Trilobite Eyes

NOTE: For many of these references, I have pdfs. This list includes all known papers on the structure of fossil trilobite eyes. If you think I am missing something, please let me know!

Bennett, C. E., M. Williams, M. J. Leng, M. R. Lee, M. Bonifacie, D. Calmels, R. A. Fortey, J. R. Laurie, A. W. Owen, A. A. Page, A. Munnecke, and T. R. A. Vandenbroucke. 2018. Oxygen isotope analysis of the eyes of pelagic trilobites: Testing the application of sea temperature proxies for the Ordovician. Gondwana Research 57: 157-169.

Bruton, D. L., and W. Haas. 2003. The puzzling eye of Phacops. Special Papers in Palaeontology 70: 349-361.

Bruton, D. L. 2006. A reconstruction of Telephina bicuspis, a pelagic trilobite from the Middle Ordovician of the Oslo Region, Norway. Lethaia 39: 359-364.

Budil, P., and F. Hörbinger. 2007. Exoskeletal structures and ultrastructures in Lower Devonian dalmanitid trilobites of the Prague Basin (Czech Republic). Bulletin of Geosciences 82: 27-36.

Campbell, K. S. W. 1975. The functional anatomy of phacopid trilobites: musculature and eyes. Journal and Proceedings, Royal Society of New South Wales 108: 168-188.

Clarkson, E. N. K. 1966. Schizochroal eyes and vision of some Silurian acastid trilobites. Palaeontology 9: 1-29.

Clarkson, E. N. K. 1966. Schizocroal eyes and vision in some phacopid trilobites. Palaeontology 9: 464-487.

Clarkson, E. N. K. 1967. Environmental significance of eye-reduction in trilobites and recent arthropods. Marine Geology 5: 367-375.

Clarkson, E. N. K. 1967. Fine structure of the eye in two species of Phacops (Trilobita). Palaeontology 10: 603-616.

Clarkson, E. N. K. 1968. Structure of the eye of Crozonaspis struvei (Trilobita, Dalmanitidae, Zeliszkellinae). Senckenbergiana lethaea 49: 383-393.

Clarkson, E. N. K. 1969. Dimorphism of the eye in Weberides shunnerensis (King) [Trilobita]. In Sexual dimorphism in fossil Metazoa and taxonomic implications, G. E. G. Westermann, ed. E. Schweizerbart’sche Verlagsbuchhandlung (Nägele u. Obermiller), Stuttgart, p. 185-195.

Clarkson, E. N. K. 1969. On the schizocroal eyes of three species of Reedops (Trilobita, Phacopidae) from the Lower Devonian of Bohemia. Transactions of the Royal Society of Edinburgh 68: 183-205.

Clarkson, E. N. K. 1971. On the early schizocroal eyes of Ormathops (Trilobita, Zeliskellinae). Mémoires du Bureau de Recherches Géologiques et Minières 73: 51-63.

Clarkson, E. N. K. 1973. Morphology and evolution of the eye in Upper Cambrian Olenidae (Trilobita). Palaeontology 16: 735-763.

Clarkson, E. N. K. 1973. The eyes of Asaphus raniceps Dalman (Trilobita). Palaeontology 16: 425-444.

Clarkson, E. N. K. 1975. The evolution of the eye in trilobites. Fossils and Strata 4: 7-31.

Clarkson, E. N. K., and R. Levi-Setti. 1975. Trilobite eyes and the optics of Des Cartes and Huygens. Nature 254: 663-667.

Clarkson, E. N. K. 1979. The visual system of trilobites. Palaeontology 22: 1-22.

Clarkson, E. N. K., R. Levi-Setti, and G. Horváth. 2006. The eyes of trilobites; the oldest preserved visual system. Arthropod Structure & Development 35: 247-259.

Cowen, R., and J. S. Kelley. 1976. Stereoscopic vision within the schizochroal eye of trilobites. Nature 261: 130-131.

Crônier, C., and E. N. K. Clarkson. 2001. Variation of eye-lens distribution in a new late Devonian phacopid trilobite. Transactions of the Royal Society of Edinburgh: Earth Sciences 92: 103-113.

Crônier, C., R. Feist, and J.-C. Auffray. 2004. Variation in the eye of Acuticryphops (Phacopina, Trilobita) and its evolutionary significance: a biometric and morphometric approach. Paleobiology 30: 471-481.

