2026
Qing-hao GUO, Jie YANG, Brigitte SCHOENEMANN, Xiguang ZHANG
An early Cambrian bivalved euarthropod with specialised limbs and highly sclerotised abdomen
Earth and Environmental Science Transactions of the Royal Society of Edinburgh 2025;116(3-4):102-114
Earth and Environmental Science Transactions of The Royal Society of Edinburgh , Volume 116 - Special Issue 3-4 - October 2025:
A Palaeontological Odyssey: A tribute to Euan Clarkson DSc, FRSE (1937–2024)
DOI: 10.1017/S1755691025101011
https://www.cambridge.org/core/journals/earth-and-environmental-science-transactions-of-royal-society-of-edinburgh/article/an-early-cambrian-bivalved-euarthropod-with-specialised-limbs-and-highly-sclerotised-abdomen/3B1744E2E87C3C85A15510E3AA42BE66
Brigitte SCHOENEMANN, Jaap EYZENGA
Insights into the genetic program of compound-eye formation by an injured phacopine trilobite
Earth and Environmental Science Transactions of the Royal Society of Edinburgh 2025;116(3-4):170-176
Earth and Environmental Science Transactions of The Royal Society of Edinburgh , Volume 116 - Special Issue 3-4 - October 2025:
A Palaeontological Odyssey: A tribute to Euan Clarkson DSc, FRSE (1937–2024)
DOI: 10.1017/S1755691025100984
https://www.cambridge.org/core/journals/earth-and-environmental-science-transactions-of-royal-society-of-edinburgh/article/insights-into-the-genetic-program-of-compoundeye-formation-by-an-injured-phacopine-trilobite/8A68BDFE5DF511A9024379B3DEF0EA6F
Mehdi Ghaedi, Brigitte Schoenemann, Ali Bahrami, Hemen Moradi-Salimi & Wenkun Qie
Beyond the obvious fossil record: microscopic remains of the Late Devonian to earliest Carboniferous trilobite elements in Alborz Basin, Iran.
Carbonates Evaporites 41, 56 (2026).
DOI: 10.1007/s13146-026-01259-5
https://link.springer.com/article/10.1007/s13146-026-01259-5
2025
Michel Schmidt*, Brigitte Schoenemann*, Xianguang Hou, Roland R. Melzer, Yu Liu
* contributed equally
Pygmaclypeatus daziensis, a unique lower Cambrian arthropod with two different compound eye systems
Communications Biology 8, 317 (2025)
DOI: 10.1038/s42003-025-07664-1
https://www.nature.com/articles/s42003-025-07664-1
Brigitte Schoenemann
Trilobite Eyes and Their Evolution
Arthropoda 2025, 3(1), 3
DOI: 10.3390/arthropoda3010003
https://doi.org/10.3390/arthropoda3010003
2024
Schoenemann, B., Hoekstra, H.J.W.M., Horváth, G., Clarkson, E.N.K.
Vision of Trilobites and Polarized Light
In: Horváth, G. (eds) Polarization Vision and Environmental Polarized Light
Springer Series in Vision Research. Springer, Cham (2024)
DOI: 10.1007/978-3-031-62863-4_22
https://doi.org/10.1007/978-3-031-62863-4_22
Schoenemann, B., Hughes, N.C., Myrow, P.M.
Eye structure and function in Ameura (Trilobita) from the Upper Carboniferous Minturn Formation at Bond, Colorado, USA
Irish Journal of Earth Sciences 42(1), 299-307 (2024)
DOI: 10.1353/ijes.2024.a935035
https://dx.doi.org/10.1353/ijes.2024.a935035
2023
Brigitte Schoenemann & Euan N.K. Clarkson
The median eyes of trilobites
Scientific Reports 13, 3917 (2023)
DOI: 10.1038/s41598-023-31089-7
https://www.nature.com/articles/s41598-023-31089-7
2021
B. Schoenemann, E.N.K. Clarkson, C. Bartels, W. Südkamp, U. Ryck
A 390 million-year-old hyper-compound eye in Devonian phacopid trilobites
Scientific Reports 11, 19505 (2021)
DOI: 10.1038/s41598-021-98740-z
https://www.nature.com/articles/s41598-021-98740-z
Brigitte Schoenemann & Euan N.K. Clarkson
Points of view in understanding trilobite eyes
Nature Communications 12, 2081 (2021)
DOI: 10.1038/s41467-021-22227-8
https://www.nature.com/articles/s41467-021-22227-8
Brigitte Schoenemann
An overview on trilobite eyes and their functioning
Arthropod Structure & Development 61, 2021, 101032
DOI: 10.1016/j.asd.2021.101032
https://www.sciencedirect.com/science/article/pii/S1467803921000062
2020
Brigitte Schoenemann & Euan N.K. Clarkson
Insights into a 429-million-year-old compound eye
Scientific Reports 10, 12029 (2020)
DOI: 10.1038/s41598-020-69219-0
https://www.nature.com/articles/s41598-020-69219-0
2019
Brigitte Schoenemann, Markus Poschmann & Euan N.K. Clarkson
Insights into the 400 million-year-old eyes of giant sea scorpions (Eurypterida) suggest the structure of Palaeozoic compound eyes.
