Heterostracan vertebrates and the Great Eodevonian Biodiversification Event—an essay

Abstract

The oldest vertebrates are Early Cambrian, cephalized unossified species (craniates) from China. The oldest armoured species (euvertebrates) are Ordovician in age. After Talimaa’s Gap, vertebrates have their first adaptive radiation during the Silurian when jawless species (“ostracoderms”) are dominant and their second radiation during the Devonian when jawed species (gnathostomes), and particularly placoderms (armoured fishes), are dominant. A Lochkovian peak of diversity is registered in various Lower Devonian siliciclastic series all around the Old Red Sandstone Continent and Siberia, for ostracoderms in general, and heterostracan pteraspidomorphs in particular. It occurs at different time slices in the Lochkovian, depending on the localities, and may be followed by another smaller peak in the Pragian. Both events correspond to the rise of the Devonian Nekton Revolution as defined for marine invertebrates. This appears to be the second main biodiversification event in the Palaeozoic, following the Great Ordovician Biodiversification Event or GOBE, when euvertebrates appeared. Taking into account most recent palaeobiological studies on heterostracans that suggest they were microphagous suspension feeders or feeding upon microscopic epiphytes from filamentous algae, the origin of this Great Eodevonian Biodiversification Event (GEBE) of heterostracans is questioned. Both abiotic (sea level, tectonic events, climatic changes—oceanic oxygenation and temperature, continental surface temperature) and biotic (plankton diversity, marine primary productivity, competition with vertebrates and invertebrates, including eurypterids, macroecological turnover) factors are examined. No plausible global evolutionary scenario seems to be presently available.

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References

  1. Aberhan, M., & Kiessling, W. (2012). Phanerozoic marine biodiversity: a fresh look at data, methods, patterns and processes. In J. A. Talent (Ed.), Earth and Life: Global biodiversity, extinction intervals and biogeographic perturbations through time (pp. 3–22). Dordrecht: Springer Verlag, IYPE Series.

    Google Scholar 

  2. Allmon, W. D., & Martin, R. E. (2014). Seafood through time revisited: the Phanerozoic increase in marine trophic resources and its macroevolutionary consequences. Paleobiology, 40(2), 256–287.

    Google Scholar 

  3. Alroy, J. (2010). The shifting balance of diversity among major marine animal groups. Science, 329, 1191–1194.

    Google Scholar 

  4. Alroy, J., Aberhan, M., Bottjer, D. J., Foote, M., Fürsich, F. T., Harries, P. J., Hendy, A. J. W., Holland, S. M., Ivany, L. C., Kiessling, W., Kosnik, M. A., Marshall, C. R., McGowan, A. J., Miller, A. I., Olszewski, T. D., Patzkowsky, M. E., Peters, S. E., Villier, L., Wagner, P. J., Bonuso, N., Borkow, P. S., Brenneis, B., Clapham, M. E., Fall, L. M., Ferguson, C. A., Hanson, V. L., Krug, A. Z., Layou, K. M., Leckey, E. H., Nürnberg, S., Powers, C. M., Sessa, J. A., Simpson, C., Tomasovych, A., & Visaggi, C. C. (2008). Phanerozoic trends in the global diversity of marine invertebrates. Science, 321, 97–100. doi:10.1126/science.1156963.

    Article  Google Scholar 

  5. Bambach, R. K. (2002). Supporting predators: changes in the global ecosystem inferred from changes in predator diversity. Paleontological Society Papers, 8, 319–351.

    Google Scholar 

  6. Benton, M. J. (Ed.). (1993). The Fossil Record 2. London: Chapman and Hall. 845 p.

    Google Scholar 

  7. Blieck, A., (1984). Les Hétérostracés Ptéraspidiformes, Agnathes du Silurien-Dévonien du Continent nord-atlantique et des blocs avoisinants: révision systématique, phylogénie, biostratigraphie, biogéographie. Cahiers de Paléontologie (Vertébrés), 199 p.

  8. Blieck, A. (1985). Paléoenvironnements des Hétérostracés, Vertébrés agnathes ordoviciens à dévoniens. In J.-C Fischer. organ., Journées d’étude sur les indicateurs paléobiologiques de milieux (RCP 641, Paris, 26–27 mars 1984). Bulletin du Muséum National d’Histoire Naturelle, 4e série, 7, C (2), 143–155.

