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Schindewolf, Otto Heinrich


(b. Hanover, Germany, 7 June 1896; d. Tübingen, Germany, 10 June 1971), geology, paleontology, evolutionary theory, philosophy of science.

“Evolutionary Theory is in a crisis”: This assessment was Otto Schindewolf’s starting point for his Paleontology, Development, and Genetics: A Critique and Synthesis(1936). George Gaylord Simpson, a leading architect of the mid-twentieth-century “modern synthesis” of evolutionary theory, praised Schindewolf as “the first person to attempt a genuine general synthesis of evolutionary genetics and paleontology. For its time that was an original and brilliant achievement … his initial work in this field was one of the stimuli for the Modern Synthesis, to which he is himself so bitterly opposed” (Simpson, 1952, pp. 388–389). The opposition, according to Simpson, was an old one boldly posed in new terms by Schindewolf as typology versus population concepts. In one of the milestones of the modern synthesis, Ernst Mayr’s Animal Species and Evolution(1963), Schindewolf earned just one approving citation, which referred to his emphasis on the fact that morphological differences between different “types” of invertebrate animals often originate in early developmental stages. But as the modern synthesis hardened, the time seemed right in 1993 to publish an English translation of Schindewolf’s Grundfragen der Paläontologie(1950). Called Basic Questions in Paleontology, it was introduced by one of the most prominent critics of the modern synthesis, Stephen J. Gould, as “a trenchant and utterly consistent (if spectacularly flawed) account of how an uncompromisingly anti-Darwinian, but fully evolutionary, worldview can operate” (p. xi).

Academic Life Schindewolf first became exposed to geology and paleontology in his hometown, Hanover, during his high school years through the Geological Society of Lower Saxony (Niedersächsischer Geologischer Verein). In 1914, Schindewolf enrolled in the natural sciences program at the University of Göttingen, where he majored in paleontology under Rudolf Wedekind, a specialist in biostratigraphy (the identification of the relative geological age of successive sedimentary layers by means of index fossils). In 1917, Schindewolf followed Wedekind to the University of Marburg, where he defended his PhD thesis in 1919. The prize-winning thesis on the biostratigraphy of Late Devonian and Early Carboniferous (Paleozoic) ammonites (fossil cephalopods) from northeastern Bavaria and their paleogeographical significance established Schindewolf’s reputation as a brilliant biostratigrapher. Expanding his research interests, Schindewolf began publishing on other groups of fossil invertebrates as well, most notably corals. His efforts earned him the Habilitation in 1921 (this required the defense of a second, larger thesis on a different topic resulting in the subsequent title of Privatdozent, a step on the way to full professorship) and the title of honorary professor in 1927. It was within these and subsequent years that Schindewolf developed and consolidated his ideas on macroevolutionary processes, leafing through the fossil-bearing strata, much like through the pages of a book, presenting a literal account of evolutionary history. Gaps in the fossil record on the one hand, and the sudden appearance of new types of organisms on the other, seemed to suggest a saltational mode of macroevolutionary change.

Shortly after his promotion in 1927, Schindewolf moved to Berlin, where he had accepted the post of director of the Prussian Geological Survey. Pursuing his interests in macroevolution, Schindewolf inferred a cyclical pattern of evolution from the fossil record, where evolutionary lineages would go through stages of birth and adolescence, maturity, and senescence and death. The biostratigrapher Karl Beurlen independently developed similar ideas, but Schindewolf rejected Beurlen’s vitalism (Wille zur Macht, literally will to power), which he felt to be motivated by Beurlen’s public sympathy for Nazi ideology. Indeed, Schindewolf probably eschewed contact with Beurlen for political reasons. He preferred not to appear at a workshop, Paleontology and Its Relation to Phylogeny, organized by the German Paleontological Society in 1937 in Göttingen with Beurlen as chair. Schindewolf privately distanced himself from the Nazi regime and refused collaboration, in contrast to Beurlen, who set out to criticize Schindewolf ’s work during the Third Reich (before choosing to leave for BRāzīl in 1949). Much later, upon his retirement, Schindewolf would draw parallels between the historical cycles that govern not only evolutionary lineages but also cultures, invoking such authors as Oswald Spengler and Arnold J. Toynbee. This led some to highlight the irony that in his earlier work, Schindewolf used the words “decadence,” “abnormal,” and “degeneration” in characterizing terminal stages of evolutionary lineages, which were terms that were common in Nazi vocabulary. But Schindewolf’s integrity during the Nazi regime is beyond doubt and earned him national and international respect from colleagues and peers.

