The reception of Darwin in late nineteenth-century German paleontology as a case of pyrrhic victory

Studies in History and Philosophy of Biological and Biomedical Sciences xxx (2017) 1e9

Contents lists available at ScienceDirect

Studies in History and Philosophy of Biological and Biomedical Sciences journal homepage: www.elsevier.com/locate/shpsc

The reception of darwin in late nineteenth-century German paleontology as a case of pyrrhic victory Marco Tamborini Institute of Philosophy, Technische Universität Darmstadt, Landwehrstraße 54, D-64293 Darmstadt, Germany

a r t i c l e i n f o

a b s t r a c t

Article history: Received 30 March 2017 Received in revised form 16 September 2017 Available online xxx

This paper investigates German-speaking paleontologists’ reception of Darwin’s thought and the ways in which they negotiated their space of knowledge production accordingly. In German-speaking regions, the majority of paleontologists welcomed Darwin’s magnum opus, since it granted paleontology an independent voice within biology, and thus a new institutional setting. However, in the process of negotiating the features of paleontology within the Darwinian framework, German paleontologists constrained their practices too narrowly, for fear of leaving open possible results at odds with the burgeoning Darwinian biological community. In doing so, they also limited the further development of German paleontology. In other words, paleontologists Karl Alfred von Zittel (1839e1904) and Melchior Neumayr (1845e1890) advocated for a handmaid’s role for paleontology, which increased biologists’ dependence on paleontologists for empirical evidence, but which limited paleontologists’ theoretical autonomy. By analyzing both the institutional strategies and the methodology of German-speaking paleontology at the end of the nineteenth century, this paper shows the importance of scientists’ ability to enter into and negotiate their place within the broader biological community. Ó 2017 Elsevier Ltd. All rights reserved.

Keywords: Darwin Series of forms Karl Alfred von Zittel Melchior Neumayr Labeling Morphology

“The armies separated; and, it is said, Pyrrhus replied to one that gave him joy of his victory that one other such victory would utterly undo him.”

Richards, 2008). Yet historians have paid little attention to paleontology. The reading of Darwin’s masterpiece deeply transformed German-speaking1 paleontology; they changed their ways of doing

Plutarch, Pyrrhus Within the comparative studies on Darwin’s reception, Germany assumes a particularly prominent role. As philosopher Robin Craw put it, “Darwinism found more supporters among biologists and the general intellectual audience in Germany than in any other country” (Craw, 1992, p. 68). Paleontologist and historian of science Wolf-Ernst Reif (1945e2009) even wrote that “Darwinism was so fully accepted in the first decades after publication of Origin that it wiped out the memory and influence of the Naturphilosophie and idealists” within the German scientific community (Reif, 1986, p. 110). The reception of Darwin’s theory among some disciplines and scientists within the German academic landscape has been investigated in depth: The rise and decline of evolutionary morphology endorsed by Ernst Haeckel (1834e1919) and Karl Gegenbauer (1826e1903) has been, for instance, a highly debated topic (Breidbach, 2003; Di Gregorio, 2005, 2008; Nyhart, 1987, 1995;

E-mail address: [email protected].

1 My focus here is not merely on the confederation of states that had a political affiliation throughout the nineteenth century and eventually made up the unified German nation-state in 1871, but rather on the German-speaking paleontologists. As I show, despite the political differences, a particular intellectual link connected the German-speaking paleontologists throughout the nineteenth and twentieth centuries. All these paleontologists studied and worked together. They continually exchanged knowledge and data, thus creating a scientific network, which put aside local and regional conflicts. For instance, after his education in Bavaria and Baden, paleontologist Melchior Neumayr became professor for paleontology in Vienna and continued networking with his German colleagues. This, though, does not mean that I treat the German-language area as one in terms of approach. Conversely, I am deeply aware of the different regional contexts, which in turn shaped different practices and approaches to biology. For instance, as Deborah Coen noted, “in Prussia and Baden, with their burgeoning electrical and chemical industries, liberals concerned themselves more immediately with industrialization than their Habsburg counterparts and promoted science for practical ends” (Coen, 2007, p. 12). Indeed, during the first decades of the twentieth century, a different premium on the industrialization of science gave rise to distinct and contrasting models of knowledge production and dissemination, advocated respectively by the director of the Geological-Paleontological Museum in Berlin Wilhelm von Branca (1844e1928) and the Austrian paleontologist Othenio Abel (1875e1946). A detailed investigation of early twentieth-century natural history and German polycentrism can be found in (Tamborini, 2016).

https://doi.org/10.1016/j.shpsc.2017.09.004 1369-8486/Ó 2017 Elsevier Ltd. All rights reserved.

