Initial diversification of macroboring ichnofossils and exploitation of the macroboring niche in the lower Paleozoic

The traces of macroboring organisms are known throughout the Phanerozoic, with diversification and exploitation of the macroboring niche paralleling variations in the development of skeletal metazoa. The oldest macroboring biota is an abundant yet low diversity fauna in hardgrounds and reefs of Lower Cambrian age. Following the extinction of archaeocyathids at the end of the Lower Cambrian (and thus the demise of skeletal reefs until the Middle Ordovician), boring organisms appear to be restricted to submarine hardgrounds. With the development of skeletal reefs in the Middle Ordovician the macroboring fauna shows a rapid speciation and a dramatic increase in diversity. This same pattern occurs again in the Devonian. This record appears to represent refuge of the fauna in low stress, hardground environments when skeletal reefs were not present and radiation in the high stress environment of the reef when large skeletal metazoa were abundant and diverse.