Eakin, R. M. The Third Eye (Univ. of California Press,1973).
Lamb, T. D. Evolution of phototransduction, vertebrate photoreceptors and retina. Prog. Retinal Eye Res. 36, 52–119 (2013).
Janvier, P. Early Vertebrates (Clarendon,1996).
Darwin, C. On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life 1st edn (Murray, 1859).
Benoit, J., Abdala, F., Manger, P. R. & Rubidge, B. S. The sixth sense in mammalian forerunners: variability of the parietal foramen and the evolution of the pineal eye in South African Permo-Triassic Eutheriodont Therapsids. Acta Palaeontol. Pol. 61, 777–789 (2016).
Lamb, T. D., Collin, S. P. & Pugh, E. N. Evolution of the vertebrate eye: opsins, photoreceptors, retina and eye cup. Nat. Rev. Neurosci. 8, 960–976 (2007).
Clements, T. et al. The eyes of Tullimonstrum reveal a vertebrate affinity. Nature 532, 500–503 (2016).
Gabbott, S. E. et al. Pigmented anatomy in Carboniferous cyclostomes and the evolution of the vertebrate eye. Proc. Biol. Sci. 283, 20161151 (2016).
Conway Morris, S. & Caron, J.-B. A primitive fish from the Cambrian of North America. Nature 512, 419–422 (2014).
Gabbott, S. E., Sansom, R. S. & Purnell, M. A. Systematic analysis of exceptionally preserved fossils: correlated patterns of decay and preservation. Palaeontology 64, 789–803 (2021).
Braun, K. & Stach, T. Structure and ultrastructure of eyes and brains of Thalia democratica (Thaliacea, Tunicata, Chordata). J. Morphol. 278, 1421–1437 (2017).
Shu, D.-G. et al. Lower Cambrian vertebrates from south China. Nature 402, 42–46 (1999).
Shu, D.-G. et al. Head and backbone of the Early Cambrian vertebrate Haikouichthys. Nature 421, 526–529 (2003).
Hou, X.-G., Aldridge, R. J., Siveter, D. J., Siveter, D. J. & Feng, X.-H. New evidence on the anatomy and phylogeny of the earliest vertebrates. Proc. Biol. Sci. 269, 1865–1869 (2002).
Zhao, J., Li, G.-B. & Selden, P. A. A poorly preserved fish-like animal from the Chengjiang Lagerstätte (Cambrian series 2, stage 3). Palaeogeogr. Palaeoclimatol. Palaeoecol. 520, 163–172 (2019).
Zhang, X.-G. & Hou, X.-G. Evidence for a single median fin-fold and tail in the Lower Cambrian vertebrate Haikouichthys ercaicunensis. J. Evol. Biol. 17, 1162–1166 (2004).
Liu, Y. et al. Comparisons of the structural and chemical properties of melanosomes isolated from retinal pigment epithelium, iris and choroid of newborn and mature bovine eyes. Photochem. Photobiol. 81, 510–516 (2005).
Shawkey, M. D., D’Alba, L., Xiao, M., Schutte, M. & Buchholz, R. Ontogeny of an iridescent nanostructure composed of hollow melanosomes. J. Morphol. 276, 378–384 (2015).
D’Alba, L. & Shawkey, M. D. Melanosomes: biogenesis, properties, and evolution of an ancient organelle. Physiol. Rev. 99, 1–19 (2019).
Roy, A., Pittman, M., Kaye, T. G. & Saitta, E. T. Sediment-encased pressure–temperature maturation experiments elucidate the impact of diagenesis on melanin-based fossil color and its paleobiological implications. Paleobiology 49, 712–732 (2023).
Agrup, G., Hansson, C., Rorsman, H. & Rosengren, E. The effect of cysteine on oxidation of tyrosine, dopa, and cysteinyldopas. Arch. Dermatol. Res. 272, 103–115 (1982).
Bush, W. D. et al. The surface oxidation potential of human neuromelanin reveals a spherical architecture with a pheomelanin core and a eumelanin surface. Proc. Natl Acad. Sci. USA 103, 14785–14789 (2006).