Crônier, C., P. Budil, O. Fatka, and L. Laibl. 2015. Intraspecific bimodal variability in eye lenses of two Devonian trilobites. Paleobiology 41: 554-569.

Egri, Á., and G. Horváth. 2012. Possible optical functions of the central core in lenses of trilobite eyes: spherically corrected monofocality or bifocality. Journal of the Optical Society of America A 29: 1965-1976.

Fan, Q., W. Xu, X. Hu, W. Zhu, T. Yue, C. Zhang, F. Yan, L. Chen, H. J. Lezec, Y. Lu, A. Agrawal, and T. Xu. 2022. Trilobite-inspired neural nanophotonic light-field camera with extreme depth-of-field. Nature Communications 13: 2130.

Fatka, O., P. Budil, and O. Zicha. 2021. Exoskeletal and eye repair in Dalmanitina socialis (Trilobita): An example of blastemal regeneration in the Ordovician. International Journal of Paleopathology 34: 113-121.

Fordyce, D., and T. W. Cronin. 1989. Comparison of fossilized schizochroal compound eyes of phacopid trilobites with eyes of modern marine crustaceans and other arthropods. Journal of Crustacean Biology 9: 554-569.

Fordyce, D., and T. W. Cronin. 1993. Trilobite vision: a comparison of schizochroal and holochroal eyes with the compound eyes of modern arthropods. Paleobiology 19: 288-303.

Greenberger, R. E. 2020. Investigating rare biomineralization structures in trilobites. Department of Geological Sciences M.S.: 1-58.

Gál, J., G. Horváth, E. N. K. Clarkson, and O. Haiman. 2000. Image formation by bifocal lenses in a trilobite eye? Vision Research 40: 843-853.

Gál, J., G. Horváth, and E. N. K. Clarkson. 2000. Reconstruction of the shape and optics of the lenses in the abathochroal-eyed trilobite Neocobboldia chinlinica. Historical Biology 14: 193-204.

Haack, S. C. 1987. The evolution and acuity of the schizochroal eye in trilobites. Evolutionary Theory 8: 69-72.

Han, N.-R., and Y. Zhang. 1985. Holochroal eyes of Cyclopyge. Geological Review 31: 390-395.

Han, N.-R. 2001. The eyes of Ordovician trilobite Telephina convexa Lu. Acta Palaeontologica Sinica 40: 399-408.

Harzsch, S., and G. Hafner. 2006. Evolution of eye development in arthropods: Phylogenetic aspects. Arthropod Structure & Development 35: 319-340.

Hessler, R. R. 1976. An eye for an eye. Paleobiology 2: 94-97.

Horváth, G. 1989. Geometric optics of trilobite eyes: A theoretical study of the shape of the aspherical interface in the cornea of schizochroal eyes of phacopid trilobites. Mathematical Biosciences 96: 79-94.

Horváth, G. 1996. The lower lens unit in schizochroal trilobite eyes reduces reflectivity; On the possible optical function of the intralensar bowl. Historical Biology 12: 83-92.

Horváth, G., E. N. K. Clarkson, and W. Pix. 1997. Survey of modern counterparts of schizochroal trilobite eyes: Structural and functional similarities and differences. Historical Biology 12: 229-263.

Isberg, O. 1917. Ein regeneriertes trilobitenauge. Geologiska Föreningens i Stockholm Förhandlingar 39: 593-596.

Jell, P. A. 1975. The abathochroal eye of Pagetia, a new type of trilobite eye. Fossils and Strata 4: 33-43.

Klug, C., H. Schulz, and K. deBaets. 2009. Red Devonian trilobites with green eyes from Morocco and the silicification of the trilobite exoskeleton. Acta Palaeontologica Polonica 54: 117-123.

Krueger, H.-H. 1974. Die Porenöffnungen an den Augenstielen von Asaphus (Neoasaphus) cornutus und Asaphus (Neoasaphus) kowalewskii [The pore openings on the eyestalks of Asaphus (Neoasaphus) cornutus and Asaphus (Neoasaphus) kowalewskii]. Zeitschrift für Geologische Wissenschaften 2: 1085-1091.