Scientific Reports 9, 17797 (2019)
DOI: 10.1038/s41598-019-53590-8
https://www.nature.com/articles/s41598-019-53590-8
Oskar Fischer & Brigitte Schoenemann
Why are Bones in Vertebrate Eyes? Morphology, Development and Function of Scleral Ossicles in Vertebrate eyes - a Comparative Study
Journal of Anatomy and Physiological Studies 3(2), 1-26 (2019)
2018
Brigitte Schoenemann
Evolution of Eye Reduction and Loss in Trilobites and Some Related Fossil Arthropods
Emerging Science Journal 2, 5, 272-286 (2018)
DOI: 10.28991/esj-2018-01151
www.ijournalse.org/index.php/ESJ/article/download/114/57
2017
Brigitte Schoenemann, Helje Pärnaste & Euan N. K. Clarkson
Structure and function of a compound eye, more than half a billion years old
PNAS 114 (51) 13489-13494
DOI: 10.1073/pnas.1716824114
www.pnas.org/content/early/2017/11/28/1716824114.full
Brigitte Schoenemann, Euan N. K. Clarkson & Magne Høyberget
Traces of an ancient immune system – how an injured arthropod survived 465 million years ago.
Scientific Reports 7, 40330 (2017)
DOI: 10.1038/srep40330
www.nature.com/articles/srep40330
Sidong Wu, Tao Jiang, Gexiang Zhang, Brigitte Schoenemann, Ferrante Neri, Ming Zhu, Chunguang Bu, Jianda Han & Klaus-Dieter Kuhnert
Artificial compound eye: a survey of the state-of-the-art
Artificial Intelligence Review 48, 573–603 (2017)
DOI: 10.1007/s10462-016-9513-7
link.springer.com/article/10.1007/s10462-016-9513-7
Vizcaíno S. F., Clarkson E.N.K. & Schoenemann B. (Editors)
Form, Function and Palaeobiology.
Special Issue Transactions of the Royal Society of Edinburgh 106(4), 207, 2017
DOI: 10.1017/S1755691016000244
https://www.cambridge.org/core/services/aop-cambridge-core/content/view/1BECB5603283FCEC86A6767CA4B705FF/S1755691016000244a.pdf/form_function_and_palaeobiology_preface.pdf
Schoenemann B. & Clarkson E.N.K.
Vision in fossilised eyes.
Special Issue Transactions of the Royal Society of Edinburgh 106(4), 209-220, 2017
DOI: 10.1017/S1755691016000232
www.cambridge.org/core/services/aop-cambridge-core/content/view/64E008DCF10488F6C77B07E29BD22293/S1755691016000232a.pdf/vision_in_fossilised_eyes.pdf
2016
Vannier J.*, Schoenemann B.* , Gillot T., Charbonnier S.
amp; Clarkson E.N.K.
* contributed equally
Exceptional preservation of eye structure in arthropod visual predators from the Middle Jurassic.
Nature Communications 7, 10320
DOI: 10.1038/ncomms10320
www.nature.com/ncomms/2016/160119/ncomms10320/full/ncomms10320.html
Schoenemann B (2016)
A Miniscule System of a ‘Sailing Rock’.
Journal of Geology & Geophysics 5:256
DOI: 10.4172/2381-8719.1000256
www.longdom.org/open-access/a-miniscule-system-of-a-8216sailing-rock8217-40836.html
Poschmann M., Schoenemann B. & McCoy V.