Please cite this article in press as: Tamborini, M., The reception of darwin in late nineteenth-century German paleontology as a case of pyrrhic victory, Studies in History and Philosophy of Biological and Biomedical Sciences (2017), https://doi.org/10.1016/j.shpsc.2017.09.004

2

M. Tamborini / Studies in History and Philosophy of Biological and Biomedical Sciences xxx (2017) 1e9

science in their encounter with the Origin of Species. As the great German paleontologist Karl Alfred von Zittel (1839e1904) asserted, “Darwin’s conception of the origin of species could not fail to enhance the interest of paleontology. That study was realized to be no longer merely a descriptive and comparative practice, or the means, by which useful material was brought to the sciences of botany and zoology, but rather a branch of knowledge to be studied for its own intrinsic interest” [italics mine] (Zittel, 1901, p. 379). With the establishment of the Darwinian theory of evolution, paleontology acquired a central role. It could settle questions about the tempo and mode of evolution: for example, by proving or disproving whether evolution happened steadily, gradually, and by means of natural selection. According to German Darwinian paleontologist Melchior Neumayr (1845e1890), paleontology also obtained “a particular influence over the arrangement of the biological sciences, since Darwin’s Origin of Species put at the center the questions, whether the various plant and animal forms originated from a gradual reorganization, which powers caused this, and according to which laws they proceed.”2 (Neumayr, 1889, p. 26). This paper investigates German-speaking paleontologists’ reception of Darwin’s thought and the ways in which they negotiated their space of knowledge production accordingly. In Germanspeaking regions, the majority of paleontologists welcomed Darwin’s magnum opus, since it granted paleontology an independent voice within biology, and thus a new institutional setting. Around 1859, paleontology was indeed a complex, disunited, and disorganized discipline. On the one hand, due to the richness of the variety and distribution of invertebrate fossils, invertebrate paleontology was mainly used to draw a stratigraphic column of time.3 On the other hand, although vertebrate paleontology had adopted a morphological, i.e. a more biologically oriented attitude, ever since the publication of George Cuvier’s Recherches sur les ossemens fossiles de quadrupèdes (1812), it remained particularly difficult to find well-preserved records. Only in exceptional cases where specimens were entirely entombed in earth layers could paleontologists deal with vertebrate animals. In German-speaking countries, this scene changed through the positive reception of Darwin’s works. The Darwinian label provided the necessary epistemic cornerstone to emancipate paleontology from both geological interests as well as from the poor quality of its data. Darwin’s theory modified the social, institutional, and epistemic status of paleontology. It established paleontology as an autonomous biological discipline and allowed for the foundation of paleontological institutes detached from geology (for instance in Munich and Vienna). Austrian paleontologist and geologist Rudolf Hoernes (1850e1912) concisely summarized the development of late nineteenth-century paleontology4 when he maintained that “paleontology received a completely new meaning through the theory of evolution” (R. Hoernes, 1911, p. 312). In this paper I argue that German-speaking paleontologists, struggling to assert autonomy, opportunistically applied the Darwinian label to their disciplinedhence accepting Darwin’s theory of evolutiondbecause it established a new biological space for their discipline. This strategy was successful on the one hand: Zittel, for example, gained the first professorship for Paleontology in Germany at Munich in 1866, while the University of Vienna

2

All translations are mine unless otherwise noted. 3 See (Oppel, 1856e1858; Quenstedt, 1858). 4 In 1911, Rudolf Hoernes was expressing his last words on the importance of Darwin’s thought for paleontology, thus concluding a twenty-year controversy with his Austrian colleague Theodor Fuchs (1842e1925). See (R. Hoernes, 1880, 1884) and particularly (Tamborini, 2015b).