  9. Blieck, A. (2011). The André Dumont medallist lecture — From adaptive radiations to biotic crises in Palaeozoic vertebrates: a geobiological approach. Geologica Belgica, 14(3–4), 203–227; also World Wide Web address: http://popups.ulg.ac.be/Geol/docannexe.php?id=3426.

  10. Blieck, A. (2012). Du plus ancien insecte au plus ancien tétrapode. Science & pseudo-sciences, Chroniques: Du côté de la science, 29 sept. 2012; World Wide Web address: http://www.pseudo-sciences.org/spip.php?article1935.

  11. Blieck, A. (2015). An Early Devonian peak of biodiversity: the case of heterostracan vertebrates. In Mottequin, B., Denayer, J., Königshof, P., Prestianni, C., & Olive, S. (Eds.), IGCP 596 – SDS Symposium: Climate change and biodiversity patterns in the Mid-Palaeozoic (Brussels, Sept. 20–22, 2015). Strata, Série 1: communications, vol. 16, 16–17.

  12. Blieck, A., & Elliott, D.K. (2016). Pteraspidomorphs (Vertebrata), the Old Red Sandstone, and the special case of the Brecon Beacons National Park, Wales, U.K. In Howe, S.R. organ., [ORS Symposium Volume]. Proceedings of the Geologists’ Association; doi:10.1016/j.pgeola.2016.07.003 [Online publication: 6-AUG-2016].

  13. Blieck, A., & Heintz, N. (1983). The Cyathaspids of the Red Bay Group (Lower Devonian) of Spitsbergen (The Downtonian and Devonian vertebrates of Spitsbergen: XIV). Polar Research, 1, n.s(1), 49–74.

    Google Scholar 

  14. Blieck, A., & Styza, A. (2014). Devonian sandstones from the Liévin shaft n° 8, Avion (Pas-de-Calais, France): historical context and additional vertebrate rema. Annales de la Société Géologique du Nord, 21(2e série), 25–33.

    Google Scholar 

  15. Blieck, A., Goujet, D., & Janvier, P. (1987). The vertebrate stratigraphy of the Lower Devonian (Red Bay Group and Wood Bay Formation) of Spitsbergen. Modern Geology, 11(3), 197–217.

    Google Scholar 

  16. Blieck, A., Mark-Kurik, E., & Märss, T. (1988). Biostratigraphical correlations between Siluro-Devonian invertebrate-dominated and vertebrate-dominated sequences: the East Baltic example. In McMillan, N.J., Embry, A.F., & Glass, D.J. (Eds.), Devonian of the World (Proceedings of the IInd International Symposium on the Devonian System, Calgary, 1987). Canadian Society of Petroleum Geologists, Memoir 14, vol. III, 579–587.

  17. Blieck, A., Goujet, D., Janvier, P., & Meilliez, F. (1995). Revised Upper Silurian-Lower Devonian ichthyostratigraphy of northern France and southern Belgium (Artois-Ardenne). In Arsenault, M., Lelièvre, H., & Janvier, P. (Eds.), Etudes sur les Vertébrés inférieurs (VIIe Symposium International, Parc de Miguasha, Québec, 9–22 Juin 1991). Bulletin du Muséum National d’Histoire Naturelle, 4e série, 17, C (1–4), 447–459; also World Wide Web address: http://www.mnhn.fr/publication/geodiv/g95n1som.html.

  18. Blieck, A., & Cloutier, R., with contributions by Elliott, D.K., Goujet, D., Loboziak, S., Reed, R.C., Rodina, O., Steemans, P., Valiukevičius, J.J., V’yushkova, L., Yolkin, E.A., & Young, V.T. (2000). Biostratigraphical correlations of Early Devonian vertebrate assemblages of the Old Red Sandstone Continent. In Blieck, A., & Turner, S. (Eds.), Palaeozoic Vertebrate Biochronology and Global Marine/Non-Marine Correlation — Final Report of IGCP 328 (1991–1996). Courier Forschungsinstitut Senckenberg, 223, 223–269.

