inference, and the desirability of combining gene trees. Taxon 41, 3–10 (1992).

    Article 

    Google Scholar
     

  • Ragan, M. A. Phylogenetic inference based on matrix representation of trees. Mol. Phylogenet. Evol. 1, 53–58 (1992).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Schliep, K. P. phangorn: phylogenetic analysis in R. Bioinformatics 27, 592–593 (2011).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Weeks, B. C. et al. Shared morphological consequences of global warming in North American migratory birds. Ecol. Lett. 23, 316–325 (2020).

    Article 
    PubMed 

    Google Scholar
     

  • Eastman, J. M., Harmon, L. J. & Tank, D. C. Congruification: support for time scaling large phylogenetic trees. Methods Ecol. Evol. 4, 688–691 (2013).

    Article 

    Google Scholar
     

  • Smith, S. A. & O’Meara, B. C. treePL: divergence time estimation using penalized likelihood for large phylogenies. Bioinformatics 28, 2689–2690 (2012).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Claramunt, S. et al. Calibrating the genomic clock of modern birds using fossils. Proc. Natl Acad. Sci. USA 121, e2405887121 (2024).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Claramunt, S. CladeDate: calibration information generator for divergence time estimation. Methods Ecol. Evol. 13, 2331–2338 (2022).

    Article 

    Google Scholar
     

  • Parham, J. F. et al. Best practices for justifying fossil calibrations. Syst. Biol. 61, 346–359 (2012).

    Article 
    PubMed 

    Google Scholar
     

  • Prum, R. O. et al. A comprehensive phylogeny of birds (Aves) using targeted next-generation DNA sequencing. Nature 526, 569–573 (2015).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Jarvis, E. D. et al. Whole-genome analyses resolve early branches in the tree of life of modern birds. Science 346, 1320–1331 (2014).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Kimball, R. T. et al. A phylogenomic supertree of birds. Diversity 11, 109 (2019).

    Article 

    Google Scholar
     

  • Felsenstein, J. Phylogenies and the comparative method. Am. Nat. 125, 1–15 (1985).

    Article 

    Google Scholar
     

  • Hansen, T. F. & Martins, E. P. Translating between microevolutionary process and macroevolutionary patterns: the correlation structure of interspecific data. Evolution 50, 1404–1417 (1996).

    Article 
    PubMed 

    Google Scholar
     

  • Hansen, T. F. Stabilizing selection and the comparative analysis of adaptation. Evolution 51, 1341–1351 (1997).

    Article 
    PubMed 

    Google Scholar
     

  • Blomberg, S. P., Garland, T. Jr. & Ives, A. R. Testing for phylogenetic signal in comparative data: behavioral traits are more labile. Evolution 57, 717–745 (2003).

    Article 
    PubMed 

    Google Scholar
     

  • Jablonski, D. Background and mass extinctions: the alternation of macroevolutionary regimes. Science 231, 129–133 (1986).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • O’Meara, B. C., Ané, C., Sanderson, M. J. & Wainwright, P. C. Testing for different rates of continuous trait evolution using likelihood. Evolution 60, 922–933 (2006).

    Article 
    PubMed 

    Google Scholar
     

  • Rabosky, D. L. Automatic detection of key innovations, rate shifts, and diversity-dependence on phylogenetic trees. PLoS ONE 9, e89543 (2014).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Uyeda, J. C., Zenil-Ferguson, R. & Pennell, M. W. Rethinking phylogenetic comparative methods. Syst. Biol. 67, 1091–1109 (2018).

    Article 
    PubMed 

    Google Scholar
     

  • Eastman, J. M., Alfaro, M. E., Joyce, P., Hipp, A. L. & Harmon, L. J. A novel comparative method for identifying shifts in the rate of character evolution on trees. Evolution 65, 3578–3589 (2011).

    Article 
    PubMed 

    Google Scholar
     

  • Revell, L. J., Mahler, D. L., Peres-Neto, P. R. & Redelings, B. D. A new phylogenetic method for identifying exceptional phenotypic diversification. Evolution 66, 135–146 (2012).

