ALBERT

Notes: Summary Six schools of thought can be detected in the development of evolutionary theory in German paleontology between 1859 and World War II. Most paleontologists were hardly affected in their research by Darwin's Origin of Species. The traditionalists (School 1) accepted evolution within lower taxa (genera and families) but not for organisms in general. They also rejected Darwin's theory of selection. The early Darwinians (School 2) accepted Darwin's theory of transmutation and theory of selection as axioms and applied them fruitfully to the fossil record, thereby laying the foundation for the new research areas of phylogeny and paleo-biology. The enthusiasm of the early Darwinians faded when the fossil record and the problems of its interpretion became more widely known. The pluralists of the turn of the century (School 3) invented and adopted a wealth of hypothetical mechanisms in order to explain individual features of the fossil record. They failed, however, to provide one coherent theory. Dissatisfaction with this situation led to adoption of a dogmatic neo-Lamarckism (School 4), which was regarded as a coherent theory providing a fruitful research program. The rejection of the Lamarckian mechanism early in this century left paleontologists with only one kind of evolutionary mechanism: inner causes. Like many neo-Lamarckians several orthogeneticists (School 5) were highly interested in adaptation and did not see any contradiction between the inner causes of evolution and adaptation. The dominance of stratigraphical research programs in paleontology led in the 1930s and 1940s to a decrease in interest in adaptation. Stratigraphical records of taxa were accepted as meaningful in the context of evolutionary theory. Orthogenesis and the new concepts of saltation and cyclicism were amalgamated into one theory: typostrophism (School 6). This theory dominated German paleontology for decades after the war and only recently has the synthetic theory been seriously considered. Evolution was never very intensively discussed in German paleontology in the hundred years after Darwin's book. Most information used here comes from textbooks or from papers given on special occasions. It has been impossible to summarize how members of one school defended their views or discussed the ideas of competing schools.

Notes: Conclusion With the rejection of group selectionist derivations of ecological phenomena so incisively given by George Williams in 1966,43 Nicholson's long-ignored messages met with acceptance. Species benefit became, explicitly, incidental. But the reorientation was not just about a point of ecological theory. It was more fundamentally about theoretical style, the element shared by Wynne-Edwards' work and the newer, evolutionary ecology. That current approach is well expressed in an already classic paper by the British plant ecologist John Harper: Ultimately all the discoveries of descriptive, production or population ecology must find their meaning in evolutionary phenomena... Evolutionary thinking concentrates attention on the behaviour of the individual and his descendants. If nothing in biology has meaning except in the light of evolution and if evolution is about individuals and their descendants — i.e. fitness — we should not expect to reach any depth of understanding from studies that are based at the level of the superindividual... What we see as the organised behaviour of systems is the result of the fate of individuals.44 The emphasis in this passage is on style of thinking more than on study matter. What is so different from earlier ecologists is the rejection of the superorganism and its associated baggage of selfimposed constraints in an end-oriented process, such as balance of nature or species-level adaptation. The hallmark of post-1959 evolutionary ecology is the recognition, the making explicit of the question that the selective, genetical process of fitness (fates of genes) has to be separated from the observations of different levels of adaptiveness. An easy conflation of cause and result is no longer satisfying; now attempts to disentangle the complex strands tying population phenomena to the different levels of selection are providing the excitement of a new research program. In this program evolutionary theory is used to define the appropriate questions, as exemplified by the organization of the textbook Evolutionary Ecology.45 Our discussion of the invention of a new approach, with a characteristic style and set of interests, must also be about how the practitioners of ecology have perceived themselves. The early attempts of Poulton and Nicholson were not taken up by the vigorously growing science of ecology, which developed first - and intentionally — along a narrower, hierarchically restrained set of questions about population. The depth of Poulton's and Nicholson's submergence is indicated by the fact that even Wynne-Edwards did not argue in the manner of one rescuing a minor tradition. Rather, he thought he had a new approach, that of placing evolution in the foundations of ecology. Although this approach was not exactly original, we should be cautious in grouping all evolutionary concerns within ecology in one intellectual tradition. The particular style of selectionist deduction invigorated by Wynne-Edwards was successful only in the volatile situation of a rapidly growing discipline, full of self-critical examination and new rigor. Whereas in 1930 evolutionary concerns did not provide that rigor, by 1960 they could - and did.