In 1947, with the Nazis ousted from power, Schindewolf became associate, and soon thereafter full, professor at the Humboldt University in Berlin. He left Berlin in 1948 to take up the position of chair of geology and paleontology at the Eberhard Karls University in Tübingen. In 1956–1957 he served as president (Rektor) of that University, from which he retired in 1964. He continued research on ammonite phylogeny, and in 1969, he published a theoretical paper in which he offered important qualifiers with regards to his magnum opus of 1950, Grundfragen der Paläontologie. The 1950 text was written in Berlin under difficult conditions during World War II, and its publication was delayed due to the poor economic conditions that prevailed in Germany after the war. The English translation testifies to the fact that it was the Grundfragen above all else that established Schindewolf’s reputation as a preeminent if controversial evolutionary theoretician, not only in Germany, but internationally. During his long and active life as a teacher and researcher, Schindewolf served as editor of various professional journals (Paläontologisches Zentralblatt, Fortschritte der Paläontologie, Palaeontographica, and Neues Jahrbuch für Geologie und Paläontologie), initiated the highly influential book series Handbuch der Paläozoologie, and was member of various professional societies and higher academic bodies (Akademie der Wissenschaften und der Literatur Mainz, Akademie der Naturforscher Leopoldina in Halle, and the academies of sciences of Heidelberg, Germany; Vienna, Austria; and Lund, Sweden).

Biostratigraphy and Macroevolution Schindewolf was first and foremost a biostratigrapher. The method of biostratigraphy requires clearly and unequivocally identifiable and re-identifiable species, or taxa, of fossils that by virtue of their distribution through time characterize a certain time horizon line in the column of sedimentary layers of rock laid down during earth’s history. Naturally, biostratigraphers look for species, or taxa, that (1) are well demarcated from other fossil taxa, (2) make a relatively sudden appearance in the fossil record, (3) persist relatively unchanged, and (4) disappear relatively suddenly again. In his studies of ammonites and their use as index fossils, Schindewolf took great care to establish the degree of individual variation in order to ensure that the biozones he recognized were delimited by clearly diagnosable species. This required the investigation of the individual development of ammonite fossils, because much variation can potentially be expressed during ontogeny, such that juveniles and adults must not be mistaken for different species. His interest in the ontogeny of ammonites led to inquiries into the causal mechanisms that drive and regulate the developmental process, which through all variation nevertheless preserves the “typical” form of the species. With his Paläontologie, Entwicklungslehre, und Genetik(1936), Schindewolf became the first author to attempt a synthesis of (pre–World War II) genetics with the study of the ontogeny and of the morphology of fossil organisms, a synthesis that he naturally pursued against the background of his biostratigraphical research program.

The theories of genetics that best seemed to match the stratigraphic record of fossils—marked by gaps and the sudden appearance of new forms of life, some as start-ingly new as the earliest bird Archaeopteryx—were those of Richard Goldschmidt. Schindewolf’s adviser, Wedekind, was well acquainted with Goldschmidt’s work, and in “Über den ‘Typus’ in morphologischer und phylogenetischer Biologie Schindewolf” (1969), Schindewolf himself reminisced about many discussions he had with Gold-schmidt when they both lived and worked in Berlin. Goldschmidt’s primary interest was the genetic control of developmental processes, where relatively minor early deviations were thought to potentially result in significant changes at the adult stage. Goldschmidt’s claim to fame was accordingly his theory of macromutations, which later developments in genetics proved to be mistaken. Yet the result of Schindewolf’s early synthesis, which preceded the spread of population genetics triggered in 1937 by Theodosius Dobzhansky, was accordingly a theory of saltational macroevolution known as typostrophe theory (typostrophentheorie). Its central tenets were that (1) internal (structural), not external (natural selection) factors drive macroevolution, which therefore, unlike microevolution, is not adaptive; (2) macroevolution is saltational; and (3) macroevolution is cyclic. During the first phase of an evolutionary cycle, called typogenesis, a “type,” or Bau-plan, is laid down that is adapted only in the sense that it is capable of surviving environmental changes through time. In stage two, called typostasis, the general “type,” or Bauplan, is maintained as it diversifies through time and space in adaptation to special and varying conditions of life at low taxonomic levels. Stage three, called typolysis, leads to overspecialization and eventual extinction of the evolutionary lineage representing the “type,” or Bauplan. With his Der Zeitfaktor in Geologie und Paläontologie(1950), Schindewolf became the first author in modern times to invoke extraterrestrial causes such as cosmic radiation generated by a supernova (1954, 1963) to explain the mass extinctions at the end of the Paleozoic and Mesozoic periods.