    Article 
    PubMed 

    Google Scholar
     

  • Uyeda, J. C. & Harmon, L. J. A novel Bayesian method for inferring and interpreting the dynamics of adaptive landscapes from phylogenetic comparative data. Syst. Biol. 63, 902–918 (2014).

    Article 
    PubMed 

    Google Scholar
     

  • Beaulieu, J. M., Jhwueng, D.-C., Boettiger, C. & O’Meara, B. C. Modeling stabilizing selection: expanding the Ornstein-Uhlenbeck model of adaptive evolution. Evolution 66, 2369–2383 (2012).

    Article 
    PubMed 

    Google Scholar
     

  • Landis, M. J., Schraiber, J. G. & Liang, M. Phylogenetic analysis using Lévy processes: finding jumps in the evolution of continuous traits. Syst. Biol. 62, 193–204 (2013).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Thomas, G. H. & Freckleton, R. P. MOTMOT: models of trait macroevolution on trees. Methods Ecol. Evol. 3, 145–151 (2012).

    Article 
    CAS 

    Google Scholar
     

  • Khabbazian, M., Kriebel, R., Rohe, K. & Ané, C. Fast and accurate detection of evolutionary shifts in Ornstein–Uhlenbeck models. Methods Ecol. Evol. 7, 811–824 (2016).

    Article 

    Google Scholar
     

  • Ingram, T. & Mahler, D. L. SURFACE: detecting convergent evolution from comparative data by fitting Ornstein-Uhlenbeck models with stepwise Akaike information criterion. Methods Ecol. Evol. 4, 416–425 (2013).

    Article 

    Google Scholar
     

  • Bastide, P., Mariadassou, M. & Robin, S. Detection of adaptive shifts on phylogenies by using shifted stochastic processes on a tree. J. R. Stat. Soc. B 79, 1067–1093 (2017).

    Article 

    Google Scholar
     

  • Revell, L. J. & Harmon, L. J. A discrete character evolution model for phylogenetic comparative biology with Γ-distributed rate heterogeneity among branches of the tree. Preprint at bioRxiv https://doi.org/10.1101/2024.05.25.595896 (2024).

  • Uyeda, J. C., Pennell, M. W., Miller, E. T., Maia, R. & McClain, C. R. The evolution of energetic scaling across the vertebrate tree of life. Am. Nat. 190, 185–199 (2017).

    Article 
    PubMed 

    Google Scholar
     

  • Bastide, P., Ané, C., Robin, S. & Mariadassou, M. Inference of adaptive shifts for multivariate correlated traits. Syst. Biol. 67, 662–680 (2018).

    Article 
    PubMed 

    Google Scholar
     

  • Bartoszek, K. et al. Fast mvSLOUCH: multivariate Ornstein–Uhlenbeck-based models of trait evolution on large phylogenies. Methods Ecol. Evol. 15, 1507–1515 (2024).

    Article 

    Google Scholar
     

  • Gould, S. J., Lewontin, R. C., Maynard Smith, J. & Holliday, R. The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme. Proc. R. Soc. Lond. B 205, 581–598 (1979).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Revell, L. J. Size-correction and principal components for interspecific comparative studies. Evolution 63, 3258–3268 (2009).

    Article 
    PubMed 

    Google Scholar
     

  • Bishop, C. M. Pattern Recognition and Machine Learning (Springer, 2006).

  • Clavel, J. & Morlon, H. Reliable phylogenetic regressions for multivariate comparative data: illustration with the MANOVA and application to the effect of diet on mandible morphology in phyllostomid bats. Syst. Biol. 69, 927–943 (2020).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Alfaro, M. E. et al. Nine exceptional radiations plus high turnover explain species diversity in jawed vertebrates. Proc. Natl Acad. Sci. USA 106, 13410–13414 (2009).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Phillips, P. C. & Arnold, S. J. Hierarchical comparison of genetic variance-covariance matrices. I. Using the Flury hierarchy. Evolution 53, 1506–1515 (1999).