created a professorship for Paleontology for Neumayr in 1873. However, in the process of negotiating the features of paleontology within the Darwinian framework, German paleontologists constrained their practices too narrowly, for fear of leaving open possible results at odds with the burgeoning Darwinian biological community. In so doing, they also limited the further development of German-speaking paleontology. In other words, Zittel and Neumayr, and following them the majority of paleontologists, won key battles in establishing paleontology as an independent biological science. They applied the Darwinian label to paleontology in order to accomplish their aim and to be accepted by their fellow biologists. Their approach garnered them professorships and international recognitions. These victories, though, inflicted such a devastating toll on the discipline that their strategy ultimately was rejected. In fact, the course they set with their investigations resulted in a paleontological practice so limited in scope and methodology that the discipline lost both its biological space of knowledge and its significance. Following Plutarch, I call this a Pyrrhic victory. As a result, the analysis of German reception of Darwin offers an excellent case study to see how scientists’ labeling strategies shape the development of their discipline. Labeling can be both a boon and a detriment: It might allow for new kinds of practices, works, and financial support, and yet also limit the spread of research agendas: Under which conditions does labeling open up, establish, or restrict research agendas?

1. Paleozoology: the fossil record as valuable biological data The translator5 of the first two editions of the Origin of Species was the prominent Heidelberg paleontologist Heinrich Georg Bronn (1800e1862). Bronn’s translationsdthe first of which appeared in 1860dbrought immediate attention to Darwin among German paleontologists and biologists. However, Bronn also rejected some of the Origin’s key conclusions (Gliboff, 2008). This was mainly due to Bronn’s practice and results: He was an advocate of a quantitative method that could not be easily assimilated with Darwin’s theory. Indeed, those paleontologists who practiced a mathematical approach to datadas, for instance, Joachim Barrande (1799e1833) or Theodor Fuchs (1842e1925)dchallenged the evolutionary mechanisms proposed by Charles Darwin. They doubted whether evolution was the slow, gradual, and steady phenomenon produced means of natural selection proposed by Darwin, leading them to consider alternative mechanisms: Fuchs asserted that evolution was not characterized by “a continuous and uniformly progressive change”, but rather by a long time span of “relative calm with shorter periods of transformation [relativer Ruhe mit kürzeren Epochen der Umwandlung]” (Fuchs, 1879, p. 39)6 (Sepkoski & Tamborini, forthcoming; Tamborini, 2015b). Several influential German paleontologists at the height of their fame at the end of the 1850s shared Bronn’s ambiguous reception of Darwin’s theory. For instance, geologist and paleontologist Carl Albert Oppel (1831e1865) equally manifested sympathy and strong repulsion toward Darwin. Austrian geologist Eduard Suess (1831e1914) for his part, though an admirer of Darwin,7 seldom recognized his importance publicly.

5 On Darwin the German translations of the Origin of Species see (Junker & Backenköhler, 1999). 6 See also (Barrande, 1872). 7 “After Darwin’s book was published, views in all areas of biology drastically changed. In fact, besides the great discoveries of Galileo and Copernicus there is no other example of such a profound influence on the general ideas of the naturalists.” (Suess, 1901, p. 2).

Please cite this article in press as: Tamborini, M., The reception of darwin in late nineteenth-century German paleontology as a case of pyrrhic victory, Studies in History and Philosophy of Biological and Biomedical Sciences (2017), https://doi.org/10.1016/j.shpsc.2017.09.004

M. Tamborini / Studies in History and Philosophy of Biological and Biomedical Sciences xxx (2017) 1e9

Several students, who started their career by exhaustively reading the Origin of Species, transformed the ambiguous reaction of their teachers into an enthusiastic and positive acceptance. They carefully studied Darwin’s book, took up its central problem, and sought to disentangle its main difficulty: the imperfect and incomplete nature of the fossil record.8 In fact, as Melchior Neumayr observed: If someone follows and checks the biological literature in this period, he will find almost in every field a noticeable difference in the views and way of expression between those who, as formed scientific individualities, stood at the time of the publication of Darwin’s work and the younger generation, who started under the influence of Darwinian doctrine and accepted it from the beginning (Neumayr, 1889, p. 3). Therefore, German paleontologists neither immediately nor entirely accepted Darwin’s thought.9 There was rather a generational fracture10 between the already established paleontologists and the younger generation of naturalists, who were keen on discussing and accepting Darwin’s results as these emphasized the biological importance and institutional independence of paleontology. The names Karl Alfred von Zittel and Melchior Neumayr stand out among the young scientists who took up this challenge. These paleontologists took advantage of the new Darwinian framework as well as of German polycentrism,11 which celebrated regional and borderland identities instead of the idea of nation, thus elevating paleontology towards an autonomous biological discipline. Despite their common studies and supervisors,12 Zittel’s practice immediately gained national success and international echo. He

