Morphological optimization in the largest living foraminifera: implications from finite element analysis

Abstract

Benthic foraminifera have attained gigantic sizes many times throughout geologic history. To understand the selective processes underlying foraminiferal gigantism, we used a computer-based, three-dimension, solid finite element model to analyze the mechanical strength of different discoid forms, including two of the largest living foraminifera—Cycloclypeus carpenteri Brady and Marginopora vertebralis Quoy and Gaimard. These two species enlarge by cyclic, planar growth, resulting in slightly biconvex (C. carpenteri) and slightly biconcave (M. vertebralis) forms. As the tests enlarge, the maximum stresses induced by a standard bending moment decrease in both species. Such stress-reducing growth plans apparently allow growth to extraordinarily large sizes and allow volume to increase with minimal lowering of the surface-to-volume ratio, a critical functional factor in ensuring increased surface area for photosynthesis by endosymbionts contained within the tests and for chemical exchange by the foraminifera with the external environment. Of the two species, M. vertebralis has a stronger construction and a lower surface-to-volume ratio. These features indicate optimal constructional solutions to environmental constraints (degree of turbulence and light availability) in their disparate natural habitats: M. vertebralis in the mechanically rigorous inter- to subtidal, and C. carpenteri on the light-minimal and hydraulically quieter, deeper sea beds. We conclude that morphological design in larger foraminifera is constrained by a biomechanical factor, and that gigantism and biomechanical optimization are demonstrably related.