Lee, M. R., C. Torney, and A. W. Owen. 2007. Magnesium-rich intralensar structures in schizochroal trilobite eyes. Palaeontology 50: 1031-1037.

Lindgren, J., D. E. Nilsson, P. Sjövall, M. Jarenmark, S. Ito, K. Wakamatsu, B. P. Kear, B. P. Schultz, R. L. Sylvestersen, H. Madsen, J. R. LaFountain, C. Alwmark, M. E. Eriksson, S. A. Hall, P. Lindgren, I. Rodríguez-Meizoso, and P. Ahlberg. 2019. Fossil insect eyes shed light on trilobite optics and the arthropod pigment screen. Nature 573: 122-125.

Lindström, G. 1901. Researches on the visual organs of the trilobites. Kongliga Svenska Vetenskapsakademiens Handlingar 34: 1-87.

Lorenz, P. 1991. Die Variabilität und Ontogenie des Komplexauges von Phacops granulatus (Münster 1840) (Trilobita; Ober-Devon) [Variability and ontogeny of Phacops granulatus (Münster 1840) complexes (Trilobita, Upper Devonian)]. Geologica et Palaeontologica 25: 47-55.

Miller, J., and E. N. K. Clarkson. 1980. The post-ecdysial development of the cuticle and the eye of the Devonian trilobite Phacops rana milleri Stewart 1927. Philosophical Transactions of the Royal Society of London B. Biological Sciences 288: 461-480.

Packard, A. S., Jr. 1880. The structure of the eye of trilobites. American Naturalist 14: 503-508.

Richter, R. 1922. Über einen fall äußerster rückbildung des schizochroalen trilobiten-auges. Centralblatt für Mineralogie und P 11: 344-352.

Rose, J. N. 1968. The eyes of Isotelus and Nileus. The Proceedings of the Iowa Academy of Science 74: 178-185.

Ruedemann, R. 1916. The presence of a median eye in trilobites. New York State Museum Bulletin 189: 127-143.

Schoenemann, B. 2007. Fossile augensysteme: bericht über zur zeit durchgeführte analysen fossiler augensysteme. Freiberger Forschungshefte C 524: 85-96.

Schoenemann, B. 2007. Trilobite eyes and a new type of neural superposition eye in an ancient system. Palaeontographica Abteilung A 281: 63-91.

Schoenemann, B., and E. N. K. Clarkson. 2008. Did the trabecula in phacopid lenses act as light-guides? Cuadernos del Museo Geominero, n°9 Advances in Trilobite Research: 351-354.

Schoenemann, B., E. N. K. Clarkson, and A. Franz. 2008. Sublensar capsules in phacopid eyes. Cuadernos del Museo Geominero, n°9 Advances in Trilobite Research: 355-359.

Schoenemann, B., E. N. K. Clarkson, P. Ahlberg, and M. E. D. Álvarez. 2010. A tiny eye indicating a planktonic trilobite. Palaeontology 53: 695-701.

Schoenemann, B., and E. N. K. Clarkson. 2011. Light guide lenses in trilobites? Earth and Environmental Science Transactions of the Royal Society of Edinburgh 102: 17-23.

Schoenemann, B., and E. N. K. Clarkson. 2011. The eyes of Bohemian trilobites. Geologické výzkumy na Moravĕ a ve Slezsku 18: 45-50.

Schoenemann, B., and E. N. K. Clarkson. 2012. Compound eyes in the Chengjiang Biota. Acta Universitatis Latviensis, Earth and Environmental Sciences 783: 57-71.

Schoenemann, B., and E. N. K. Clarkson. 2012. Insights to eyes of phacopid trilobites. Scientific Papers. University of Latvia. Earth and Environmental Sciences 783: 72-75.

Schoenemann, B., and E. N. K. Clarkson. 2013. Discovery of some 400 million year-old sensory structures in the compound eyes of trilobites. Science Reports 3: 1429.

Schoenemann, B., and E. N. K. Clarkson. 2015. Eyes and vision in the coeval Furongian trilobites Sphaerophthalmus alatus (Boeck, 1938) and Ctenopyge (Mesoctenopyge) tumida Westergård, 1922, from Bornholm, Denmark. Palaeontology 58: 133-140.

Schoenemann, B., E. N. Clarkson, and G. Horváth. 2015. Why did the UV-A-induced photoluminescent blue-green glow in trilobite eyes and exoskeletons not cause problems for trilobites? PeerJ 3: e1492.