The “Type” Concept The concept that lies at the core of Schindewolf’s typostrophism is that of a “type” of organization, or Bauplan. The use of this concept identifies Schindewolf as an exponent of the neo-idealistic morphology that became popular among German-speaking morphologists and paleontologists after World War I. Following the rise of Darwinism, morphological research was caught between “pattern” versus “process” views of nature. The architects of the modern synthesis of evolutionary theory proclaimed “population thinking,” with population genetics at its heart, as the only proper way to understand Darwinian evolution. Ever so slight variations and the force of natural selection would result in gradual evolutionary change in a continuous chain of populations extending through time and space. If intermediate populations were absent in the fossil record, their existence had to be inferred on the basis of evolutionary theory. Idealist morphologists, on the other hand, emphasized the primacy of structure, and of the hierarchical pattern that results from the comparative analysis of anatomy, over its historical explanation (evolution). Morphology was to provide the empirical base for the natural system, and if evolutionary theory was to offer a causal-historical explanation of that system, it had to take into account its structure, even if that meant to account for gaps to be bridged by saltational change.

Schindewolf first laid down his views on idealistic morphology as the basis for biosystematics and phylogenetic research in his influential paper of 1928, “Prinzipienfragen der biologischen Systematik,” which had its origin in a seminar he presented at Marburg University in 1921. He revisited his type concept in his magnum opus of 1950, and once again returned to its defense in 1969 with his “Über den ‘Typus.’” Ernst Mayr, a leading author of the modern synthesis, dismissed the type concept as a Platonic idea (eidos), rendering it the hallmark of the worst possible fallacy in evolutionary biology, typological thinking. In “Über den ‘Typus,’” Schindewolf naturally rejected Mayr’s identification of the type concept as merely a Platonic idea, or a phantasmagoria, as it was called by the botanist Walter Zimmermann. All such arguments Schindewolf considered as resulting from a lack of understanding of the paleontological method. Ichthyostega stands for the first tetrapod, Archaeopteryx is the first representative of the “type” of birds, and both make a sudden appearance in the fossil record. In that sense, tetrapods, or birds, represent a “type,” which is a genuinely scientific concept, and as such objective and real. The sudden appearance of new “types” is what paleontologists observe, and—according to Schindewolf— scientific speculation should not exceed the empirical base to an irresponsible degree. This, however, is exactly what Schindewolf found “population thinking” to be guilty of, as it bridged the gaps between types with hypothetical intermediate populations for which there was no empirical evidence: “I cannot offer up populations,” Schindewolf wrote (“Über den ‘Typus,’” p. 88).

