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Phylogenetic, taxonomic and biogeographical studies in the Pithophoraceae (Cladophorales, Chlorophyta) Boedeker, C.

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Phylogenetic, taxonomic and biogeographical studies in the Pithophoraceae (Cladophorales, Chlorophyta)

Boedeker, C.

Citation

Boedeker, C. (2010, December 21). Phylogenetic, taxonomic and biogeographical studies in the Pithophoraceae (Cladophorales, Chlorophyta). Retrieved from https://hdl.handle.net/1887/16263

Version: Corrected Publisher’s Version

License: Licence agreement concerning inclusion of doctoral thesis in the Institutional Repository of the University of Leiden Downloaded from: https://hdl.handle.net/1887/16263

Note: To cite this publication please use the final published version (if applicable).

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