    PubMed 

    Google Scholar
     

  • Stepanova, N., Boyko, J. D., Lin, J., Davis Rabosky, A. R. & Rabosky, D. L. Punctuated versus gradual shifts in the multivariate evolutionary process: a test with paired radiations of scincid lizards. Syst. Biol. 74, 1–17 (2025).

    Article 

    Google Scholar
     

  • Mahler, D. L., Revell, L. J., Glor, R. E. & Losos, J. B. Ecological opportunity and the rate of morphological evolution in the diversification of Greater Antillean anoles. Evolution 64, 2731–2745 (2010).

    Article 
    PubMed 

    Google Scholar
     

  • Konishi, S. & Kitagawa, G. Generalised information criteria in model selection. Biometrika 83, 875–890 (1996).

    Article 

    Google Scholar
     

  • Schwarz, G. Estimating the dimension of a model. Ann. Stat. 6, 461–464 (1978).

    Article 

    Google Scholar
     

  • Revell, L. J. phytools: an R package for phylogenetic comparative biology (and other things). Methods Ecol. Evol. 3, 217–223 (2012).

    Article 

    Google Scholar
     

  • Burnham, K. P. & Anderson, D. R. Model Selection and Multimodel Inference: A Practical Information-Theoretic Approach 2nd edn, Vol. 2 (Springer-Verlag, 2002).

  • Burnham, K. P. & Anderson, D. R. Multimodel inference: understanding AIC and BIC in model selection. Sociol. Methods Res. 33, 261–304 (2004).

    Article 

    Google Scholar
     

  • Berv, J. S. et al. Supplementary code for: Rates of passerine body plan evolution in time and space. v1.0.0. Zenodo https://doi.org/10.5281/zenodo.19211076 (2026).

  • Spence, A. J. Scaling in biology. Curr. Biol. 19, R57–R61 (2009).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Wilcoxon, F. Individual comparisons by ranking methods. Biom. Bull. 1, 80–83 (1945).

    Article 

    Google Scholar
     

  • Kolmogorov, A. N. Sulla determinazione empírica di uma legge di distribuzione. G. Ist. Ital. Attuari 4, 83–91 (1933).


    Google Scholar
     

  • Smirnov, N. Table for estimating the goodness of fit of empirical distributions. Ann. Math. Stat. 19, 279–281, 273 (1948).

    Article 

    Google Scholar
     

  • Marsaglia, G., Tsang, W. W. & Wang, J. Evaluating Kolmogorov’s distribution. J. Stat. Softw. 8, 1–4 (2003).

    Article 

    Google Scholar
     

  • Moss, J. univariateML: An R package for maximum likelihood estimation of univariate densities. J. Open Source Softw. 4, 1863 (2019).

    Article 

    Google Scholar
     

  • Title, P. O. et al. The macroevolutionary singularity of snakes. Science 383, 918–923 (2024).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • BirdLife International & Handbook of the Birds of the World. Bird Species Distribution Maps of the World. Version 2021.1 edn (BirdLife International, 2021).

  • Pebesma, E. Simple features for R: standardized support for spatial vector data. R J. 10, 439–446 (2018).

    Article 

    Google Scholar
     

  • Pebesma, E. & Bivand, R. Spatial Data Science: With Applications in R. (Chapman and Hall/CRC, 2023).

  • H3: A Hexagonal Hierarchical Geospatial Indexing System (Uber Technologies, 2018).

  • O’Brien, L. h3jsr: Access Uber’s H3 Library (CRAN, 2024).

  • GBIF Secretariat. GBIF Backbone Taxonomy. Checklist dataset edn (GBIF Secretariat, 2023).

  • Redding, D. W. & Mooers, A. O. Incorporating evolutionary measures into conservation prioritization. Conserv. Biol. 20, 1670–1678 (2006).

    Article 
    PubMed 

    Google Scholar
     

  • Ord, K. Estimation methods for models of spatial interaction. J. Am. Stat. Assoc. 70, 120–126 (1975).

    Article 

    Google Scholar
     

  • Bivand, R. & Piras, G. Comparing implementations of estimation methods for spatial econometrics. J. Stat. Softw. 63, 1–36 (2015).

