Davidsonia and Rugodavidsonia (new genus), cryptic Devonian atrypid brachiopods from Europe and South China

The fixosessile, perireefal to reefal Devonian epibiont Davidsonia cemented itself by its entire ventral valve to the underside of alveolitid tabulate corals, or to stromatoporoid sponges. It occupied a cryptic niche in perireefal biostromal thickets to reef complexes in the Old World faunal realm (Europe, western North America), and Uralian–Central Asian faunal province. A new species, D. septata, is here reported in abundance for the first time from Middle Devonian (Givetian) rocks of Yunnan, thus extending its range to South China. The usual absence of the brachial valve is suggested to be due to postmortem taphonomy as a result of its unique, cemented mode of life on the undersides of corals or sponges. A new genus of coarsely ribbed, partly libero-, partly fixosessile davidsoniid, Rugodavidsonia, type Davidsonia woodwardiana (DeKoninck), mimicking unrelated Late Paleozoic orthotetids, is also described, demonstrating that ribs evolved secondarily in a family dominated from its inception by smooth shells. Its paleogeographic distribution ranged from Europe through western North America (and possibly China), commonly accompanying Davidsonia in similar assemblages, but with a slightly different mode of life. Reexamination of available material, using thin sections, peels and SEM, confirms that the impunctate shell structure, dorsally directed spiralia, nature of the hinge mechanism, the deltidial plates, absence of a chilidium, “atrypoid” musculature and vascular markings, show features in Davidsonia and Rugodavidsonia that are all typical of the Order Atrypida. Thus, Davidsonia demonstrates that ventral valve cementation evolved independently more than once in articulate brachiopods such as strophomenids, productids, atrypids, and thecideid spiriferids. A revised classification of the Davidsonioidea and Palaferelloidea is proposed. Relationships between various genera are outlined in a new family tree for these groups. The various families discussed, including the Symmatrypidae, are united under a new suborder, the Davidsoniidina, a primarily carbonate platform and perireefal dwelling group, which first evolved during the Ludlovian (Late Silurian) in the Uralian–Central Asian tropical shelf region, diversified over a period of ~47 Ma, and then died out during the late Givetian, 10 Ma prior to the Frasnian–Famenian mass extinction events.