AN ABSTRACT OF THE THESIS OF Joel Bartholemew Hagen for the degree ofDoctor of Philosophy in General Science (Biological Science)presented on 31 July 1981 Title: Experimental Taxonomy, 1930-1950: The Impact of Cytology, Ecology, and Genetics on Ideas of Biological Classification Redacted for privacy Abstract approved: Paul Lawrence Farber During the years between 1930 and 1950 a number of attempts were made to introduce techniques and ideas from cytology, ecology, and genetics into the field of taxonomy. Advocates of this "experimental taxonomy" believed that cytological, ecological, and genetic techniques would provide a rigorous, objective methodology to replace the traditional descriptive methods of taxonomy. Between 1930 and 1950 a number of impressive experimental taxonomic research programs were instituted. Despite the enthusiastic activity in experimental taxonomy, the introduction of cytological, ecological, and genetic methods into taxonomy was problematic. Taxonomists who borrowed from other biological fields were not taking methods and ideas from fully "mature" sciences. Cytology, ecology, and genetics were themselves going through major theoretical and methodological changes. The integration of methods and ideas from a number of complex, semi-autonomous biological disciplines was difficult. Experimental taxonomists, themselves, disagreed on the form this integration should take. Finally, although experimental taxonomy proved to be a viable and fruitful line of research, it did not constitute a complete reform of general taxonomic theory and practice. By 1950 most taxonomists realized that experimental taxonomy was a valuable, but limited, area of research within the more comprehensive discipline of general systematics. qD Copyright by Joel Bartholemew Hagen August 28, 1981 All Rights Reserved Experimental Taxonomy, 1930-1950: The Impact of Cytology, Ecology, and Genetics on Ideas of Biological Classification by Joel Bartholemew Hagen A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Commencement June 1982 APPROVED Redacted for privacy Associate Professor of History of Science in charge of major Redacted for privacy Chairman of Department of General Science Redacted for privacy Dean of GraG.,,, , .., V CT Date thesis is presented July 311 1981 Typed by Jean Kreitlow for Joel Bartholemew Hagen ACKNOWLEDGMENTS At the completion of such a long project it is a pleasure to acknowledge the help that so many people have given me along the way. In particular I owe a debt of gratitude to my adviser, Paul Farber. As teacher, critic, and friend he has been instrumental in bringing this work to a successful conclusion. Without his pa- tience and optimistic encouragement throughout the years this pro- ject would certainly never have been completed. I am also grate- ful to the other members of my dissertation committee. Peter Daw- son, Brookes Spencer, and Charles Warren have all been extremely generous with their time and have provided keen insights and cri- ticism. Lawrence Small was generous enough to serve as graduate school representative on very short notice. I owe much to both formal and informal teachers at Oregon State University. In particular, Paul Farber, Dan Jones, Jim Morris, and Brookes Spencer have contributed toward the develop- ment of my interest in history of science. I have learned much history of biology in a very enjoyable manner through the "Dinner and Reading Club" consisting of Marshall Allen, Keith Benson, David Chapin, and Paul Farber. I have benefited greatly from long discussions on the nature of modern science with Marshall Allen and Walter Hellmann. Depart- To my friends and co-workersin the General Science support and en- ment at Oregon StateUniversity I am grateful for helped to couragement throughout the years. Many friends have like buoy my spirits during thisproject. In particular I would Goldenberg, Diane to thank Kit Ctiesla, Margaret Coleman, Debbie Bunting, Finn Pond, and JeanKreitlow. Jean Kreitlow was generous enough to type this workduring a rather hecticspring. completed Much of the preliminary work onthis project was College. I am during a year asScholar-in-Residence at Cornell of Cornell thankful to the faculty,students, and administration for a most enjoyable periodof unimpeded study. My family has been extremelysupportive and encouraging to my wife, throughout this project. A special thanks goes the thankless tasksof Susan. She enthusiastically undertook Most of filing bibliography cardsand proof-readingdrafts. throughout the allI am grateful for herconstant encouragement completion of this project. TABLE OF CONTENTS Chapter Page INTRODUCTION 1 NOTES 5 1 The New Systematics and the Modern Synthesis 6 NOTES 33 2 The Developing Relationships Between Taxonomy and Cytology, Ecology, and Genetics 43 NOTES 70 3 Experimental Studies in Taxonomy 77 NOTES 106 4 The Critical Reception of Experimental Studies in Taxonomy 113 NOTES 142 5 Experimental Systems of Classification 149 NOTES 185 CONCLUSION 193 BIBLIOGRAPHY 205 LIST OF FIGURES Figure Page 1 Diagram symbolizing the total variability found within a freely interbreeding group of organisms. 