    Article 

    Google Scholar
     

  • Bivand, R., Hauke, J. & Kossowski, T. Computing the Jacobian in Gaussian spatial autoregressive models: an illustrated comparison of available methods. Geogr. Anal. 45, 150–179 (2013).

    Article 

    Google Scholar
     

  • Fick, S. E. & Hijmans, R. J. WorldClim 2: new 1-km spatial resolution climate surfaces for global land areas. Int. J. Climatol. 37, 4302–4315 (2017).

    Article 

    Google Scholar
     

  • Shepard, D. A two-dimensional interpolation function for irregularly-spaced data. In Proc. 1968 23rd ACM National Conference, 517–524 (Association for Computing Machinery, 1968).

  • Moran, P. A. P. Notes on continuous stochastic phenomena. Biometrika 37, 17–23 (1950).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Willis, K. J. et al. The tempo of avian diversification during the Quaternary. Phil. Trans. R. Soc. Lond. B 359, 215–220 (2004).

    Article 

    Google Scholar
     

  • Lovette, I. J. Glacial cycles and the tempo of avian speciation. Trends Ecol. Evol. 20, 57–59 (2005).

    Article 
    PubMed 

    Google Scholar
     

  • Nadachowska-Brzyska, K., Li, C., Smeds, L., Zhang, G. & Ellegren, H. Temporal dynamics of avian populations during Pleistocene revealed by whole-genome sequences. Curr. Biol. 25, 1375–1380 (2015).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Weir, J. T. & Schluter, D. Ice sheets promote speciation in boreal birds. Proc. R. Soc. Lond. B 271, 1881–1887 (2004).

    Article 

    Google Scholar
     

  • Jablonski, D. Lessons from the past: evolutionary impacts of mass extinctions. Proc. Natl Acad. Sci. USA 98, 5393–5398 (2001).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Atchley, W. R. M-Statistics and morphometric divergence. Science 208, 1059–1060 (1980).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Huttegger, S. M. & Mitteroecker, P. Invariance and meaningfulness in phenotype spaces. Evol. Biol. 38, 335–351 (2011).

    Article 

    Google Scholar
     

  • Csardi, G. & Nepusz, T. The igraph software package for complex network research. InterJ. Complex Syst. 1695, 1–9 (2006).


    Google Scholar
     

  • Schuurman, T. & Bruner, E. Modularity and community detection in human brain morphology. Anat. Rec. 307, 345–355 (2024).

    Article 

    Google Scholar
     

  • Padi, M. & Quackenbush, J. Detecting phenotype-driven transitions in regulatory network structure. npj Syst. Biol. Appl. 4, 16 (2018).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Adams, D. C. A generalized K statistic for estimating phylogenetic signal from shape and other high-dimensional multivariate data. Syst. Biol. 63, 685–697 (2014).

    Article 
    PubMed 

    Google Scholar
     

  • Collyer, M. L., Baken, E. K. & Adams, D. C. A standardized effect size for evaluating and comparing the strength of phylogenetic signal. Methods Ecol. Evol. 13, 367–382 (2022).

    Article 

    Google Scholar
     

  • Baken, E. K., Collyer, M. L., Kaliontzopoulou, A. & Adams, D. C. geomorph v4.0 and gmShiny: enhanced analytics and a new graphical interface for a comprehensive morphometric experience. Methods Ecol. Evol. 12, 2355–2363 (2021).

    Article 

    Google Scholar
     

  • Benjamini, Y. & Yekutieli, D. The control of the false discovery rate in multiple testing under dependency. Ann. Stat. 29, 1165–1188 (2001).

    Article 

    Google Scholar
     

  • Berv, J. et al. Supplementary data archive for Rates of passerine body plan evolution in time and space. v1.0.0. Zenodo https://doi.org/10.5281/zenodo.19198393 (2026).

  • Berv, J. S. et al. bifrost: an R package for scalable inference of phylogenetic shifts in multivariate evolutionary dynamics. Preprint at bioRxiv https://doi.org/10.64898/2026.04.12.718036 (2026).