Benthic foraminifera (Protista) as tools in deep-water palaeoceanography: environmental influences on faunal characteristics.

Foraminiferal research lies at the border between geology and biology. Benthic foraminifera are a major component of marine communities, highly sensitive to environmental influences, and the most abundant benthic organisms preserved in the deep-sea fossil record. These characteristics make them important tools for reconstructing ancient oceans. Much of the recent work concerns the search for palaeoceanographic proxies, particularly for the key parameters of surface primary productivity and bottom-water oxygenation. At small spatial scales, organic flux and pore-water oxygen profiles are believed to control the depths at which species live within the sediment (their 'microhabitats'). Epifaunal/shallow infaunal species require oxygen and labile food and prefer relatively oligotrophic settings. Some deep infaunal species can tolerate anoxia and are closely linked to redox fronts within the sediment; they consume more refractory organic matter, and flourish in relatively eutrophic environments. Food and oxygen availability are also key factors at large (i.e. regional) spatial scales. Organic flux to the sea floor, and its seasonality, strongly influences faunal densities, species compositions and diversity parameters. Species tend to be associated with higher or lower flux rates and the annual flux range of 2-3 g Corg m-2 appears to mark an important faunal boundary. The oxygen requirements of benthic foraminifera are not well understood. It has been proposed that species distributions reflect oxygen concentrations up to fairly high values (3 ml l-1 or more). Other evidence suggests that oxygen only begins to affect community parameters at concentrations < 0.5 ml l-1. Different species clearly have different thresholds, however, creating species successions along oxygen gradients. Other factors such as sediment type, hydrostatic pressure and attributes of bottom-water masses (particularly carbonate undersaturation and current flow) influence foraminiferal distributions, particularly on continental margins where strong seafloor environmental gradients exist. Epifaunal species living on elevated substrata are directly exposed to bottom-water masses and flourish where suspended food particles are advected by strong currents. Biological interactions, e.g. predation and competition, must also play a role, although this is poorly understood and difficult to quantify. Despite often clear qualitative links between environmental and faunal parameters, the development of quantitative foraminiferal proxies remains problematic. Many of these difficulties arise because species can tolerate a wide range of non-optimal conditions and do not exhibit simple relationships with particular parameters. Some progress has been made, however, in formulating proxies for organic fluxes and bottom-water oxygenation. Flux proxies are based on the Benthic Foraminiferal Accumulation Rate and multivariate analyses of species data. Oxygen proxies utilise the relative proportions of epifaunal (oxyphilic) and deep infaunal (low-oxygen tolerant) species. Yet many problems remain, particularly those concerning the calibration of proxies, the closely interwoven effects of oxygen and food availability, and the relationship between living assemblages and those preserved in the permanent sediment record.