A new model for the calcification of the green macro-alga Halimeda opuntia (Lamouroux)

Halimeda opuntia is a cosmopolitan marine calcifying green alga in shallow tropical marine environments. Besides Halimeda's contribution to a diverse habitat, the alga is an important sediment producer. Fallen calcareous segments of Halimeda spp. are a major component of carbonate sediments in many tropical settings and play an important role in reef framework development and carbonate platform buildup. Consequently the calcification of H. opuntia accounts for large portions of the carbonate budget in tropical shallow marine ecosystems. Earlier studies investigating the calcification processes of Halimeda spp. have tended to focus on the microstructure or the physiology of the alga, thus overlooking the interaction of physiological and abiotic processes behind the formation of the skeleton. By analyzing microstructural skeletal features of Halimeda segments with the aid of scanning electron microscopy and relating their occurrence to known physiological processes, we have been able to identify the initiation of calcification within an organic matrix and demonstrate that biologically induced cementation is an important process in calcification. For the first time, we propose a model for the calcification of Halimeda spp. that considers both the alga's physiology and the carbon chemistry of the seawater with respect to the development of different skeletal features. The presence of an organic matrix and earlier detected external carbonic anhydrase activity suggest that Halimeda spp. exhibit biotic precipitation of calcium carbonate, as many other species of marine organisms do. On the other hand, it is the formation of micro-anhedral carbonate through the alga's metabolism that leads to a cementation of living segments. Precisely, this process allows H. opuntia to contribute substantial amounts of carbonate sediments to tropical shallow seas.