Phosphatic sediments of the Late Neoproterozoic (ca. 600 million years old [Myr]) Doushantuo Formation at Weng'an, South China, contain fossils of multicellular algae preserved in anatomical detail. As revealed by light microscopy and scanning electron microscopy, these fossils include both simple pseudoparenchymatous thalli with apical growth but no cortex-medulla differentiation and more complex thalli characterized by cortex-medulla differentiation and structures interpretable as carposporophytes, suggesting a multiphasic life cycle. Simple pseudoparenchymatous thalli, represented by Wengania, Gremiphyca, and Thallophycoides, are interpreted as stem group florideophytes. In contrast, complex pseudoparenchymatous thalli, such as Thallophyca and Paramecia, compare more closely to fossil and living corallinaleans than to other florideophyte orders, although they also differ in some important aspects (e.g., lack of biocalcification). These more complex thalli are interpreted as early stem group corallinaleans that diverged before Paleozoic stem groups such as Arenigiphyllum, Petrophyton, Graticula, and Archaeolithophyllum. This phylogenetic interpretation implies that (1) the phylogenetic divergence between the Florideophyceae and its sister group, the Bangiales, must have taken place before Doushantuo time-an inference supported by the occurrence of bangialean fossils in Mesoproterozoic rocks; (2) the initial diversification of the florideophytes occurred no later than the Doushantuo time; and (3) the corallinalean clade had a "soft" (uncalcified) evolutionary history in the Neoproterozoic before evolving biocalcification in the Paleozoic and undergoing crown group diversification in the Mesozoic.

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The fine structure of pit plugs was examined in 63 species of red algae representing 34 families. The number of plug cap layers was found to be a taxonomically reliable character when tested against a recent revision of the Rhodymeniales which established the order Palmariales. In a broader survey involving all the orders of red algae possessing pit plugs, three basic plug cap configurations were detected. These differ in having one, two, or no plug cap layers. Some taxa have plug caps in which the outer cap layer is greatly elaborated, however, this layer was shown to be positionally and cytochemically identical, thus probably homologous, to the outer cap layer of the typical two-layered plug cap. The pit plug character confirms the validity of the order Palmariales and indicates that the Nemaliales and Cryptonemiales are heterogeneous assemblages. Revision of these groups along lines indicated by the plug character is supported by significant features of morphology and reproduction. These correlations lead us to propose that Rhodophysema and Coriophyllum be transferred to the Palmariales, that the Gelidiales be resurrected, that the Bonnemaisoniales (including the Naccariaceae) is deserving of recognition, that the Batrachospermaceae be raised to a higher rank which will also receive the Lemaneaceae and the Thoreaceae, and that the Corallinaceae and Hildenbrandiaceae both be raised to ordinal status. The authors would like to thank

Representatives of the freshwater red algal familyThoreaceae were studied to resolve their taxonomic and phylogenetic status. Three specimens of Nemalionopsis and five collections of Thorea were examined for pit plug ultrastructure and analyzed for the sequences of the genes coding for the large subunit of RUBISCO ( rbc L) and the small subunit of rRNA (18S rRNA). The phylogenetic trees generated from the two genes, and a combined tree all showed the Thoreaceae to be contained in a well-supported monophyletic clade that is separate from the other two families currently classified in the Batrachospermales, the Batrachospermaceae and the Lemaneaceae. In addition, secondary structure elements of the 18S rRNA gene were observed at positions 650 and 1145 ( Escherichia coli numbering system) that are not present in other members of the Rhodophyta. The pit plugs of the gametophytic and chantransia stages of the Thoreaceae contain two cap layers, the outer one of which is typically plate-like, though occasionally inflated ones have been seen. No pit plug cap membrane has been observed. These findings indicate the Thoreaceae has been misclassified in the Batrachospermales and should be placed in its own order, the Thoreales. This order is characterized by having freshwater representatives with multiaxial gametophytes, a uniaxial chantransia stage, and pit plugs with two cap layers, the outer one of which is usually plate-like.