From Haeckel to Hennig: the early development of phylogenetics in German-speaking Europe

An outline of the development of phylogenetic thinking and methodology in German literature published between 1862 and 1942 is presented. Central European biologists and palaeontologists of the first post-Darwinian generation of biologists holding evolutionary views were directly stimulated by Darwin. Members of the second generation, mostly born after 1850, were largely influenced also by colleagues of the first post-Darwinian generation, mainly by Haeckel. Among them were O. Abel, V. Franz, R. Hertwig, A. Naef, L. Plate, and R. v. Wettstein. Opinions on the relationship between systematics and phylogeny differed considerably. Many authors admitted that phylogeny must be mirrored in systematics but at the same time shared Haeckel's views on classification, which permitted paraphyletic groupings. Particularly Abel and Naef took systematics several steps further, and many important elements of phylogenetic systematics were developed several decades before Hennig. Naef presented a definition of a phylogenetic group that exactly matches Hennig's definition of monophyly. He also formulated a species concept that was implicitly based on reproductive isolation. This was an important presupposition for viewing speciation as the splitting of a stem species into daughter species. However, many authors of the first half of the 20th century repeated old, but established views on phylogenetics, while others overlooked or misunderstood earlier progressive views thus causing slow development of phylogenetic systematics in Central Europe. Its development almost stopped between 1925 and 1950, because of a widespread shift towards typology and extreme idealistic morphology. During that time very few persons such as W. Zimmermann and W. Hennig assembled elements of phylogenetic systematics and combined them with their own thoughts to create a sound theory and methodology.