In his highly influential book on evolutionary theory, Deszendenzlehre (1922), Sinai Tschulok sought a compromise between “Machian permissiveness” (a reference to Ernst Mach) and “Millian rigidity” (John Stuart Mill). With reference to evolutionary theory, he distinguished three issues: (1) the “principle issue” (Grundsatzfrage), which settles the question that evolution did, in fact, occur; (2) the “phylogenetic tree issue” (Stammbaumproblem), which is concerned with the reconstruction of the history of life on Earth; and (3) the “causal issue” (Faktorenproblem), which is concerned with the causes that drive the evolutionary process. According to Tschulok, it simply did not make sense either to search for phylogenetic relationships, or to investigate the causes of evolutionary change, without prior acceptance of the fact that evolution as such did occur. The Göttingen workshop of 1937 that Schindewolf had preferred not to attend had concluded that paleontology is the only science that has direct epistemic access to the evolutionary process, that the goal of paleontology therefore must be to reconstruct the history of that process, and that the causes driving this process could at best be inferred from its systematic reconstruction. Regretting his mistake of having accepted Goldschmidt’s macromutations in his early theorizing, Schindewolf followed the eminent paleontologist Walter Gross in arguing that the paleontologist has nothing to say about the causal issue. The task of paleontology, Schindewolf wrote in ““Über den ‘Typus,’” is simply “the explanatory description of the empirically accessible regularities apparent in the Fossil Record and nothing else” (p. 107). He accordingly distinguished paleontology as a nomographical science, in contradistinction to the idiographic and nomothetical sciences that had been recognized by Wilhelm Windelband. On that account, paleontology comes out as a primarily historical science in the sense of Leopold von Ranke, (1795–1886), who said “to state, how it [actually] was” (Schindewolf, 1969, p. 106). It was not, accordingly, one concerned with the understanding of historically unique situations, as were Windelband’s idiographic sciences, nor one concerned with the discovery of universal laws of nature, as were Windelband’s nomothetical sciences, but one that seeks the “explanatory description” of regularities from which empirical generalizations can be inferred.

The difficulty of understanding the type concept may have its roots in what appear to be inconsistencies in the idealistic morphological literature. For the early Schindewolf in “Prinzipienfragen der biologischen Systematik” (1928), biosystematics was a purely ordering science and its categories, consequently, are purely conceptual, that is, spatiotemporally unrestricted. These categories are the hierarchically nested and/or stratigraphically stacked “types,” or Baupläne. In his Grundfragen, Schindewolf maintained the need for logical abstraction that is required to conceptualize “types,” but he denied that these represent mere mental constructs. Instead, the “types” marked out by morphology are the result and the expression of the concrete and real phylogenetic process. The hierarchy of “types” must therefore be considered a hierarchy of historical entities, marked out by “typical” characters that have real, indeed vital, significance for the token organisms that exemplify the corresponding “types.” In his later reflections, Schindewolf again stressed the “objective biological reality” of the “type,” a “scientific reality” that equals the reality of populations. Schindewolf went on to discuss the latest applications of the “type” concept by other authors both in biology and anthropology, and registered his satisfaction that the philosopher Marjorie Grene, in her discussion of “Two Evolutionary Theories” (1958), had come down closer to his own views than to those of George G. Simpson. Indeed, Schindewolf, like most other neo-idealistic morphologists, followed the lead of Adolf Naef, who had called the “type” a concept, or a kind, that is, a natural kind, in his Idealistische Morphologie und Phylogenetik (1919). To consider the “type” of neo-idealistic morphology a natural kind raises the question whether these kinds represent immutable and timeless Aristotelian classes of naturally occurring “stuff” or “things,” or whether they can also be exemplified by natural historical entities such as species and supraspecific taxa. On a charitable reading, it seems that from 1928 to 1969, Schindewolf progressed from the former to the latter interpretation of types in his writings.

Schindewolf’s Legacy During his long career, Schindewolf published 187 books, book chapters, monographs, and papers and trained more than sixty graduate students. His scientific output, his dedication to teaching and education, and his personal integrity established him as the leading paleontologist of his time in Germany. He was the first author in Germany to attempt a synthesis of genetics, development, morphology, and paleontology, and in so doing developed his own internally coherent but nonselectionist and saltational theory of macroevolution. Naturally, his typostrophism found little acclaim on an international stage that was dominated by the modern synthesis of evolutionary thought. There is good reason to believe that Schindewolf’s towering presence delayed the acceptance of the synthetic theory of evolution in German paleontology. In the late twentieth century, however, proponents of the “punctuated equilibria” theory of evolution found in Schindewolf—the man who had famously adopted the statement “the first bird hatched from a reptile’s egg” (Paläontologie, Entwicklungslehre, und Genetik,

p. 59)—a brother in arms, even if only a distant one. Both Schindewolf’s typology and the model of punctuated equilibrium of evolution have their roots in a biostrati-graphic research program, which, for its successful execution, requires sharp boundaries between taxa.