Notes: Conclusion The distinction between taxonomic plant geography and ecological plant geography was never absolute: it would be historically inaccurate to portray them as totally divergent. Taxonomists occasionally borrowed ecological concepts, and ecologists never completely repudiated taxonomy. Indeed, some botanists pursued the two types of geographic study. The American taxonomist Henry Allan Gleason (1882–1975), for one, made noteworthy contributions to both. Most of Gleason's research appeared in short articles, however. He never published a major synthetic work comparable in scope or influence to the ecological texts of Clements, Schimper, and Warming. Despite exceptions such as Gleason, most plant geographers throughout the twentieth century have emphasized the distinction between ecological and taxonomic plant geographies. Why have these distinct traditions developed? In his book Geographical Ecology, Robert MacArthur has suggested a psychological explanation for the dichotomy: “Unraveling the history of a phenemenon has always appealed to some people and describing the machinery of the phenomenon to others... The ecologist and physical scientist tend to be machinery oriented, whereas paleontologists and most biogeographers tend to be history oriented.”46 Without necessarily rejecting MacArthur's explanation, my study suggests a more complex relationship between taxonomic and ecological plant geographies. At the turn of the century a group of botanists self-consciously defined a new area of botanical research. These ecologists defined their new discipline in opposition to what they believed was a moribund, nineteenth-century, natural-history tradition. They turned from historically oriented, descriptive, taxonomic plant geography to experimental physiology. The new ecological plant geography was to focus on communities rather than on species, on proximate environmental causes rather than on historical explanations, and on physiological experiments rather than on morphological descriptions. As we look back, much of the “revolt from morphology” was rhetorical. Ecologists never completely replaced species as units of distribution, nor did they set geography on an explicitly physiological basis. Indeed, much of early ecological research was, quite simply, descriptive. Plant communities were defined in terms of dominant species, representative life forms, or general physiognomy. The underlying physiological basis for community characteristics was more often assumed than demonstrated by experiments. Despite the fact that ecological plant geography was not a truly physiological specialty, it was significantly different from more traditional taxonomic plant geography. First, ecologists were less explicitly evolutionary in their approach than were taxonomists. Following Darwin, most taxonomic plant geographers viewed distribution in historical terms. In contrast, early ecologists tended to ignore the traditional geographic problems. Most ecologists were skeptical of historical explanations, emphasizing instead the proximate, environmental causes of distribution. While some nineteenth-century biogeographers had studied the correlation between climate and vegetation, twentieth-century ecologists focused much more sharply on the interactions between plant and environment. Plant ecologists did not place biogeography on a physiological basis, but by emphasizing physiology they laid the foundation for a more detailed understanding of adaption. This emphasis on physiology and environmental causation was a second distinguishing characteristic of ecological plant geography. Finally, the idea of the plant community, articulated by Eugenius Warming in 1895, provided ecologists with a unique perspective on the distribution of plants. For early ecologists, the community was more than an assemblage of species; it was an integrated unit. The distribution of these units became the major focus of ecological plant geography. Communities never completely replaced species as geographic units, and the distinction between flora and vegetation was often blurred. Nonetheless, ecologists were innovative in studying the distribution of structurally and functionally integrated groups of plants. In the twentieth century plant geography has occupied an anomalous position in biology. It has not developed into an autonomous discipline, nor has it been incorporated into the developing discipline of ecology. Ecologists and taxonomists have pursued fairly distinct styles of geographic research, with the result that two relatively independent approaches to the study of plant distribution have persisted.

