Quantifying the effect of seasonal and vertical habitat tracking on planktonic foraminifera proxies

The composition of planktonic foraminiferal (PF) calcite is routinely used to reconstruct climate variability. However, PF ecology leaves a large imprint on the proxy signal: seasonal and vertical habitats of PF species vary spatially, causing variable offsets from annual mean surface conditions recorded by sedimentary assemblages. PF seasonality changes with temperature in a way that minimises the environmental change that individual species experience and it is not unlikely that changes in depth habitat also result from such habitat tracking. While this behaviour could lead to an underestimation of spatial or temporal trends as well as of variability in proxy records, most palaeoceanographic studies are (implicitly) based on the assumption of a constant habitat. Up to now, the effect of habitat tracking on foraminifera proxy records has not yet been formally quantified on a global scale. Here we attempt to characterise this effect on the amplitude of environmental change recorded in sedimentary PF using core top delta O-18 data from six species. We find that the offset from mean annual near-surface delta O-18 values varies with temperature, with PF delta O-18 indicating warmer than mean conditions in colder waters (on average by -0.1%(equivalent to 0.4 degrees C) per degrees C), thus providing a firstorder quantification of the degree of underestimation due to habitat tracking. We use an empirical model to estimate the contribution of seasonality to the observed difference between PF and annual mean delta O-18 and use the residual Delta delta O-18 to assess trends in calcification depth. Our analysis indicates that given an observation-based model parametrisation calcification depth increases with temperature in all species and sensitivity analysis suggests that a temperature-related seasonal habitat adjustment is essential to explain the observed isotope signal. Habitat tracking can thus lead to a significant reduction in the amplitude of recorded environmental change. However, we show that this behaviour is predictable. This allows accounting for habitat tracking, enabling more meaningful reconstructions and improved data-model comparison.