Review: Freshwater and Soil Foraminifera – A Story of Long-Forgotten Relatives

ABSTRACT Foraminifera are a primarily marine taxon widespread in all oceanic habitats, from shallow, brackish-water settings to deep-seafloor and pelagic realms. Their diversity is remarkable with several thousand species described and a fossil record tracing back to the Cambrian. While foraminifera represent one of the best-studied groups of marine meiofauna, much less is known about their non-marine relatives. The first freshwater foraminifera were described in the 19th century by European and North American protozoologists, but interest in them lapsed during much of the 20th century and was not rekindled until the advent of molecular systematics provided a fresh impetus to their study. Several new species, genera, and families have been described recently based on morphological and molecular data derived from cultured specimens. In parallel, environmental genomic studies revealed that foraminifera are highly diverse and ubiquitous in freshwater and soil environments. Molecular phylogenetic analyses places non-marine foraminifera in a few clades among the large array of single-chambered (monothalamous) lineages, suggesting that several independent colonization events of freshwater and terrestrial habitats occurred. Non-marine foraminifera are turning from obscure curiosities to being recognized as an important part of soil and freshwater microbial communities, a major component of these complex environments.