The effects of mid-Phanerozoic environmental stress on bryozoan diversity, paleoecology, and paleogeography

Evidence of sustained environmental degradation associated with the end-Guadalupian, end-Permian, and end-Triassic extinctions has been inferred from numerous geochemical and sedimentological studies, but the long-term impacts of this extinction-associated stress on the evolutionary trajectories of marine invertebrates have not been explored. An examination of the diversity, extinction, paleoenvironmental range, and geographical distribution of marine stenolaemate bryozoans during the Permian to Jurassic interval provides striking new evidence of the taxonomic and ecological influence of these mid-Phanerozoic extinctions on one of the most abundant components of the Paleozoic Fauna. Elevated bryozoan extinction rates during the Late Permian and Late Triassic were coupled with major changes in their habitats. Bryozoans gradually disappeared from deep-water offshore settings during the Late Permian and from nearshore and offshore settings during the Late Triassic. Re-colonization of these environments in the wake of each crisis was delayed but coupled with increases in global generic diversity. The taxonomic effects of the end-Guadalupian extinction were milder than previously described, even though ecologically bryozoans were becoming restricted to nearshore settings. The end-Permian mass extinction remained the largest for bryozoans, drastically reducing global and assemblage generic diversity and triggering a permanent change in their paleoenvironmental preferences from nearshore to mid-shelf settings. The 285 Myr dominance of stenolaemate bryozoans ended during the Late Triassic when all but one order (Cyclostomata) became extinct, initiating a taxonomic switch between stenolaemate and gymnolaemate bryozoans. Moreover, spatio-temporal variations in the paleoenvironmental history of bryozoans imply that Late Permian and Late Triassic marine environmental instability resulted largely from some stressful deep-water phenomenon. High extinction rates in nearshore environments in the Late Permian provide a link between marine and terrestrial/atmosphere extinction-related perturbations.