Interrelations between framework-building and encrusting skeletal organisms and microbes: more-refined growth history of Lower Devonian bindstones



The framework-building stromatoporoid Stachyodes, the encrusting calcimicrobe Rothpletzella and encrusting Graticula-like red algae are major contributors to red algal–calcimicrobial–stromatoporoid bindstones in the Lower Devonian Elmside Formation of the Yass Basin, New South Wales, Australia. The distribution and accumulation patterns of encrusting organisms within the red algal–calcimicrobial–stromatoporoid bindstones observed by optical microscopy and SEM imply a biotic interrelationship between skeletal organisms and microbes that reflects environmental changes. Rothpletzella is characterized by prostrate filaments with frequent branching and a high angle of bifurcation. Filaments of Graticula-like red algae exhibit rare branching and a relatively low angle of bifurcation. In addition, they are prostrate at the base before becoming erect. Both Rothpletzella and the red algae successively encrust the surfaces of skeletal frameworks, but exhibit different distributions. Rothpletzella and other calcimicrobes cover both the lower and upper surfaces of frameworks, whereas red algae are limited to the upper surfaces. Their individual distributions are thus significantly influenced by the frameworks formed by the thin, laminar stromatoporoid Stachyodes, which create different microenvironments as by-products. The limited distribution of the red algae was probably related to light levels or phototropism. Upper framework surfaces are variously encrusted by calcimicrobes and the red algae to form thick crusts with varying accumulation patterns. Micritization around the algal thalli, covering of calcimicrobes such as Wetheredella, and microbial micrites between algal thalli all suggest interruptions of algal growth that correspond to episodes of harsh environmental conditions. Transitions from Graticula-like red algae to Rothpletzella reflect periods of deteriorating environmental change for skeletal organisms, which resulted in the predominance of microbial growth. In contrast, a resurgence of red algae on calcimicrobes suggests improved environmental change. Repeated accumulation patterns between Graticula-like red algae and Rothpletzella indicate changing habitat environments and competitive relations within skeletal organisms and microbes. These relationships provide insight into understanding how skeletal organisms and microbes utilized space and how they interrelated with each other to produce Devonian reefal limestones.