While biostratigraphy was the main motivating factor for Schindewolf’s typostrophe theory, he also found that developmental biology lent support to a typological perspective of organismic diversity. Deviation of developmental pathways during early stages of ontogeny may cause changes in adult morphology that create gaps in the continuity of organic forms. Such gaps in morphospace, at the turn of the twentieth to the twenty-first century the subject matter of the evolution and development research program, were prominently used by Gerry Webster and Brian Goodwin in “The Origin of Species: A Structuralist Approach” (1982), their critique of neo-Darwinism. Their typological perspective is motivated by a field theory of generative mechanisms. In most general terms, a “field is a spatial domain in which every part has a state determined by the state of neighboring parts so that the whole has a specific relational structure” (Goodwin, “What Are the Causes of Morphogenesis?” 1984, p. 228). The result is that any disturbance of the field requires the whole spatial pattern to be reconstituted. Morphogenetic fields that play out during development thus condition “types,” that is, they constrain development to typical relations of morphology that mark out empirically discovered natural kinds. The goal is a structuralist morphology that delivers a rational classification of organisms in the form of a hierarchy of “types,” which in turn is causally explained by the physico-chemical properties of the underlying morphogenetic fields and their transformations. The answer to Darwinian theories of variation and natural selection is that “no Darwinist has ever understood typology, because he can not possibly understand it” (Webster and Goodwin, “The Origin of Species,” p. 26). In Schindewolf’s case, the success of applied biostratigraphy may have made it impossible for him to understand the biological processes underlying Darwinian evolutionary theory.


A complete bibliography of Schindewolf is given in Heinrich K. Erben, “Nachruf auf Otto Heinrich Schindewolf,” Jahrbuch 1971 der Akademie der Wissenschaften und der Literatur in Mainz (1971), pp. 75–86. A selection of Schindewolf’s most important publications in biostratigraphy, evolution, and systematics, was compiled by Wolf-Ernst Reif in his “Afterword” for Otto Heinrich Schindewolf, Basic Questions in Paleontology: Geologic Time, Organic Evolution, and Biological Systematics, translated by Judith Schaefer (Chicago: University of Chicago Press, 1993), pp. 435–453. Universitätsarchiv Tübingen, Wilhelmstrasse 32, D-72074 Tübingen.


“Prinzipienfragen der biologischen Systematik.” Paläontologische Zeitschrift 9 (1928): 122–166.

“Ontogenie und Phylogenie.” Paläontologische Zeitschrift 11 (1929): 54–67.

Paläontologie, Entwicklungslehre, und Genetik: Kritik und Synthese[Paleontology, development, and genetics: A critique and synthesis]. Berlin: Gebrüder Bornträger, 1936.

“Beobachtungen und Gedanken zur Deszendenzlehre.” Acta Biotheoretica 3 (1937): 195–221.

“Zum Kampf um die Gestaltung der Abstammungslehre.” Die Naturwissenschaften 31 (1944): 269–282.

Grundfragen der Paläontologie: Geologische Zeitmessung, organische Stammesentwicklung, biologische Systematik. Stuttgart, Germany: E. Schweizerbart, 1950.

Der Zeitfaktor in Geologie und Paläontologie. Stuttgart, Germany: E. Schweizerbart, 1950.

“Über die möglichen Ursachen der grossen erdgeschichtlichen Faunenschnitte.” Neues Jahrbuch für Geologie und Paläontlogie, Monatshefte (1954): 457–465.

“Neue Systematik.” Paläontologische Zeitschrift 36 (1962): 59–78.”

Neokatastrophismus?” Zeitschrift der Deutschen Geologischen Gesellschaft 114 (1963): 430–445.

“Erdgeschichte und Weltgeschichte.” Akademie der Wissenschaften und der Literatur (Mainz), Abhandlungen der Mathematisch-Naturwissenschaftlichen Klasse 2 (1964): 56–104.

“Über den ‘Typus’ in morphologischer und phylogenetischer Biologie.” Akademie der Wissenschaften und der Literatur (Mainz), Abhandlungen der Mathematisch-Naturwissenschaftlichen Klasse 4 (1969): 55–131.

Basic Questions in Paleontology: Geologic Time, Organic Evolution, and Biological Systematics. Translated by Judith Schaefer. Edited by Wolf-Ernst Reif. Foreword by Stephen Jay Gould. Chicago: University of Chicago Press, 1993.