  19. Blieck, A.R.M., Karatajūtė-Talimaa, V.N., & Mark-Kurik, E. (2002). Upper Silurian and Devonian heterostracan pteraspidomorphs (Vertebrata) from Severnaya Zemlya (Russia): a preliminary report with biogeographical and biostratigraphical implications. Geodiversitas, 24 (4), 805–820; also World Wide Web address: http://www.mnhn.fr/publication/geodiv/g02n4a6.html.

  20. Brett, C.E., & Walker, S.E. (2002). Predators and predation in Palaeozoic marine environments. In Kowalewski, M., & Kelley, P.H. (Eds.), The Fossil Record of Predation. Paleontological Society Papers, 8, 93–118.

  21. Broad, D. S., & Dineley, D. L. (1973). Torpedaspis, a new Upper Silurian and Lower Devonian genus of Cyathaspididae (Ostracodermi) from Arctic Canada. Geological Survey of Canada, Bulletin, 222, 52–91.

  22. Butterfield, N.J. (2011). Was the Devonian radiation of large predatory fish a consequence of rising atmospheric oxygen concentration ? Proceedings of the National Academy of Sciences of the United States of America, 108 (9), E28; doi:10.1073/pnas.1018072108.

    Google Scholar 

  23. Carr, R.K., & Jackson, G.L. (2009). The vertebrate fauna of the Cleveland Member (Famennian) of the Ohio Shale. In Guide to the Geology and Paleontology of the Cleveland Member of the Ohio Shale (68th Annual Meeting of the Society of Vertebrate Paleontology, Cleveland, Ohio, Oct. 15–18, 2008). Ohio Geological Survey, Guidebook 22, 17 p.

  24. Cowen, R. (2003). Diversity of life through time, including Sepkoski’s analysis: mini-essay. University of California, Davis campus (UC Davis), Earth and Planetary Sciences, MyGeologyPage, Richard Cowen’s Home Page: 10 p.; World Wide Web address: http://mygeologypage.ucdavis.edu/cowen/~GEL107/PTect.html.

  25. Dahl, T. W., Hammarlund, E. U., Anbar, A. D., Bond, D. P. G., Gill, B. C., Gordon, G. W., Knoll, A. H., Nielsen, A. T., Schovsbo, N. H., & Canfield, D. E. (2010). Devonian rise in atmospheric oxygen correlated to the radiations of terrestrial plants and large predatory fish. Proceedings of the National Academy of Sciences of the United States of America, 107(42), 17911–17915. doi:10.1073/pnas.1011287107.

    Article  Google Scholar 

  26. Dineley, D. L. (1976). New species of Ctenaspis (Ostracodermi) from the Devonian of Arctic Canada. In C. S. Churcher (Ed.), Athlon - Essays on Palaeontology in Honour of Loris Shano Russell (pp. 26–44). Toronto: Royal Ontario Museum.

    Google Scholar 

  27. Donoghue, P. C. J., Forey, P. L., & Aldridge, R. J. (2000). Conodont affinity and chordate phylogeny. Biological Reviews, 75, 191–251.

    Google Scholar 

  28. Dupret, V. (2010). Revision of the genus Kujdanowiaspis Stensiö, 1942 (Placodermi, Arthrodira, “Actinolepida”) from the Lower Devonian of Podolia (Ukraine). Geodiversitas, 32(1), 5–63. doi:10.5252/g2010n1a1.

    Article  Google Scholar 

  29. Dupret, V., & Blieck, A. (2009). The Lochkovian-Pragian boundary in Podolia (Lower Devonian, Ukraine) based upon placoderm vertebrates. Comptes Rendus Geoscience, 341(1), 63–70. doi:10.1016/j.crte.2008.09.007.

    Article  Google Scholar 

  30. Elliott, D. K., & Blieck, A. R. M. (2010). A new ctenaspid (Agnatha, Heterostraci) from the Early Devonian of Nevada, with comments on taxonomy, paleobiology and paleobiogeography. In D. K. Elliott, J. G. Maisey, X. Yu, & D. Miao (Eds.), Morphology, Phylogeny and Paleobiogeography of Fossil Fishes – Honoring Meemann Chang (pp. 25–38). München: Verlag Dr. Friedrich Pfeil.