Notes: Conclusion We should now be able to come to some general conclusions about the main lines of Cuvier's development as a naturalist after his departure from Normandy. We have seen that Cuvier arrived in Paris aware of the importance of physiology in classification, yet without a fully worked out idea of how such an approach could organize a whole natural order. He was freshly receptive to the ideas of the new physiology developed by Xavier Bichat. Cuvier arrived in a Paris also torn by many overlapping debates on the nature of classification, and in particular that between the natural and artificial systems. The very validity of the enterprise of classification was questioned in many quarters. Cuvier's achievement on his entry into the Parisian world of science was not simply to establish himself as a highly competent anatomist: far more important, he also began to use ideas from many different specialties to change completely the notion of what was involved in natural history.124 At the same time that he himself swung away from the guiding image of the field naturalist as the ideal of the specialty, he took ideas from the new physiology to answer questions about the order of the animal world, and from comparative anatomy to resurrect extinct creation — and to come to conclusions from that creation about the history of the forms of life and the manner of their succession. He showed himself able to alter the relationships between natural history and many other fields of study in a way that implied, rightly or wrongly, his own complete mastery over such a movement. Partly he was able to do this because the ideas he borrowed were not themselves logically articulated and thus could be easily adapted and refocused for many different specific purposes. The value of the heuristic possibilities inherent in the idea of life, for example, far outweighed its inability to generate full systems of classification. Cuvier also consistently refused to consider in science matters relating to the first causes of events. Freed from the consideration of first-order phenomena, he was able to use second-order explanations across a far wider field of applicability. Personal doubts about the validity of a theology that had used science in order to bolster its own claims were combined here with the strong influence of the Kantian critique of the limits of human reason.125 Cuvier's characteristic mode of procedure was that of intellectual appropriation and a bold capacity for altering the relationships between different fields of knowledge, rather than, with the exception of taxonomy, the technique of expanding their subject matter. His claims to originality came, first, from this reappropriation and reorientation and, second, from the sheer scope of his work, which aimed at nothing less than the cataloging and classification of all animate objects.126 They rested also on his acute use of his assertion of a certain relationship with the past of his subject. Very often he would present this history in such a way as to obscure his own intellectual genealogy, and often too he would give differing accounts of the priority of use of an idea in order to distract attention from the questionable exactitude of his own claims to originality. Cuvier came to Paris at precisely the time when society and institutions were most profitably malleable for a newcomer; it was also a time when many scientific disciplines had reached a stage advanced in terms of their factual content, yet relatively inadequate in conceptual organization. They were ripe for takeover by large-scale organizing ideas such as the animal economy and the subordination of characteristics. Paleontology is a particularly good example of a specialty in this particular form of underdevelopment in 1795. Cuvier paid a high price for his initial success. His electic applications of large-scale organizing ideas tended to mean that little of his own work had complete coherence at all levels. Ideas, as we have seen, that proved capable of providing a complete reform of the larger groups of the animal kingdom were incapable of producing its detailed working-out in the taxonomy of smaller groups, which had to be supplied from observed analogical correlations. Further, his physiological approach to classification involved the breakdown of strict correspondence between organs and functions, which left the way open for workers such as Geoffroy St. Hilaire gradually to tilt the balance away from the study of the correlations of hierarchies of functions, and toward morphology as the basis of the order of nature. Cuvier's brilliant appropriations from physiology from the beginning, therefore, contained the seeds of conflict with Geoffroy. Cuvier's eclectic approach made it very nearly impossible for him to present a clear idea of the ways in which the life sciences could be said to be lawful. In spite of his efforts to assimilate them to the position of the physical sciences in this respect, he was forced in the end to accord only an ambiguous status as “laws” to observational correlations. From this area of failure came much of the attempt to give his own two laws — the correlation of parts and the subordination of characteristics — predictive qualities, particularly in relation to paleontological research. It is not surprising that Cuvier's title as the “legislator” of natural history should represent more a claim than a reality. How, then, was he able to emerge as the leading French naturalist of his day? First of all must be adduced the sheer scale of his undertakings. Then comes his expertise as a practical anatomist, and the range of different topics toward which he turned his interest. His collaborators cannot be given credit for his output nor, as we have seen, for slavish adherence to his ideas. Cuvier was able to successfully claim to have dominated the underdeveloped specialties, such as paleontology, and turned them into a major heuristic input into both geology and comparative anatomy; but in other fields, such as physiology, he appropriated concepts and encouraged research but made little impact on the field himself. His attempts in 1812 to head off, or neutralize and absorb the growth of morphological studies landed him in a dangerously rigid position, which despite his encouragement of the new physiological research under the Restoration made further elaboration of his own conceptual underpinnings almost impossible. Cuvier's authority in the scientific world would in any case have been great because of his substantive achievement in taxonomy, but the rest of his work had enough ambiguities and dislocations for it to need the support of his political and social power. Cuvier's detractors seized on a vital fragment of the truth when they accused him of finding the political dimension all-important: it obscured the disjunctions in his theories and at the same time gave him the authority to make new claims for the status of the observational sciences - and for their relations of power with their surrounding specialties. Cuvier's science both thrived on and was halted by the power games of intellectual appropriation, manipulation of the past to confirm the present, and continual claims for hegemony.