"Organismic" positions in early German-speaking ecology and its (almost) forgotten dissidents

In early German ecology, the key concept used to refer to a synecological unit was Biozönose (biocoenosis). Taken together with the concept of the Biotop (biotope), it was also understood as an integrated higher-order unit of life, sometimes called a "Holozön" (holocoen). These units were often perceived as having properties similar to those of individual organisms, and they informed the mainstream of German ecology until at least the late 1960s. Here I ask how "organismic" these concepts really were and what conceptual problems they entailed. To do so, I focus on some almost forgotten dissident positions, especially those of (German-born) Friedrich Simon Bodenheimer and Fritz Peus, which I contrast with the mainstream German ecology of the time. In a radical paper published in 1954 that postulated the "dissolution of the concepts of biocoenosis and biotope", Peus in particular elicited a forceful response from many prominent German ecologists. An analysis of the ensuing debate, including especially a colloquium held in 1959 that was partly inspired by Peus' paper, is helpful for sifting the various arguments proffered with respect to a quasi-organismic perception of the biocoenosis in German speaking ecology. Although German mainstream ecologists rejected the notion of the biocoenosis as a superorganism, ontological holism was quite common among them. Additionally, the mainstream concept of the biocoenosis was plagued by several methodological problems and much conceptual confusion, to which the "dissidents" rightly pointed. Some of these problems are still pertinent today, e.g. in connection with more modern concepts such as "ecosystem".