    Google Scholar 

  31. Elliott, D. K., & Petriello, M. A. (2011). New poraspids (Agnatha, Heterostraci) from the Early Devonian of the western United States. Journal of Vertebrate Paleontology, 31(3), 518–530.

    Google Scholar 

  32. Elliott, D. K., Reed, R. C., & Loeffler, E. J. (2004). A new species of Allocryptaspis (Heterostraci) from the Early Devonian, with comments on the structure of the oral area in cyathaspidids. In G. Arratia, M. V. H. Wilson, & R. Cloutier (Eds.), Recent Advances in the Origin and early Radiation of Vertebrates – Honoring Hans-Peter Schultze (pp. 455–472). Verlag Dr. Friedrich Pfeil: München.

    Google Scholar 

  33. Elliott, D. K., Schultze, H.-P., & Blieck, A. (2015). A new pteraspid (Agnatha, Heterostraci) from the Lower Devonian Drake Bay Formation, Prince of Wales Island, Nunavut, Arctic Canada, and comments on environmental preferences of pteraspids. Journal of Vertebrate Paleontology, 35(6), e1005098. doi:10.1080/02724634.2015.1005098. 10 p.

    Article  Google Scholar 

  34. Friedman, M., & Sallan, L. C. (2012). Five hundred million years of extinction and recovery: a Phanerozoic survey of large-scale diversity patterns in fishes. Palaeontology, 55(4), 707–742.

    Google Scholar 

  35. Gensel, P. G. (2008). The earliest land plants. Annual Review of Ecology, Evolution, and Systematics, 39, 459–477.

    Google Scholar 

  36. George, D., & Blieck, A. (2011). Rise of the earliest tetrapods: An Early Devonian origin from marine environment. PLoS ONE, 6(7), 1–7. doi:10.1371/journal.pone.0022136.

    Google Scholar 

  37. Gornitz, V. (Ed.). (2009). Encyclopedia of Paleoclimatology and Ancient Environments. Dordrecht: Springer Netherlands. xxviii + 1049 p.

    Google Scholar 

  38. Grahn, Y., & Paris, F. (2011). Emergence, biodiversification and extinction of the chitinozoan group. Geological Magazine, 148(2), 226–236.

    Google Scholar 

  39. Gross, W. (1937). Die Wirbeltiere des rheinischen Devons. Teil II. Abhandlungen der Preußischen Geologischen Landesanstalt, N. F. 176, 83 p.

  40. Gross, W. (1963). Drepanaspis gemuendenensis Schlüter. Neuuntersuchung. Palaeontographica A, 121(4–6), 133–155.

    Google Scholar 

  41. Hairapetian, V., Roelofs, B.P.A., Trinajstic, K.M., & Turner, S. (2015). Famennian survivor turiniid thelodonts of North and East Gondwana. In Becker, R.T., Königshof, P., & Brett, C. (Eds.), Devonian Climate, Sea Level and Evolutionary Events. Geological Society of London, Special Publication 423. doi:10.1144/SP423.3, 17 p.

    Google Scholar 

  42. Hansen, M. C., & Mapes, R. H. (1990). A predator–prey relationship between sharks and cephalopods in the Late Palaeozoic. In A. J. Boucot (Ed.), Evolutionary Paleobiology of Behavior and Coevolution (pp. 189–192). Amsterdam: Elsevier.

    Google Scholar 

  43. Haq, B. U., & Schutter, S. R. (2008). A chronology of Paleozoic sea-level changes. Science, 322(5898), 64–68.

    Google Scholar 

  44. Holland, S. M., & Sclafani, J. A. (2015). Phanerozoic diversity and neutral theory. Paleobiology, 41, 369–376.

    Google Scholar 

  45. Janvier, P. (1996). Early Vertebrates. Oxford: Oxford Science Publications and Clarendon Press, Oxford Monographs on Geology and Geophysics, 33. 393 p.

  46. Joachimski, M. M., Breisig, S., Buggisch, W., Talent, J. A., Mawson, R., Gereke, M., Morrow, J. R., Day, J., & Weddige, K. (2009). Devonian climate and reef evolution: Insights from oxygen isotopes in apatite. Earth and Planetary Science Letters, 284(3–4), 599–609.