Schoenemann, B., H. Pärnaste, and E. N. K. Clarkson. 2017. Structure and function of a compound eye, more than half a billion years old. Proceedings of the National Academy of Science of the United States of America 114: 13489-13494.

Schoenemann, B., E. N. Clarkson, and M. Høyberget. 2017. Traces of an ancient immune system - how an injured arthropod survived 465 million years ago. Science Reports 7: 40330.

Schoenemann, B., and E. N. K. Clarkson. 2017. Vision in fossilised eyes. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 106: 209-220.

Schoenemann, B. 2018. Evolution of eye reduction and loss in trilobites and some related fossil arthropods. Emerging Science Journal 2: 272-286.

Schoenemann, B., and E. N. K. Clarkson. 2020. Insights into a 429-million-year-old compound eye. Science Reports 10: 12029.

Schoenemann, B., E. N. K. Clarkson, C. Bartels, W. Südkamp, G. E. Rössner, and U. Ryck. 2021. A 390 million-year-old hyper-compound eye in Devonian phacopid trilobites. Scientific Reports 11: 19505.

Schoenemann, B. 2021. An overview on trilobite eyes and their functioning. Arthropod Structure & Development 61: 101032.

Schoenemann, B., and E. N. K. Clarkson. 2021. Points of view in understanding trilobite eyes. Nature Communications 12: 2081.

Scholtz, G., A. Staude, and J. A. Dunlop. 2019. Trilobite compound eyes with crystalline cones and rhabdoms show mandibulate affinities. Nature Communications 10: 2503.

Scholtz, G., A. Staude, and J. A. Dunlop. 2021. Reply to “Points of view in understanding trilobite eyes”. Nature Communications 12: 2084.

Shaw, F. C., and A. R. Ormiston. 1964. The eye socle of the trilobites. Journal of Paleontology 38: 1001-1002.

Spencer, W. K. 1903. The hypostomic eyes of trilobites. Geological Magazine 10: 489-492.

Stockton, W. L., and R. Cowen. 1976. Stereoscopic vision in one eye: Paleophpysiology of the schizocroal eye of trilobites. Paleobiology 2: 304-315.

Tanaka, G., B. Schoenemann, K. El Hariri, T. Ono, E. Clarkson, and H. Maeda. 2015. Vision in a Middle Ordovician trilobite eye. Palaeogeography, Palaeoclimatology, Palaeoecology 433: 129-139.

Tasch, P. 1952. Adaptive trend in eyeline development in the Olenellidae. Journal of Paleontology 26: 484-488.

Thomas, A. T. 1998. Variation in the eyes of the Silurian trilobites Eophacops and Acaste and its significance. Palaeontology 41: 897-911.

Thomas, A. T. 2005. Developmental palaeobiology of trilobite eyes and its evolutionary significance. Earth-Science Reviews 71: 77-93.

Torney, C., M. R. Lee, and A. W. Owen. 2008. An electron backscatter diffraction study of Geesops: a broader view of trilobite vision? Cuadernos del Museo Geominero, n°9 Advances in Trilobite Research: 389-394.

Torney, C. 2011. Mineral eyes: lessons from the natural world. School of Geographical and Earth Sciences Ph.D: 1-414.

Torney, C., M. R. Lee, and A. W. Owen. 2014. Microstructure and growth of the lenses of schizochroal trilobite eyes. Palaeontology 57: 783-799.

Towe, K. 1973. Trilobite eyes: Calcified lenses in vivo. Science 179: 1007-1009.

van der Straaten, R. 2006. Trilobieten hebben ogen van kristal [Trilobites have eyes of crystal]. Grondboor en Hamer 60: 121-122.

Walcott, C. D. 1883. Injury sustained by the eye of a trilobite at the time of the moulting of the shell. American Journal of Science 26: 302.

Zhang, X.-G., and E. N. K. Clarkson. 1990. The eyes of Lower Cambrian eodiscid trilobites. Palaeontology 33: 911-932.

Zydorowicz, T. 1993. Geometria holochroiciznego oka trylobita a problemy uniformitaryzmu w geologii. Geometry of the trilobite’s holochroic eye and the issue of uniformitarism in geology. Przeglad Geologiczny 41: 432-434.