Revision of Lower Devonian clam shrimp (Branchiopoda, Diplostraca) from the Rhenish Massif (Eifel, SW-Germany), and the early colonization of non-marine palaeoenvironments

Three collections of clam shrimp from the Lower Devonian (upper lower Emsian) Klerf Formation of Willwerath and Waxweiler in the western Eifel Mountains are herein studied. Four discernible morphotypes are present in the Willwerath assemblage. These morphotypes correspond to the previously described species from this locality: Pseudestheria diensti (Gross), Pseudestheria subcircularis Raymond, and Palaeolimnadiopsis? eifelensis Raymond. The fourth morphotype also closely corresponds to Pseudestheria diensti, but is slightly different in outline. Asmussia willweratica (Novozhilov) is an objective synonym of Pseudestheria subcircularis. In the Willwerath clam shrimp sample, it is difficult to disentangle taphonomic/preservational versus ontogenetic/sexual variation. If taphonomic/preservational causes could be confirmed, all the Willwerath specimens may represent just one taxon and Pseudestheria subcircularis and Palaeolimnadiopsis? eifelensis would be potential subjective synonyms of Pseudestheria diensti. However, we refrain from a formal taxonomic act, which requires a larger number of specimens to be analysed.

In addition, we investigated two samples from Waxweiler, which originate from distinct centimetre-thick layers and therefore represent associations. One of these associations comprises two discernible clam shrimp morphotypes. Because of limited preservation of morphological characters, one is here left in open nomenclature as Spinicaudata incertae sedis, whereas the other is recognised as a new species of the family Palaeolimnadiopseidae, Palaeolimnadiopsis frankeorum sp. nov. The second association from Waxweiler comprises at least four, possibly six different clam shrimp taxa. In addition to Pseudestheria diensti, Palaeolimnadiopsis sp., and poorly preserved Asmussia- and Ulugkemia-like specimens, two taxa can be described as new, namely the vertexiid Cornia wasvilrensis sp. nov. and the leaiine Eicheleaia wenndorfi gen. nov. et sp. nov. Altogether, the Klerf Formation at Waxweiler yields a total of at least five (possibly eight) clam shrimp taxa. The palaeoenvironment of these early clam shrimp from the Klerf Formation is interpreted as a non-marine, relatively proximal deltaic setting with no or very minor marine influence, such as a deltaic freshwater pond or lake. The comparatively high diversity of these lower Devonian clam shrimp suggests a considerably earlier origin and cryptic evolution of the group.