    Google Scholar 

  47. Kleesment, A., & Mark-Kurik, E. (1997). Lower Devonian. In A. Raukas & A. Teedumäe (Eds.), Geology and Mineral Resources of Estonia (pp. 107–112). Tallinn: Estonian Academy Publishers.

    Google Scholar 

  48. Klug, C., Kröger, B., Kiessling, W., Mullins, G.L., Servais, T., Frýda, J., Korn, D., & Turner, S. (2010). The Devonian nekton revolution. Lethaia, 43, 465–477. doi:10.111/j.1502-3931.2009.00206.x.

  49. Klug, C., Frey, L., Korn, D., Jattiot, R., & Rücklin, M. (2016). The oldest Gondwanan cephalopod mandibles (Hangenberg Black Shale, Late Devonian) and the Mid-Palaeozoic rise of jaws. Palaeontology, Early View Article, 19 p. doi:10.111/pala.12248.

  50. Lamsdell, J. C., & Braddy, S. J. (2010). Cope’s Rule and Romer’s theory: patterns of diversity and gigantism in eurypterids and Palaeozoic vertebrates. Biology Letters, 2010(6), 265–269. doi:10.1098/rsbl.2009.0700.

    Article  Google Scholar 

  51. Le Hérissé, A., Mullins, G. L., Dorning, K. J., & Wicander, R. (2009). Global patterns of organic-walled phytoplankton biodiversity during the Late Silurian to earliest Devonian. Palynology, 33, 25–75.

    Google Scholar 

  52. Le Hir, G., Donnadieu, Y., Godderis, Y., Meyer-Berthaud, B., Ramstein, G., & Blakey, R. C. (2011). The climate change caused by the land plant invasion in the Devonian. Earth and Planetary Science Letters, 310, 203–212.

    Google Scholar 

  53. Lebedev, O.A., Mark-Kurik, E., Karatajūtė-Talimaa, V., Lukševičs, E., & Ivanov, A. (2009). Bite marks as evidence of predation in early vertebrates. In Ahlberg, P.E., Blom, H., & Boisvert, C.A. (Eds.), Forty Years of Early Vertebrates (11th International Symposium on Early and Lower Vertebrates, Uppsala, 2007). Acta Zoologica, Special Issue, Supplement to vol. 90, 344–356.

  54. Long, J. A. (1990). Fishes. In K. J. McNamara (Ed.), Evolutionary Trends (pp. 255–278). London: Belhaven Press.

    Google Scholar 

  55. Long, J. A., Large, R. R., Lee, M. S. Y., Benton, M. J., Danyushevsky, L. V., Chiappe, L. M., Halpin, J. A., Cantrill, D., & Lottermoser, B. (2016). Severe selenium depletion in the Phanerozoic oceans as a factor in three global mass extinction events. Gondwana Research. doi:10.1016/j.gr.2015.10.001. 10 p.

    Article  Google Scholar 

  56. Mallatt, J. (1984). Feeding ecology of the earliest vertebrates. Zoological Journal of the Linnean Society, 82(3), 261–272. doi:10.1111/j.1096-3642.1984.tb00643.x.

    Article  Google Scholar 

  57. Mark-Kurik, J. E. (1966). On some alterations of the exoskeleton of psammosteids (Agnatha). In Organism and its environment in the geological past. Moskva: Nauka, 55–60. [In Russian.]

  58. Mark-Kurik, E., & Põldvere, A. (2012). Devonian stratigraphy in Estonia: current state and problems. Estonian Journal of Earth Sciences, 61(1), 33–47. also World Wide Web address: http://www.kirj.ee/public/Estonian_Journal_of_Earth_Sciences/2012/issue_1/earth-2012-1-33-47.pdf.

    Google Scholar 

  59. Martin, R. E. (1996). Secular Increase in Nutrient Levels through the Phanerozoic: Implications for Productivity, Biomass, and Diversity of the Marine Biosphere. Palaios, 11(3), 209–219.

    Google Scholar 

  60. May, A. (1996). Relationships among sea-level fluctuation, biogeography and bioevents of the Devonian: an attempt to approach a powerful, but simple model for complex long-range control of biotic crises. Geolines, 3(1995), 38–49.

