Theoretical morphology of colonial meshworks in the Fenestrata (Bryozoa): modelling hydraulic-resistance constraints and minimum fenestrule widths

Abstract

Ancient fenestrate bryozoans (Fenestellidae, Polyporidae) exhibit a range of fenestrule widths that can be investigated to gain insights about constraints on meshwork morphology. Theoretical morphospace analyses of branch width and spacing reveal that there is a minimum fenestrule width common to both groups. We propose that the minimum fenestrule width observed in fenestrate colonies was constrained by hydraulic resistance (the hydraulic-resistance-constraint hypothesis). Modelling that considers both the viscous and inertial components of fluid-flow resistance through hypothetical fenestrate meshworks corroborates this hypothesis for the smaller fenestrate meshworks of the biserial fenestellids and for polyporids with branch widths less than about 0.6 mm. However, the geometry of the polyporid meshworks in species with branches wider than 0.6 mm does not appear to have been determined by a hydraulic-resistance constraint. Instead, maximum branch densities in the larger-branched polyserial polyporid colonies appear to have been constrained by a limit on the number of zooid rows that these colonies could develop on their branches (the zooid-row-limit hypothesis). The observed decrease in the minimum fenestrule width in these colonies is the indirect geometric result of the inability of the larger polyserial polyporid colonies to develop meshworks with high branch densities.