Systematics and palaeoecology of Changhsingian (Late Permian) Ambocoeliidae brachiopods from South China and implications for the end-Permian mass extinction

Four Ambocoeliidae brachiopod species including one new species (Crurithyris tazawai sp. nov., Crurithyris sp., Paracrurithyris pygmaea and Attenuatella mengi) are described from the Changhsingian (Late Permian) deep-water facies of South China. Analysis of the morphology, palaeoecology and palaeogeographical and temporal distributions of these species revealed that the presence of a delthyrium and/or the micro-ornaments among three of the four species (Crurithyris tazawai sp. nov., Paracrurithyris pygmaea and Attenuatella mengi) favoured an epifaunal (epiphytic) lifestyle. Morphological differences suggest that Paracrurithyris pygmaea may have been more effective metabolically in forming the shell compared with Attenuatella mengi and Crurithyris tazawai. The temporal and palaeogeographical distribution of Attenuatella suggests that A. mengi inhabited cool or cold deep waters. Both Crurithyris tazawai and Attenuatella mengi disappeared earlier in the stratigraphic record than Paracrurithyis pygmaea during the PermianTriassic mass extinction. These differences in timing of extinction, morphology and palaeogeographical distributions suggest that oxygen deficiency and trophic resource limitation (a consequence of the changing composition of marine phytoplankton in the seas) may have contributed to the end-Permian mass extinction.