Where should we draw the lines between dinocyst “species”? Morphological continua in Black Sea dinocysts

Abstract. The morphology of dinoflagellate cysts (dinocysts) is related not only to the genetics of the motile dinoflagellate from which it derives, but is also dependent on a range of environmental factors including salinity, temperature and nutrient status. Although this knowledge improves our understanding of the drivers behind dinocyst morphological variations, it makes the taxonomy governing their description somewhat complex. In basins such as the Black Sea, where environmental change can be extreme and occurs on relatively short (millennial) timescales, taxonomy becomes particularly challenging. Morphological continua can be observed between described forms, displaying a large range of intermediate phenotypes that do not necessarily correspond to any genetic difference. As these morphological nuances may preserve information about palaeoenvironments, it is important to find a systematic method of characterising morphotypes. Here, we show a dinocyst matrix within which dinocysts are described according to their similarity to (or difference from) described forms based on key descriptive parameters. In the example set out here, cyst shape and degree of process and/or ectophragm development are taken as two key parameters in Pyxidinopsis psilata and Spiniferites cruciformis, and can allow the description of intermediate forms even though the definitions do not overlap. We review some frequently occurring morphotypes and propose that using matrices to show the gradual variation between endmember forms is the most pragmatic approach until cyst–theca studies and genetic sequencing can be used to demonstrate relationships between genotypes and morphotypes. As prior studies propose salinity to be a primary driver of intraspecific variability, the endmembers presented may represent salinity extremes within an overall brackish environment. Although we cannot assign each morphotype to a value or a range of an environmental parameter (e.g. salinity) as the different morphotypes can occur in the same sample, using this matrix allows preservation of information about morphological variability without creating taxonomic categories that are likely to require alteration if genetic evidence becomes available.