Palaeozoic Foraminifera: Systematics, palaeoecology and responses to global changes

Because the Foraminifera are very sensitive to various environmental parameters (e.g., water temperature, salinity, light, etc.), there are important proxies used for palaeoenvironmental and palaeogeographic reconstructions. The evolution of the structure, shape and size of the mineralized tests of Foraminifera can directly reflect the variation of these parameters through geological time. Furthermore, their biostratigraphic value has been widely demonstrated. In this context, the systematics, evolution and ecological behaviour of the first mineralized Palaeozoic Foraminifera are important to discuss in order to have a clearer picture of former shallow marine environments, and finally understand their distribution through space and time. The systematics of the fossil group of Foraminifera that first developed a mineralized test remains under discussion. These early foraminifers are considered as Textulariata (as generally admitted), recrystallized Fusulinata or an independent group, sometimes called Astrorhizata. In this paper, we argue to assign the early foraminifers to the Fusulinata, and to subdivide this class into six orders: Parathuramminida, Archaediscida and Earlandiida (forming together the subclass Afusulinana n. subcl.), and Tournayellida, Endothyrida and Fusulinida (subclass Fusulinana nom. translat.). These subdivisions are discussed and linked to the first occurrences of the later classes: Miliolata, Nodosariata and Textulariata. The environmental living conditions of the first fossilized foraminifers remain enigmatic during the Early Palaeozoic (Cambrian-Silurian). During the Late Silurian, the unilocular Parathuramminida started to colonize the inner parts of ramps and platforms. The first plurilocular microgranular foraminifers (Semitextulariidae, Nanicellidae, and Eonodosariidae) developed in back-reefal systems and in deeper-water environments (“griottes”-type nodular limestone) from the late Early Devonian to the early Late Devonian. The Moravamminida, another group of possible Protista, are typical markers of Devonian inner ramp systems. The Semitextulariidae, Nanicellidae, and Eonodosariidae did not survive the Frasnian/Famennian crisis. From the Tournaisian to the Serpukhovian (Mississippian subsystem or Early Carboniferous), numerous new genera and species of Archaediscida, Tournayellida and Endothyrida flourished but remained confined to inner ramp environments. In deeper water depositional systems (i.e. coral thrombolite microbialites and/or nodular limestones), a few opportunistic Foraminifera were living up to the disphotic zone. During the Pennsylvanian (Bashkirian to Gzhelian), the habitats extended to more confined, shallower areas of the inner ramp (with Staffelloidea). During the Late Carboniferous and Permian, the larger Fusulinida (Schwagerinoidea) reached the outer platform as they have been commonly reworked in calciturbidites. During the Late Permian, some taxa were even able to live in hypersaline environments such as sabkhas and hypersaline lagoons. Two major biotic crises occurred during the Permian (post-Middle and post-Late Permian crisis), but the number of survivors after the PTE (Permian/Triassic Extinction) is probably higher than previously admitted. From the Cambrian to the Serpukhovian, the Foraminifera were probably all infaunal or living at the sediment/seawater interface. The TROX and TROX-2 models are consequently applicable. Anoxia, often suggested as triggering environmental crises, was likely not systematically lethal for many infaunal foraminifers. The late Tournaisian-Changhsingian Tetrataxis genus was probably the first epiphyte foraminifer, because of its conical, limpet-like test. The Tetrataxidae (e.g., Tetrataxis, Pseudotaxis and Abadehella) constituted the unique trochospirally coiled plurilocular foraminiferal family of the Palaeozoic. The Bradyinoidea, Ozawainelloidea, Staffelloidea, and the Pseudoschwagerinidae (Schwagerinoidea) are other examples of Pennsylvanian-Permian epiphytes but cannot be considered as planktonic taxa. All the other Schwagerinoidea are related to high-energy environments and coarse-grained substrates. Their history, as well as that of the Neoschwagerinoidea, was likely subject to the vicissitudes of their endosymbiotic algae.