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PUBLISHED: 26 JULY 2016 | ARTICLE NUMBER: 16114 | DOI: 10.1038/NMICROBIOL.2016.114 correspondence Haeckel’s 1866 tree of and the origin of

To the Editor — In their Letter describing were summarized under the an expanded version of the tree of life, Eukarya, which includes all eukaryotic Hug et al.1 refer to a key publication of micro- and macroorganisms (from amoebae and co-workers2, wherein a to humans). 16S rRNA-phylogenetic scheme of the In chapter 26 entitled ‘Phylogenetic domains , and Eukarya is theses’, Haeckel3 re-interpreted and shown. However, the first three- supplemented Darwin’s evolutionary tree of life that included microorganisms deductions of 1859. With reference to the was depicted by (pictured simplest (bacteria), which form the at around 26 years old) in his General root of his ‘oak tree’ (see Fig. 6 in ref. 5), Morphology of Organisms, a seminal book Haeckel3 concluded that “all organisms that was published one hundred and are the descendants of such autogenous fifty years ago3. Moneren”. Hence, according to this In his Origin of Species, Haeckelian interpretation of the phylogeny refers to the Linnean two-kingdom of life, the Monera (that is, Bacteria and system, with ‘Animalia’ and ‘Vegetabilia’ Archaea)2 are the progenitors of all more as the only two branches of the living complex living beings on Earth. world. Haeckel3, who was also known This 150-year-old idea is consistent with as the German Darwin4, introduced a recent discoveries and genomic analyses that three-kingdom scheme in which, as a support an archaeal origin of eukaryotes1,8. supplement to the Linnean Animalia and A ‘Lokiarchaea’-like host that had Plantae (synonymous with Vegetabilia), rudimentary phagocytotic capabilities for an the ‘Protista’ were added5. Haeckel’s / GETTY PORTFOLIO / MONDADORI / CONTRIBUTOR PORTFOLIO MONDADORI uptake of bacteria may have been the most protists included all microscopic organisms ancient, metabolically active, self-replicating known at that time (Amoebozoa, Bacterium, and provided a definition of protoplasmic unit from which all life on Myxomycetes etc.), and, as one of its eight these tiny microorganisms that included Earth evolved9. ❐ divisions, the ‘Monera’. their characteristic mode of propagation via In several chapters of his General binary fission6. References Morphology, Haeckel3 characterized the tiny Despite Cohn’s introduction of the term 1. Hug, L. A. et al. Nature Microbiol. 1, 16048 (2016). 2. Woese, C. R., Kandler, O. & Wheelis, M. L. Moneren as the “most simple organisms, Bacteria in 1872, Haeckel’s term Monera Proc. Natl Acad. Sci. USA 87, 4576–4579 (1990). without structure, homogeneous pieces survived in the biomedical literature 3. Haeckel, E. Generelle Morphologie der Organismen Vols I and II of Plasma”, and explicitly mentioned the until 1969, when Robert H. Whittaker (Georg Reimer, 1866). 4. Kutschera, U. Evo. Edu. Outreach 2, 564–566 (2009). bacterial genus as an example. Hence, introduced his five-kingdom-system of 5. Kutschera, U. Biol. Direct 6, 33 (2011). Haeckel’s Monera included microorganisms classification. In this scheme, Monera refers 6. Cohn, F. Beitr. Biol. Pflanzen 1(II), 127–224 (1872). that are today known as Bacteria (a term not to prokaryotic microorganisms (bacteria, 7. Whittaker, R. H. Science 163, 150–160 (1969). 8. Sprang, A. et al. Nature 521, 173–179 (2015). yet coined in 1866). ), that are juxtaposed to four 9. Martin, W. F., Garg, S. & Zimorski, V. Phil. Trans. R. Soc. B Six years later, the German botanist eukaryotic kingdoms (Protista, Fungi, 370, 20140330 (2015). Ferdinand Cohn published a monograph Animalia, Plantae)7. This taxonomic on the occurrence, cultivation, morphology, interpretation of extant biodiversity was behaviour, reproduction, nutrition, and replaced by the three-domains-system of U. Kutschera systematics of a group of microorganisms Woese et al.2, in which Monera are equated Institute of Biology, University of Kassel, he called Bacteria. Cohn used the word with the prokaryotic domains Bacteria and D-34132 Kassel, Germany. ‘bacteria’ with reference to the genus-name Archaea. The four remaining kingdoms e-mail: [email protected]

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