Beurlen, Karl. “Die gegenwärtige Stellung der Paläontlogie zu den Hauptproblemen der Stammesgeschichte.” Paläontologische Zeitschrift 20 (1938): 167–176.

Dobzhansky, Theodosius. Genetics and the Origin of Species. New York: Columbia University Press, 1937.

Eldredge, Niles. “Genetics and the Origin of Species.” Evolution 25 (1971): 156–167.

Erben, Heinrich K. “Nachruf auf Otto Heinrich Schindewolf.” Jahrbuch 1971 der Akademie der Wissenschaften und der Literatur in Mainz (1971): 75–86. Includes a complete bibliography.

Goodwin, Brian C. “What Are the Causes of Morphogenesis?” In Beyond Neo-Darwinism: An Introduction to the New Evolutionary Paradigm, edited by Mae-Wan Ho and Peter T. Saunders. London: Academic Press, 1984.

——. “What Are the Causes of Morphogenesis?” BioEssays 3 (1985): 32–36.

Grene, Marjorie. “Two Evolutionary Theories.” British Journal for the Philosophy of Science 9 (1958): 110–127, 185–193.

——.“Evolution, ‘Typology,’ and ‘Population Thinking.’” American Philosophical Quarterly 27 (1990): 237–244. Korn, Dieter. “Schindewolf, Otto Heinrich.” In Encyclopedia of Paleontology, edited by Ronald Singer. Vol. 2. Chicago: Fitzroy-Dearborn Publishers, 1999.

Kullmann, Jürgen, and Jost Wiedmann. “Otto H. Schindewolf.” Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 125 (1966): xi–xxvii (Festband Schindewolf).

——. “Otto Heinrich Schindewolf zum Gedächtnis, 7.6.1896–10.6.1971.” Atempto Tübingen 39–40 (1971): 120–121.

Mayr, Ernst. Animal Species and Evolution. Cambridge, MA: Harvard University Press, 1963.

Morris, Simon Conway. “Wonderfully, Gloriously Wrong.” TREE 9 (1994): 407–408.

Naef, Adolf. Idealistische Morphologie und Phylogenetik (zur Methodik der systematischen Morphologie). Jena, Germany: Gustav Fischer, 1919.

Reif, Wolf-Ernst. “Evolutionary Theory in German Paleontology.” In Dimensions of Darwinism: Themes and Counterthemes in Twentieth-Century Evolutionary Theory, edited by Marjorie Grene. Cambridge, U.K.: Cambridge University Press, 1983.

——. “The Search for a Macroevolutionary Theory in German Paleontology.” Journal of the History of Biology 19 (1986): 79–130.

——. “Afterword.” In Basic Questions in Paleontology: Geologic Time, Organic Evolution, and Biological Systematics, by Otto Heinrich Schindewolf, translated by Judith Schaefer. Chicago: University of Chicago Press, 1993.

——. “Typology and the Primacy of Morphology: The Concepts of O. H. Schindewolf.” Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 205 (1997): 355–371.

——. “Deutschsprachige Paläontologie im Spannungsfeld zwischen Makroevolutionstheorie und Neo-Darwinismus (1920–1950).” Verhandlungen Geschichte und Theorie der Biologie 2 (1999): 151–188.

Seilacher, Adolf. “Otto H. Schindewolf, 7 Juni 1896–10 Juni 1971.” Neues Jahrbuch für Geologie und Paläontologie, Monatshefte (1972): 69–71.

Simpson, George Gaylord. Obituary. Quarterly Review of Biology 27 (1952): 388–389.

Teichert, Curt. “From Karpinsky to Schindewolf—Memories of Some Great Paleontologists.” Journal of Paleontology 50 (1976): 1–12.

Tschulok, Sinai. Deszendenzlehre. Jena, Germany: Gustav Fischer, 1922.

Webster, Gerry. “The Relations of Natural Forms.” In Beyond Neodarwinism. An Introduction to the New Evolutionary Paradigm, edited by Mae-Wan Ho and Peter T. Saunders. London: Academic Press, 1984.

Webster, Gerry, and Brian C. Goodwin. “The Origin of Species: A Structuralist Approach.” Journal of Social and Biological Structures 5 (1982): 15–47.

Oliver Rieppel

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