Silurian-Devonian boundary events and their influence on cephalopod evolution: evolutionary significance of cephalopod egg size during mass extinctions

Diversity evolution of 197 of the latest Ludlow-Lochkovian cephalopod species from Bohemia (representing more that 70 percent of all species known worldwide) was analysed and compared with changes in the global carbon cycle in the marine ecosystem. Our results show a distinct relationship between cephalopod species diversity and the global carbon cycle. A progressive increase in delta C-13 values in marine carbonates from the Monograptus lochkoviensis Zone to the end of the Monograptus transgrediens Zone reflects an increase of bioproductivity in the marine ecosystem. This increase was probably caused by a shallowing of the upwelling system, thus importing nutrient-rich water. A progressive cooling during the Pridoli and earliest Lochkovian triggered these changes in the upwelling system. Our data also show an increase in total diversity of cephalopod species, which may be explained by greater availability of new food sources for a period corresponding to the progressive increase in delta C-13 values. In the latest Silurian, delta C-13 reached its maximal values while the Lochkovian is characterized by steady conditions with a decreasing tendency in delta C-13 values. Strong, selective extinction started in the latest Silurian Monograptus transgrediens Zone and continued to the earliest Devonian Monograptus uniformis Zone. This extinction affected benthic and demersal cephalopod species with relatively large eggs and long incubation times. The change in dynamics of the global carbon cycle thus coincides with the beginning of this extinction. An anoxic or hypoxic event in the period between the LAD Monograptus transgrediens Zone (Transgrediens Event) in the latest Pridoli and the base of the Devonian Monograptus uniformis Zone (documented also from offshore sequences from Poland) is considered to be a cause for the extinction of non-pelagic cephalopods. Our study also revealed a need to revise and define all earlier established bioevents in the Silurian-Devonian boundary interval more clearly.