In this volume John Murray investigates the ecological processes that control the distribution, abundance, and species diversity of benthic foraminifera in environments ranging from marsh to the deepest ocean. To interpret the fossil record it is necessary to have an understanding of the ecology of modern foraminifera and the processes operating after death leading to burial and fossilisation. This book presents the ecological background required to explain how fossil forms are used in dating rocks and reconstructing past environmental features including changes of sea level. It demonstrates how living foraminifera can be used to monitor modern-day environmental change. Ecology and Applications of Benthic Foraminifera presents a comprehensive and global coverage of the subject using all the available literature. It is supported by a website hosting a large database of additional ecological information (www.cambridge.org/0521828392) and will form an important reference for academic researchers and graduate students in Earth and Environmental Sciences.

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Ecology and Applications of Benthic Foraminifera

Mid-water oxygen minima (� 0.5 ml l � 1 dissolved O2) intercept the continental margins along much of the eastern Pacific Ocean, off west Africa and in the Arabian Sea and Bay of Bengal, creating extensive stretches of sea floor exposed to permanent, severe oxygen depletion. These seafloor oxygen minimum zones (OMZs) typically occur at bathyal depths between 200 m and 1000 m, and are major sites of carbon burial along the continental margins. Despite extreme oxygen depletion, protozoan and metazoan assemblages thrive in these envi- ronments. Metazoan adaptations include small, thin bodies, enhanced respiratory surface area, blood pigments such as haemoglobin, biogenic structure formation for stability in soupy sedi- ments, an increased number of pyruvate oxidoreductases, and the presence of sulphide-oxidising symbionts. The organic-rich sediments of these regions often support mats of large sulphide- oxidising bacteria (Thioploca, Beggiatoa, Thiomargarita ), and high-density, low-diversity meta- zoan assemblages. Densities of protistan and metazoan meiofauna are typically elevated in OMZs, probably due to high tolerance of hypoxia, an abundant food supply, and release from predation. Macrofauna and megafauna often exhibit dense aggregations at OMZ edges, but depressed densities and low diversity in the OMZ core, where oxygen concentration is lowest. Taxa most tolerant of severe oxygen depletion ( � 0.2 ml l � 1 ) in seafloor OMZs include calcare- ous foraminiferans, nematodes, and annelids. Agglutinated protozoans, harpacticoid copepods, and calcified invertebrates are typically less tolerant. High dominance and relatively low species richness are exhibited by foraminiferans, metazoan meiofauna, and macrofauna within OMZs. At dissolved oxygen concentrations below 0.15 ml l � 1 , bioturbation is reduced, the mixed layer is shallow, and chemosynthesis-based nutrition (via heterotrophy and symbiosis) becomes important. OMZs represent a major oceanographic boundary for many species. As they expand and con- tract over geological time, OMZs may influence genetic diversity and play a key role in the evo- lution of species at bathyal depths. These ecosystems may preview the types of adaptations, species, and processes that will prevail with increasing hypoxia over ecological and evolution- ary time. However, many questions remain unanswered concerning controls on faunal standing stocks in OMZs, and the physiological, enzymatic, metabolic, reproductive and molecular adap- tations that permit benthic animals to live in OMZs. As global warming and eutrophication reduce oxygenation of the world ocean, there is a pressing need to understand the functional consequences of oxygen depletion in marine ecosystems.

http://levin.ucsd.edu/publications/2003,%20OMBAR,%20Levin.pdf

Oxygen minimum zone Benthos: Adaptation and community response to hypoxia

Publisher Summary This chapter focuses on the paleoceanographical proxies based on deep-sea benthic foraminiferal assemblage characteristics, and presents the following three proxy relationships that are promising: those between benthic foraminiferal faunas and benthic ecosystem oxygenation, export productivity, and deep-sea water mass characteristics. Under most circumstances the composition of deep-sea benthic foraminiferal assemblages is controlled by a rather limited number of environmental factors. The available proxies based on benthic foraminiferal assemblage composition show that they have major potential, but further research is needed to add or improve the quantitative aspects. In many cases, such as bottom water oxygenation, and Corg flux to the ocean floor, it can be done by significantly increasing the size of existing databases. In other cases such as periodicity of the organic flux, time series observations are necessary. A major obstacle is insufficient knowledge of the differences between recent and fossil faunas due to taphonomical alterations. This phenomenon, of importance for all paleoceanographic proxies, can to some extent be solved relatively easily in the case of foraminiferal assemblages by detailed studies of their vertical succession in sediments deposited in the last 5,000 years, when environmental conditions were probably rather invariable in many areas. Proxies based on foraminiferal assemblage composition are fundamentally different from all geochemical proxies, and thus may provide independent reconstructions of essential oceanographic parameters.

Chapter Seven Paleoceanographical Proxies Based on Deep-Sea Benthic Foraminiferal Assemblage 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.

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

Benthic foraminifers inhabit a wide range of aquatic environments including open marine, brackish, and freshwater environments. Here we show that several different and diverse foraminiferal groups (miliolids, rotaliids, textulariids) and Gromia, another taxon also belonging to Rhizaria, accumulate and respire nitrates through denitrification. The widespread occurrence among distantly related organisms suggests an ancient origin of the trait. The diverse metabolic capacity of these organisms, which enables them to respire with oxygen and nitrate and to sustain respiratory activity even when electron acceptors are absent from the environment, may be one of the reasons for their successful colonization of diverse marine sediment environments. The contribution of eukaryotes to the removal of fixed nitrogen by respiration may equal the importance of bacterial denitrification in ocean sediments.

Widespread occurrence of nitrate storage and denitrification among Foraminifera and Gromiida.

In the meso-oligotrophic Bay of Biscay, a diminishing downward organic matter flux with depth is accompanied by an important decrease of the live foraminiferal density. Although bottom water oxygenation is not directly influenced by organic matter input, the oxygenation of interstitial waters and the primary redox fronts do change in response to variations of the organic matter flux. The occurrence of deep and intermediate infaunal taxa can be linked to fundamental redox fronts and putative associated bacterial consortia. Our data are in agreement with the TROX-model, which explains the benthic foraminiferal microhabitat as a function of organic flux and benthic ecosystem oxygenation. Both the depth of the principle redox fronts and the microhabitat of deep infaunal species show important increases with depth. At the deepest oligotrophic stations, deep infaunal faunas become relatively poor. Therefore, the exported flux of organic matter appears to be the main parameter controlling the composition and the vertical distribution of benthic foraminiferal faunas below the sediment-water interface. The oxygenation of pore waters plays only a minor role. A species-level adaptation of the TROX-model is presented for the Bay of Biscay.

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A Alexandre

Papers

Live benthic foraminiferal faunas from the Bay of Biscay: faunal density, composition, and microhabitats