    Google Scholar 

  61. Miles, R. S. (1973). Articulated acanthodian fishes from the Old Red Sandstone of England, with a review of the structure and evolution of the acanthodian shoulder-girdle. Bulletin of the British Museum (Natural History). Geology, 24(2), 113–213.

    Google Scholar 

  62. Nardin, E., Godderis, Y., Donnadieu, Y., Le Hir, G., Blakey, R. C., Puceat, E., & Aretz, M. (2011). Modeling the early Paleozoic long-term climatic trend. Geological Society of America Bulletin, 123(5–6), 1181–1192.

    Google Scholar 

  63. Nestor, V. (2009). Chitinozoan diversity in the East Baltic Silurian. Estonian Journal of Earth Sciences, 58(4), 311–316.

    Google Scholar 

  64. Nikiforova, O. I. (1977). Podolia. In A. Martinsson (Ed.), The Silurian-Devonian Boundary. International Union of Geological Sciences Series A, 5 (pp. 52–64). Stuttgart: E. Schweizerbart Verlag.

    Google Scholar 

  65. Novitskaya, L.I. (2004). Podklass Heterostraci [Subclass Heterostraci]. In L.I. Novitskaya,  & O.B. Afanassieva (Eds.), Iskopaemye pozvonotchnye Rossii i sopredel’nykh stran [Fossil vertebrates of Russia and adjacent countries]: Bestchelyustnye i drevnie ryby [Agnathans and early fishes]. Rossijskaya Akademiya Nauk [Academy of Sciences of Russia], Paleontologitcheskij Institut [Palaeontological Institute]. Moskva: Geos, 69–207. [In Russian].

  66. Novitskaya, L.I. (2007). Evolution of generic and species diversity in agnathans (Heterostraci: Orders Cyathaspidiformes, Pteraspidiformes). Paleontological Journal, 41 (3), 268–280. [Original Russian paper: Paleontologicheskii Zhurnal, 2007 (3), 33–46].

    Google Scholar 

  67. Novitskaya, L.I. (2008). Evolution of taxonomic diversity in amphiaspids (Agnatha, Heterostraci: Amphiaspidiformes) and the causes of extinction in ecologically favorable conditions. Paleontological Journal, 42 (2), 181–191. [Original Russian paper: Paleontologicheskii Zhurnal, 2008 (2), 75–85].

  68. Paris, F., & Nõlvak, J. (1999). Biological interpretation and palaeobiodiversity of a cryptic fossil group : the “chitinozoan animal”. Geobios, 32(2), 315–324.

    Google Scholar 

  69. Paris, F., Achab, A., Asselin, E., Chen, X.-H., Grahn, I., Nolvak, J., Obut, O., Samuelsson, J., Sennikov, N., Vecoli, M., Verniers, J., Wang, X.-F., & Winchester-Seeto, T. (2004). Chitinozoans. In B. D. Webby, F. Paris, M. L. Droser, & I. G. Percival (Eds.), The Great Ordovician Biodiversification Event (pp. 294–311). NewYork: Columbia University Press, Critical Moments and Perspectives in Earth History and Paleobiology.

    Google Scholar 

  70. Paškevičius, J. (1997). The geology of the Baltic Republics. Vilnius: Vilnius University and Geological Survey of Lithuania. 387 p.

    Google Scholar 

  71. Payne, J. L., Boyer, A. G., Brown, J. H., Finnegan, S., Kowalewski, M., Krause, R. A., Jr., Lyons, S. K., McClain, C. R., McShea, D. W., Novack-Gottshall, P. M., Smith, F. A., Stempien, J. A., & Wang, S. C. (2009). Two-phase increase in the maximum size of life over 3.5 billion years reflects biological innovation and environmental opportunity. Proceedings of the National Academy of Sciences of the United States of America, 106(1), 24–27.

    Google Scholar 

  72. Pernègre, V. N. (2005). Description d’une nouvelle espèce et analyse morpho-fonctionnelle du genre Doryaspis White (Heterostraci) du Dévonien du Spitsberg). Geobios, 38(2), 257–268.

    Google Scholar 

  73. Pernègre, V., & Blieck, A. (2016). A revised heterostracan-based ichthyostratigraphy of the Wood Bay Formation (Lower Devonian, Spitsbergen), and correlation with Russian Arctic archipelagos. Geodiversitas, 38(1), 5–20. doi:10.5252/g2016n1a1.