Telltale eyes: The lateral visual systems of Rhenish Lower Devonian eurypterids
(Arthropoda, Chelicerata) and their palaeobiological Implications.
Palaeontology 59(2), 295–304 (2016)
DOI: 10.1111/pala.12228
http://onlinelibrary.wiley.com/enhanced/doi/10.1111/pala.12228
Rust J., Bergmann A., Bartels C., Schoenemann B., Sedlmeier S., & Kühl G.
The Hunsrück biota: A unique window into the ecology of Lower Devonian arthropods.
Arthropod Structure & Development 45, 2, 2016, 140-151
DOI: 10.1016/j.asd.2016.01.004
http://dx.doi.org/10.1016/j.asd.2016.01.004
2015
Schoenemann B., Clarkson E.N.K., Horváth G.
Why did the UV-A-induced photoluminescent blue-green glow in trilobite eyes and exoskeletons
not cause problems for trilobites?
PeerJ 3:e1492
DOI: 10.7717/peerj.1492
https://peerj.com/articles/1492/
Invited Book Review
Schoenemann B.
Visual Ecology by Cronin, T.W., Johnsen, S., Marshall, N.J., Warrant, E.J.,
published by Princeton University Press
Anatomy & Physiology: CurrentResearch 2015, 5:3
DOI: 10.4172/2161-0940.1000180
www.longdom.org/open-access/visual-ecology-2161-0940-1000180.pdf
Tanaka G., Schoenemann B., El Hariri K., Ono T., Clarkson E., Maeda H.
Vision in a Middle Ordovician trilobite eye.
Palaeogeography, Palaeoclimatology, Palaeoecology 433, 2015, 129–139
DOI: 10.1016/j.palaeo.2015.05.013
http://dx.doi.org/10.1016/j.palaeo.2015.05.013
Schoenemann B. & Clarkson E.N.K.
Eyes and vision in the coeval Furongian trilobites Sphaerophthalmus alatus (Boeck, 1938)
and Ctenopyge (Mesoctenopyge) tumida Westergård, 1922, from Bornholm, Denmark.
Palaeontology 58(1), 133–140 (2015)
DOI: 10.1111/pala.12128/epdf
http://onlinelibrary.wiley.com/doi/10.1111/pala.12128/epdf
2014
Schoenemann B. & Clarkson E.N.K.
Space, Time and their Relevance to Phylogenetic Analyses (Convergences in Time).
Journal of Geology & Geophysics 3:178
DOI: 10.4172/2329-6755.1000178
www.longdom.org/open-access/space-time-and-their-relevance-to-phylogenetic-analyses-convergences-in-time-39703.html
Stewart S.E., Clarkson E.N.K., Ahlgren J., Ahlberg P. & Schoenemann B.
Sphenothallus from the Furongian (Cambrian) of Scandinavia.
GFF 137(1), 20–24.
DOI: 10.1080/11035897.2014.929173
www.tandfonline.com/doi/full/10.1080/11035897.2014.929173
Schoenemann B., Clarkson E.N.K. & Ryck U.
Colour Patterns in Devonian Trilobites.
The Open Geology Journal 8, 113-117 (2014)
DOI: 10.2174/1874262901408010113
http://benthamopen.com/contents/pdf/TOGEOJ/TOGEOJ-8-113.pdf
Schoenemann B., Clarkson E.N.K., Castellani C., Waloszek D., Maas A. & Meyer-Rochow B.V.
Description and Interpretation of the Internal Structure of a Cambrian Crustacean Compound Eye.
Bulletin of Geosciences 89(2), 311–324 (2014)
DOI: 10.3140/bull.geosci.1429
www.geology.cz/bulletin/contents/art1429
2013
Schoenemann B. & Clarkson E.N.K.
Discovery of some 400 million year-old sensory
structures in the compound eyes of trilobites.
Scientific Reports 3:1429
DOI: 10.1038/srep01429
www.nature.com/srep/2013/140313/srep01429/full/srep01429.html
related:
ScienceNOW: Looking a Trilobite in the Eye
(Lizzie Wade)
http://news.sciencemag.org/sciencenow/2013/03/looking-a-trilobite-in-the-eye.html?ref=hp
Schoenemann B.
The eyes of a tiny ‘Orsten’ crustacean – A compound eye at receptor level?
Vision Research 76, 89-93 (2013)
DOI: 10.1016/j.visres.2012.10.013
http://www.sciencedirect.com/science/article/pii/S0042698912003392
Parker A., Schoenemann B., Haug J.T. & Waloszek D.