Simon, J. D. & Peles, D. N. The red and the black. Acc. Chem. Res. 43, 1452–1460 (2010).
Colleary, C. et al. Chemical, experimental, and morphological evidence for diagenetically altered melanin in exceptionally preserved fossils. Proc. Natl Acad. Sci. USA 112, 12592–12597 (2015).
Briggs, D. E. G. & Williams, S. H. The restoration of flattened fossils. Lethaia 14, 157–164 (1981).
Dearden, R. P. et al. The three-dimensionally articulated oral apparatus of a Devonian heterostracan sheds light on feeding in Palaeozoic jawless fishes. Proc. Biol. Sci. 291, 20232258 (2024).
Gagnier, P. Sacabambaspis janvieri, vertebre Ordovicien de Bolivie. I. Analyse morphologique. Ann. Paléontol. 79, 19–69 (1993).
Fritzsch, B. & Martin, P. R. Vision and retina evolution: how to develop a retina. IBRO Neurosci. Rep. 12, 240–248 (2022).
Ekström, P. & Meissl, H. Evolution of photosensory pineal organs in new light: the fate of neuroendocrine photoreceptors. Philos. Trans. R. Soc. 358, 1679–1700 (2003).
Janvier, P. & Arsenault, M. The anatomy of Euphanerops longaevus Woodward, 1900, an anaspid-like jawless vertebrate from the Upper Devonian of Miguasha, Quebec, Canada. Geodiversitas 29, 143–216 (2007).
Gai, Z.-K., Donoghue, P. C. J., Zhu, M., Janvier, P. & Stampanoni, M. Fossil jawless fish from China foreshadows early jawed vertebrate anatomy. Nature 476, 324–327 (2011).
Erwin, D. H. et al. The Cambrian conundrum: early divergence and later ecological success in the early history of animals. Science 334, 1091–1097 (2011).
Meert, J. G., Levashova, N. M., Bazhenov, M. L. & Landing, E. Rapid changes of magnetic field polarity in the late Ediacaran: linking the Cambrian evolutionary radiation and increased UV-B radiation. Gondwana Res. 34, 149–157 (2016).
Nilsson, D.-E. Eye evolution and its functional basis. Vis. Neurosci. 30, 5–20 (2013).
Vinther, J., Porras, L., Young, F. J., Budd, G. E. & Edgecombe, G. D. The mouth apparatus of the Cambrian gilled lobopodian Pambdelurion whittingtoni. Palaeontology 59, 841–849 (2016).
Park, T.-Y. S. et al. A giant stem-group chaetognath. Sci. Adv. 10, eadi6678 (2024).
Vinther, J. et al. A fossilised ventral ganglion reveals a chaetognath affinity for Cambrian nectocaridids. Sci. Adv. 11, eadu6990 (2025).
Paterson, J. R. et al. Acute vision in the giant Cambrian predator Anomalocaris and the origin of compound eyes. Nature 480, 237–240 (2011).
Kasumyan, A. O. & Pavlov, D. S. Evolution of schooling behavior in fish. J. Ichthyol. 58, 670–678 (2018).
Bok, M. J. & Buschbeck, E. K. in Distributed Vision: From Simple Sensors to Sophisticated Combination Eyes (eds Buschbeck, E. K. & Bok, M. J.) 1–19 (Springer, 2023).
Liu, C.-X. The Dark Forest (Chongqing Press, 2008).
Rincón Camacho, L. et al. The pineal complex: a morphological and immunohistochemical comparison between a tropical (Paracheirodon axelrodi) and a subtropical (Aphyocharax anisitsi) characid species. J. Morphol. 277, 1355–1367 (2016).
Dearden, R. P. et al. The oldest three-dimensionally preserved vertebrate neurocranium. Nature 621, 782–787 (2023).
Lindgren, J. et al. Skin pigmentation provides evidence of convergent melanism in extinct marine reptiles. Nature 506, 484–488 (2014).
Lindgren, J. et al. Molecular preservation of the pigment melanin in fossil melanosomes. Nat. Commun. 3, 824 (2012).