    Article  Google Scholar 

  74. Pernègre, V. N., & Elliott, D. K. (2008). Phylogeny of the Pteraspidiformes (Heterostraci), Silurian-Devonian jawless vertebrates. Zoologica Scripta, 37(4), 391–403.

    Google Scholar 

  75. Phillips, J. (1860). Life on the Earth: Its Origin and Succession. Cambridge and London: MacMillan and Co. (Reprint edition, 1980. New York: Arno Press). (after Scott, 2015).

    Google Scholar 

  76. Plotnick, R.E. (1999). Habitat of Llandoverian-Lochkovian eurypterids. In Boucot, A.J., & Lawson, J.D. (Eds.), Paleocommunities: a case study from the Silurian and Lower Devonian. Cambridge: Cambridge University Press, World and Regional Geology Series, 11, 106–131.

  77. Purnell, M. A. (1995). Microwear on conodont elements and macrophagy in the first vertebrates. Nature, 374(6525), 798–800.

    Google Scholar 

  78. Purnell, M.A. (2001). Scenarios, selection and the ecology of early vertebrates. In Ahlberg, P.E. ed., Major Events in Early Vertebrate Evolution – Palaeontology, phylogeny, genetics and development. Systematics Association Special Volume Series, 61, 187–208.

  79. Purnell, M. A. (2002). Feeding in extinct jawless heterostracan fishes and testing scenarios of early vertebrate evolution. Proceedings of the Royal Society of London, B, 269(1486), 83–88.

    Google Scholar 

  80. Rasmussen, C. M. Ø., Ullmann, C. V., Jakobsen, K. G., Lindskog, A., Hansen, J., Hansen, T., Eriksson, M. E., Dronov, A., Frei, R., Korte, C., Nielsen, A. T., & Harper, D. A. T. (2016). Onset of main Phanerozoic marine radiation sparked by emerging Mid Ordovician icehouse. Scientific Reports, 6, 18884. doi:10.1038/srep18884. 9 p.

    Article  Google Scholar 

  81. Rohde, R. A., & Muller, R. A. (2005). Cycles in fossil diversity. Nature, 434(7030), 208–210.

    Google Scholar 

  82. Romer, A. S. (1933) Eurypterid influence on vertebrate history. Science, 78,(2015), 114–117

    Google Scholar 

  83. Sansom, R. S., Randle, E., & Donoghue, P. C. J. (2015). Discriminating signal from noise in the fossil record of early vertebrates reveals cryptic evolutionary history. Proceedings of the Royal Society B, 282, 20142245. doi:10.1098/rspb.2014.2245. 8 p.

    Article  Google Scholar 

  84. Sallan, L., & Carlton, T. (2015). The rise of fishes after the fall of the Ordovician world. In K. Trinajstic, Z. Johanson, M. Richter, & C. Boisvert (Eds.), 13th International Symposium on Early and Lower Vertebrates. Melbourne: Royal Society of Victoria. August 3–7, 2015. Abstract Volume: 28.

    Google Scholar 

  85. Schobben, M., Stebbins, A., Ghaderi, A., Strauss, H., Korn, D., & Korte, C. (2016). Eutrophication, microbial-sulfate reduction and mass extinctions. Communicative & Integrative Biology. doi:10.1080/19420889.2015.1115162.

    Article  Google Scholar 

  86. Scotese, C.R. (2002). PALEOMAP Project: Plate tectonic maps and continental drift animations. World Wide Web address: http://www.scotese.com; Arlington, Texas [dated 1998–2002; updated 2003].

  87. Scott, M. (2015). John Phillips. Strange Science; World Wide Web address: http://www.strangescience.net/phillips.htm [viewed on April 28, 2016].

  88. Sepkoski, J. J., Jr. (1981). A factor analytic description of the Phanerozoic marine fossil record. Paleobiology, 7(1), 36–53.

    Google Scholar 

  89. Sepkoski, Jr., J.J. (2002). A compendium of fossil marine animal genera. Bulletins of American Paleontology, 363. 560 p.