An unusual
cornea from a well-preserved (‘Orsten’) Cambrian compound eye.
Palaeontological Research 17(3), 251-260 (2013)
DOI: 10.2517/1342-8144-17.3.251
http://www.bioone.org/doi/abs/10.2517/1342-8144-17.3.251
2012
Schoenemann B.
The clever strategy of a tiny crustacean
eye early in the evolution of vision.
Communicative & Integrative Biology 5(2), 175–178 (2012).
DOI: 10.4161/cib.18760
www.tandfonline.com/doi/full/10.4161/cib.18760
Schoenemann B. & Clarkson E.N.K.
At First Sight – Functional Analysis of Lower Cambrian eye systems.
Palaeontographica A 297(5-6), 123-149 (2012).
DOI: 10.1127/pala/297/2012/123
https://www.schweizerbart.de/papers/pala/detail/297/85587/At_first_sight_Functional_Analysis_of_Lower_Cambrian_Eye_Systems
Schoenemann B. & Clarkson E.N.K.
Compound Eyes in the Chengjiang
Biota.
Scientific Papers University of Latvia, Earth and Environmental Sciences 783, 57-71 (2012).
https://www.apgads.lu.lv/fileadmin/user_upload/lu_portal/apgads/PDF/LUR-783_Zemes-Vides-zin.pdf
Schoenemann B. & Clarkson E.N.K.
Insights to Phacopid Eyes.
Scientific Papers University of Latvia, Earth and Environmental Sciences 783, 72-75 (2012).
https://www.apgads.lu.lv/fileadmin/user_upload/lu_portal/apgads/PDF/LUR-783_Zemes-Vides-zin.pdf
Schoenemann B. & Clarkson E.N.K.
The eyes of Leanchoilia.
Lethaia 45, 524–531 (2012).
DOI: 10.1111/j.1502-3931.2012.00313.x
http://onlinelibrary.wiley.com/enhanced/doi/10.1111/j.1502-3931.2012.00313.x/
Schoenemann, B., Castellani, C., Clarkson, E. N. K., Haug, J. T.,
Maas, A., Haug, C. & Waloszek, D.
The sophisticated visual system of a tiny Cambrian crustacean: analysis of a stalked fossil compound eye.
Proceedings of the Royal Society London B 279, 1335–1340 (2012).
DOI: 10.1098/rspb.2011.1888
http://rspb.royalsocietypublishing.org/content/early/2011/10/21/rspb.2011.1888.full.pdf+html
related:
NATIONAL GEOGRAPHIC: Fossil eyes show wraparound three-dimensional vision,
half a billion years ago.
(Ed Yong)
http://phenomena.nationalgeographic.com/2011/11/01/fossil-eyes-show-wraparound-three-dimensional-vision-half-a-billion-years-ago/
Castellani, C., Haug, J. T., Haug, C., Maas, A., Schoenemann, B. &
Waloszek, D.
Exceptionally well-preserved isolated eyes from Cambrian ‘Orsten’ fossil assemblages of Sweden.
Palaeontology 55, 553–566 (2012).
DOI: 10.1111/j.1475-4983.2012.01153.x
http://onlinelibrary.wiley.com/doi/10.1111/j.1475-4983.2012.01153.x/abstract
2011
Schoenemann B. & Clarkson E.N.K.
Eyes and vision in the Chengjiang arthropod Isoxys indicating adaptation to habitat.
Lethaia 44(2), 223-230 (2011).
DOI: 10.1111/j.1502-3931.2010.00239.x
http://onlinelibrary.wiley.com/enhanced/doi/10.1111/j.1502-3931.2010.00239.x/
Schoenemann B. & Clarkson E.N.K.
A light guide lens in ancient trilobites?
Transactions of the Royal Society of Edinburgh
102, 17-23 (2011).
DOI: 10.1017/S1755691011010097
http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=8346763&fulltextType=RA&fileId=S1755691011010097
Schoenemann B. & Clarkson E.N.K.
The Eyes of Bohemian Trilobites.
Geologické výzkumy na Moravě a ve Slezsku 18, 1, 45-50 (2011).
https://journals.muni.cz/gvms/article/download/1682/1318
2010
Schoenemann B. & Clarkson E.N.K.
The Sophisticated Strategy of
a Tiny Eye.