59 2 The Coenospecies comprised of species in the genus Zauschneria. 93 3 The Coenospecies Phleum pratense-alpinum. 97 4 Hypothetical relationships among four species in the genus Phleum. 122 5 Alternative explanations for the origin of "artificial Phleum pratense." 124 6 The Coenospecies comprising the species-complex within the genus Phleum as determined by Nordenskiold. 126 7 Turesson's original conception of the genetic and ecological aspects of species. 168 LIST OF TABLES Table Page 1 Biosystematic Units. 157 EXPERIMENTAL TAXONOMY, 1930-1950: THE IMPACT OF CYTOLOGY, ECOLOGY, AND GENETICS ON IDEAS OF BIOLOGICAL CLASSIFICATION INTRODUCTION Much recent research in the hiStory of modern biology has been directed at a limited number of biological episodes. The rise of genet- ics has received a lion's -share of historians' attention.1 The fusion of genetics and natural selection theory into the neo-Darwinian or "syn- thetic theory" of evolution has been another major focus of historical research. The introduction of experimentalism via experimental embry- ology and experimental physiology has been a third area of active his- torical inquiry. While no historian would claim that these episodes constitute all that is important in the development of modern biology, historians have drawn heavily on these episodes for their characteri- zations of twentieth century biology. Perhaps the best example of this type of characterization is pre- sented in Garland Allen's Life Science in the Twentieth Century. According to Allen, "The chief goal of this book is to show how biol- ogy as it was practiced-in the nineteenth centuryin natural history, descriptive and speculative; in physiology, largely mechanisticwas transformed into its twentieth- century mold: experimental, analytically 2 rigorous, and integrative." Allen explicitly disclaims any pretense to writing a comprehensive history of biology. Nonetheless, he portrays developments in genetics, embryology, and physiology as char- acteristic of the modern biological enterprise as a whole.3 2 Among other themes that Allen stresses in his historical account is the influence of experimentalism on the practice of modern biology. The rise of experimentalism is portrayed as coming at the expense of descriptive natural history. This change, Allen rather graphically terms the "revolt from morphology."4Although at least one historian has criticized Allen's emphasis on the antagonism between experimen- talism and descriptive biology, the "naturalist-experimentalist dichotomy" appears to be a widely held historical generalization. In a recently published article, John Dean has analyzed an epi- sode in twentieth century plant taxonomy in terms of the naturalist- experimentalist dichotomy.6The aspect of taxonomy that Dean has chosen, experimental taxonomy, is particularly intriguing because it entails the introduction of experimental methods into what is generally con- sidered a descriptive science. In contrast to Garland Allen, who argues that the experimentalists effectively won the day early in the 1900s, Dean argues that the conflict between experimentalists and descriptive 7 biologists, at least in taxonomy, is very much alive today. In fact, according to Dean, no easy resolution of this conflict is likely to occur in the near future. Experimental taxonomy is an interesting aspect of the history of systematics for a number of reasons. The problematic relationship between experimental and descriptive methods is only one point of his- torical interest. Thenewm-ethods, borrowed largely from cytology, ecology, and genetics, were introduced into systematics primarily after 1920. This was a period of intense ferment in systematics, hence the 3 popularity of the term "new systematics." The developments which sur- rounded experimental taxonomy and the new systematics closely coincided with the rise of neo-Darwinian evolutionary theory. A number of taxon- omists who view the new systematics very differently agree in their close identification of the new systematics with the synthetic theory of evolution.8 The purpose of this dissertation is to examine the introduction of cytological, ecological, and genetic methods into taxonomy and to analyze the impact that these methods had on ideas about the science of classification. In contrast to the analysis