  90. Slavik, L., Hladil, J., Chadimova, L., Valenzuela-Ríos, J.I., Huškova, A., & Liao, J.-C. (2015). Cooling or warming in the Pragian ? The sedimentary records and petrophysical logs from the key peri-Gondwanan sections. In B. Mottequin, J. Denayer., P. Königshof, C. Prestianni, & S. Olive (Eds.), IGCP 596 – SDS Symposium: Climate change and biodiversity patterns in the Mid-Palaeozoic (Brussels, Sept. 20–22, 2015). Strata, Série 1: communications, vol. 16, 130–131.

  91. Smith, A. B., & McGowan, A. J. (2008). Temporal patterns of barren intervals in the Phanerozoic. Paleobiology, 34(1), 155–161.

    Google Scholar 

  92. Smith, A. B., Lloyd, G. T., & McGowan, A. J. (2012). Phanerozoic marine diversity: rock record modelling provides an independent test of large-scale trends. Proceedings of the Royal Society B, 279, 4489–4495. doi:10.1098/rspb.2012.1793.

    Article  Google Scholar 

  93. Thomson, K. S. (1977). The pattern of diversification among fishes. In A. Hallam (Ed.), Patterns of evolution as illustrated by the fossil record (pp. 377–404). Amsterdam: Elsevier.

    Google Scholar 

  94. Turner, S., Blieck, A., & Nowlan, G.S. (2004). Vertebrates (Agnathans and Gnathostomes). In Webby, B.D., Paris, F., Droser, M.L., & Percival, I.G. (Eds.), The Great Ordovician Biodiversification Event (IGCP 410 volume). New York: Columbia University Press, "Critical moments and perspectives in Earth history and paleobiology", chapter 30, 327–335.

  95. Turner, S., Burrow, C. J., Schultze, H.-P., Blieck, A., Reif, W.-E., Rexroad, C. B., Bultynck, P., & Nowlan, G. S. (2010). False teeth: conodont-vertebrate phylogenetic relationships revisited. Geodiversitas, 32(4), 545–594. also World Wide Web address: http://www.mnhn.fr/museum/front/medias/publication/31374_g2010n4a1.pdf.

    Google Scholar 

  96. Vandenbroucke, T.R.A., Emsbo, P., Munnecke, A., Nuns, N., Duponchel, L., Lepot, K., Quijada, M., Paris, F., Servais, T., & Kiessling, W. (2015). Metal-induced malformations in early Palaeozoic plankton are harbingers of mass extinction. Nature Communications, 6 (7966), 7 p. + Supplementary Information; doi:10.1038/ncomms8966.

  97. Voichyshyn, V. (2011). The Early Devonian armoured agnathans of Podolia, Ukraine. Palaeontologia Polonica, 66, 211 p.; also World Wide Web address: http://www.palaeontologia.pan.pl/PP66/PP66.pdf.

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Acknowledgements

This paper is based on a series of oral communications that have been given in front of IGCP 596–SDS Symposium: Climate change and biodiversity patterns in the Mid-Palaeozoic (Sept. 20–22, 2015, Brussels, Belgium), the 2016 meeting of Association Française de Paléontologie (30 March–2 April 2016, Elbeuf, France), and IGCP 591 Closing meeting: The Early to Mid Palaeozoic Revolution (6–9 July 2016, Ghent, Belgium). The organisers of these meetings are warmly thanked. Several colleagues helped during the making of this paper, viz., Lauren C. Sallan (University of Pennsylvania at Philadelphia, PA, USA), Elise Nardin (Université Paul Sabatier, Toulouse, France), and some others who are cited in the text. They are thanked for this. Both reviewers David K. Elliott (Northern Arizona University, Flagstaff, USA) and Philippe Janvier (CNRS, Muséum National d’Histoire Naturelle, Paris, France) made detailed comments on the “manuscript”, that greatly improved its quality.

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This article is a contribution to the special issue “Climate change and biodiversity patterns in the mid-Palaeozoic”.

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Blieck, A. Heterostracan vertebrates and the Great Eodevonian Biodiversification Event—an essay. Palaeobio Palaeoenv 97, 375–390 (2017). https://doi.org/10.1007/s12549-016-0260-1

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Keywords

  • Early Devonian
  • Nekton Revolution
  • Ostracoderms
  • Palaeobiodiversity
  • Essay in geobiology