Dynamics of abundance of the mid- to late Pridoli conodonts from the eastern part of the Silurian Baltic basin: Multifractals, state shifts, and oscillations

The Pridoli epoch is one of the most geobiologically unstable intervals in the Silurian period. However, the ecological and evolutionary dynamics of the biota from this time are still poorly understood. Here we present a comprehensive quantitative time-series analyses of conodont abundances during the middle and the upper parts of the Pridoli epoch. The study reveals that the stratigraphic series of conodont abundances has a multifractal structure. This feature implies the presence of multiple serially correlated processes hierarchically operating on different time scales and interacting in a multiplicative fashion. Estimation of a maximal Lyapunov exponent and a test for presence of determinism also revealed a chaotic and non-linear low-dimensional nature of long-term conodont abundance dynamics. The most probable (but not exclusive) mechanism for the origin of chaos in the time series of conodont abundance is that conodont abundances tracked changes in crucial environmental parameters, most probably related to the climate that exhibits long term chaos. The complex and non-linear nature of the conodont dynamics is also revealed by a moving-window correlation analyses of conodont abundances and by an environmental proxy time series. We found three alternating episodes of different correlation patterns between conodont abundances and the δ13C record. In the middle and upper most Pridoli interval there are positive but statistically non-significant correlations between these two variables. In contrast the lower part of the upper Pridoli (middle of the Jūra formation) exhibits negative and statistically significant correlation between the variables. In addition moving-window standard deviation analyses reveals that during the transitions between these episodes, there were abrupt changes in the variance in both the carbon isotopes and the conodont abundances, which implies that these episodes represent alternative states in the way the earth system functioned that were punctuated by the critical transitions between them. The interval with negative correlations between abundances and isotopes was also marked by a profound evolutionary turnover of acanthodians and to lesser extent conodonts. This interval most probably represented an anomalous time, with differing ecological rules and increased macroevolutionary rates, which can be interpreted as a turnover pulse. The spectral cyclostratigraphic analyses of the abundance data reveals that conodont abundances also exhibited periodic oscillations at several frequencies. The cycles determined for the analysis have the period lengths of 56, 99, 127 and 950 Ka, which probably were caused by the periodic changes in the obliquity and eccentricity of Earth's orbit. The proposed proximate mechanism that controlled cyclic changes in the abundances of conodonts, based on the climatological deductions, was a variation in nutrients supplies that was due to changes in the degree of seasonality in the tropics, driven by Milankovitch cycles. It is probable that the described state shifts in the way that the ecosystem functioned were also forced by long-term Milankovitch cycles.