Zitteliana B, München 29, 92 (2010).
http://epub.ub.uni-muenchen.de/12021/1/zitteliana_2010_b29.pdf
Schoenemann B. & Clarkson E.N.K.
Neue Einblicke in das visuelle
System alter Augen – Die Augen von Phacopiden (Trilobita).
Zitteliana B, München 29, 92-93 (2010).
http://epub.ub.uni-muenchen.de/12021/1/zitteliana_2010_b29.pdf
Schoenemann B., Clarkson E.N.K., Ahlberg P. & Dies Alvarez M. E.
A Tiny Eye Indicating a Planctonic Trilobite.
Palaeontology 53, 695-701 (2010).
DOI: 10.1111/j.1475-4983.2010.00966.x
http://onlinelibrary.wiley.com/enhanced/doi/10.1111/j.1475-4983.2010.00966.x/
related:
NATIONAL GEOGRAPHIC: Tiny Trilobites Drifted in Cambrian Currents.
(Brian Switek)
http://phenomena.nationalgeographic.com/2010/06/21/tiny-trilobites-drifted-in-cambrian-currents
2009
Schoenemann B., Liu J.-n., Shu D.-g., Han J.-a. & Zhang Z.-f.
A Miniscule Optimised Visual System in the Lower Cambrian.
Lethaia 42, 265–273 (2009).
DOI: 10.1111/j.1502-3931.2008.00138.x
http://onlinelibrary.wiley.com/enhanced/doi/10.1111/j.1502-3931.2008.00138.x/
2008
Schoenemann B.
Analyse früher Augensysteme.
Freiberger Forschungsberichte, Freiberg 15, 85-96 (2008).
Schoenemann, Brigitte
Fossile Augensysteme: Bericht über zur Zeit durchgeführte Analysen fossiler Augensysteme
Paläontologie, Stratigraphie, Fazies ; H. 15 (2007)
Freiberger Forschungshefte / C;524 (2007)
Schoenemann B. & Clarkson E.N.K.
Did the trabecula in phacopid lenses act as light-guides?
In: Advances in Trilobite Research (Edts. Rabano I., Gozalo R. & Garcia-Belido D.)
Cuadernos del Museo Geominero, nº 9. Instituto Geológico y Minero de España, Madrid 351-355 (2008)
DOI: 10.13140/2.1.1806.9283
https://www.researchgate.net/publication/271769459_Advances_in_Trilobite_Research
Schoenemann B., Clarkson E.N.K. & Franz A.
Sublensar Capsules in Trilobite Eyes.
In: Advances in Trilobite Research (Edts. Rabano I., Gozalo R. & Garcia-Beliodo D.)
Cuadernos del Museo Geominero, nº 9. Instituto Geológico y Minero de España, Madrid 356-361 (2008)
DOI: 10.13140/2.1.1806.9283
https://www.researchgate.net/publication/271769459_Advances_in_Trilobite_Research
Schoenemann B., Clarkson E.N.K., Ahlberg P. & Dies Alvarez M. E.
A Furongian Polymerid Planktonic Trilobite.
In: Advances in Trilobite Research (Edts. Rabano I., Gozalo R. & Garcia-Beliodo D.)
Cuadernos del Museo Geominero, nº 9. Instituto Geológico y Minero de España, Madrid 361-365 (2008)
DOI: 10.13140/2.1.1806.9283
https://www.researchgate.net/publication/271769459_Advances_in_Trilobite_Research
2007
Schoenemann B.
Trilobite Eyes and a New Type of Neuronal Superpositional Eye in an Ancient System.
Palaeontographica A
281, 63-91 (2007).
DOI: 10.1127/pala/281/2007/63
https://www.schweizerbart.de/papers/pala/detail/281/87794/Trilobite_Eyes_and_a_New_Type_of_Neural_Superposition_Eye_in_an_Ancient_System?af=search
2006
Schoenemann B.
Cambrian View.
Palaeoworld
15, 307-314 (2006).
DOI: 10.1016/j.palwor.2006.10.012
http://www.sciencedirect.com/science/article/pii/S1871174X06000448
2005
Schoenemann B.
The Cambrian Sight of View
– Suggestions to Investigate Eyes of the Chengjiang Fauna by Comparison
with Recent Visual Systems.
Acta Micropalaeontologica Sinica 22, 169-170 (2005).
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