Why Did Some Larger Benthic Foraminifera Become so Large and Flat?

Biostromes and low-relief bioherms, some of which are characterized by exceptionally large, flat specimens of larger benthic foraminifera, are common in Palaeogene and Miocene carbonates, most notably those deposited along the Neotethys Seaway and tropical Pacific islands. By incorporating insights from palaeoceanographic research and the biology of living larger benthic foraminifera, a scenario is proposed that can account for palaeontological and sedimentological features while augmenting previous interpretations. Sexual reproduction by gamete broadcasting is common in foraminiferal taxa, including extant Nummulitidae and Amphisteginidae. Resultant zygotes can develop into tiny, resistant, easily dispersed propagules that recruit in suitable benthic-environmental conditions. The role of algal symbiosis in the biology of larger benthic foraminifera is well-documented. Palaeoceanographically, such taxa proliferated during times of reduced thermal stratification of the oceans. In regions with exceptionally clear, nutrient-depleted waters, ‘twilight-zone’ light penetration was sufficient, at least intermittently, to support some photosynthesis. On outer-shelf or promontory sites at depths of ca 100 to 200 m, the tiny propagules of larger benthic foraminiferal species, incorporating algal symbionts with the lowest light requirements, could have settled and recruited, growing very slowly, nourished by feeding on bacteria and the limited photosynthate produced by their algal symbionts. Under such conditions, thin microspheric individuals of one or two larger benthic foraminiferal taxa could have survived and grown slowly over several years to very large shell diameters, seldom reproducing asexually. Resulting carbonate accumulation rates would have been very slow, such that even rare disturbances by currents, major storms or internal waves could have produced evidence of winnowing and sedimentary structures. The fossil evidence of such habitats should include biostromes or possibly low-relief bioherms of low diversity assemblages characterized by abundant, exceptionally large, flat, microspheric larger benthic foraminifera