Using shells collected from a sediment trap series in the Madeira Basin, we investigate the effects of seasonal variation of temperature, productivity, and optimum growth conditions on calcification in three species of planktonic Foraminifera. The series covers an entire seasonal cycle and reflects conditions at the edge of the distribution of the studied species, manifesting more suitable growth conditions during different parts of the year. The seasonal variation in seawater carbonate saturation at the studied site is negligible compared to other oceanic regions, allowing us to assess the effect of parameters other than carbonate saturation. Shell calcification is quantified using weight and size of individual shells. The size–weight scaling within each species is robust against changes in environmental parameters, but differs among species. An analysis of the variation in calcification intensity (size-normalized weight) reveals species-specific response patterns. In Globigerinoides ruber (white) and Globigerinoides elongatus, calcification intensity is correlated with temperature (positive) and productivity (negative), whilst in Globigerina bulloides no environmental forcing is observed. The size–weight scaling, calcification intensity, and response of calcification intensity to environmental change differed between G. ruber (white) and G. elongatus, implying that patterns extracted from pooled analyses of these species may reflect their changing proportions in the samples. Using shell flux as a measure of optimum growth conditions, we observe significant positive correlation with calcification intensity in G. elongatus, but negative correlation in G. bulloides. The lack of a consistent response of calcification intensity to optimum growth conditions is mirrored by the results of shell size analyses. We conclude that calcification intensity in planktonic Foraminifera is affected by factors other than carbonate saturation. These factors include temperature, productivity, and optimum growth conditions, but the strength and sign of the relationships differ among species, potentially complicating interpretations of calcification data from the fossil record.

Planktonic Foraminifera are important marine calcifiers, and the ongoing change in the oceanic carbon system makes it essential to understand the influence of environmental factors on the biomineralization of their shells. The amount of calcite deposited by planktonic Foraminifera during calcification has been hypothesized to reflect a range of environmental factors. However, it has never been assessed whether their calcification only passively responds to the conditions of the ambient seawater or whether it reflects changes in resource allocation due to physiological stress. To disentangle these two end-member scenarios, an experiment is required where the two processes are separated. A natural analogue to such an experiment occurred during the deposition of the Mediterranean sapropels, where large changes in surface water composition and stratification at the onset of the sapropel deposition were decoupled from local extinctions of planktonic Foraminifera species. We took advantage of this natural experiment and investigated the reaction of calcification intensity, expressed as mean area density (MAD), of four species of planktonic Foraminifera to changing conditions during the onset of Sapropel S5 (126–121 ka) in a sediment core from the Levantine Basin. We observed a significant relationship between MAD and surface water properties, as reflected by stable isotopes in the calcite of Foraminifera shells, but we failed to observe any reaction of calcification intensity on ecological stress during times of decreasing abundance culminating in local extinction. The reaction of calcification intensity to surface water perturbation at the onset of the sapropel was observed only in surface-dwelling species, but all species calcified more strongly prior to the sapropel deposition and less strongly within the sapropel than at similar conditions during the present-day. These results indicate that the high-salinity environment of the glacial Mediterranean Sea prior to sapropel deposition induced a more intense calcification, whereas the freshwater injection to the surface waters associated with sapropel deposition inhibited calcification. The results are robust to changes in carbonate preservation and collectively imply that changes in normalized shell weight in planktonic Foraminifera should reflect mainly abiotic forcing

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Molecular genetic investigations of the highly abundant extant planktonic foraminifera plexus Globigerinella siphonifera/Globigerinella calida have recently shown this group to be the genetically most diverse one within planktonic foraminifera, separating it into 12 distinct genetic types belonging to three main genetic lineages. Independently, several morphological or physiological variants have been described within the group, but the correlation between the high genetic diversity and the phenotypic variability remains unclear. In this study, we combine genetic data with morphometric analyses of shell shape and porosity of genotyped individuals of the different genetic lineages. Our morphometric measurements suggest a differentiation of three morphotypes within the plexus, two of which possess the elongated chambers described as a typical trait of G. calida. These two morphotypes with elongated chambers are associated with two distinct genetic lineages. The G. calida morphology therefore appears to have evolved twice in parallel. Unexpectedly, we show that the two morphotypes with elongated chambers can be separated from each other by characters seen in the lateral view of their shells. This implies that the taxonomy of the extant members of the genus Globigerinella should be revised. A comparison with the original descriptions and type specimens of members of the genus shows that two genetic types of one major lineage correspond to G. calida. The second group with elongated chambers is associated with a recently diverged genetic type and we propose to reinstate the name Globigerinella radians for this distinct form. The remaining nine of the 12 genetic types correspond to the G. siphonifera morphology, and in the absence of evidence for morphological differentiation, they form a paraphyletic morpho-taxon. Our results highlight the prevalence of parallelism in the evolution of shell morphology in planktonic foraminifera even at the lowest level of relatedness represented by genetic types