8 Indeed, as Darwin famously wrote, “I look at the natural geological record, as history of the world imperfectly kept, and written in a changing dialect; of this history we possess the last volume alone, relating only to two or three countries. Of this volume, only here and there a short chapter has been preserved, and of each page, only here and there a few lines.” (Darwin, 1966, p. 310). 9 As the Austrian paleontologist Othenio Abel put it, “One would have expected a rapid turn carried by the publication of Darwin’s major work in 1859 and that paleontology would rise again to a morphologically-biological science.” (Abel, 1907, p. 69) Also, according to Austrian paleontologist Karl Diener (1862e1928), “paleontology came relatively late onto the scene to take its rightful place in the development of the theory of evolution alongside the older biological sciences.” (Diener, 1910, p. 53). 10 William M. Montgomery also recognized the generational gap among paleontologists in the acceptance of Darwin’s theory. I disagree, however, with his assertion that “the older men were imbued with catastrophist ideas and found it difficult to adjust to the idea of gradual species changes” since Heinrich G. Bronn had already dispelled the catastrophist school of thought (Montgomery, 2008, p. 89). 11 On the notion of German polycentrism see (Penny, 2001, 2012). On German polycentrism and early twentieth-century natural history see (Tamborini, 2016). On the polycentric map of mid-nineteenth-century German paleontology see (Zittel, 1895a). 12 A particular fil rouge unified their carriers. Zittel studied under Bronn and Suess, Waagen under Suess and Zittel, and Neumayr under Oppel, Suess, Waagen, and Zittel. 13 Zittel studied medicine and zoology under Heinrich G. Bronn at the University of Heidelberg. He obtained his doctorate in 1860 from Heidelberg and subsequently spent a year in Paris, going on to obtain his venia legendi in 1862 from the University of Vienna. In the Viennese period, he studied under Wilhelm Karl Ritter von Haidinger (1795e1871)da talented mineralogist whose publications are characterized by extended and detailed descriptions of the inorganic worlddand the famous Austrian geologist and paleontologist Eduard Suess. Having taught mineralogy at Karlsruhe for three years (1863e1866), he took the chair for geology and paleontology at the university of Munich, previously occupied by Carl Albert Oppel. Zittel was hired and in those years Munich became one of the most foremost centers for paleontology in the world. Many important paleontologists studied under Zittel. For instance, Mario Canavari (1855e1928), Otto Jaekel (1863e1929), Josef Felix Pompeckj (1867e1930), Karl Heinrich Ernst Freiherr Stromer von Reichenbach (1871e 1952), Wilhelm von Branca (1895e1928), and Henry Fairfield Osborn. See (Tamborini, 2015a, 2016).

3

was able to address the biological nature of paleontology and to entice several important paleontologists to Munich. The American paleontologist Henry Fairfield Osborn (1857e1935) even asserted that “Munich became the Mecca of paleontologists, young and old” under Zittel’s supervision (Osborn, 1904, p. 186). Zittel was one of the first to recognize and promote the biological value of the fossil record in light of the theory of evolution.13 As he put it, Darwin’s theory “indicated in advance to paleontological research the direction in which the most successful discoveries can be done; it forced paleontology into a solid method” (Zittel, 1876). Paleontologists were able to accomplish exceptional discoveries as soon as they embraced Darwin’s theory. Darwin indeed marked “a turning point equally essential as the one associated with the names of Werner, William Smith, [Alexandre] Brongniart, and Cuvier” (Zittel, 1876, p. 41). Zittel’s view was that, in light of Darwin’s theory, paleontology needed to become paleozoology: a purely zoological discipline.14 To accomplish this transformation, paleontologists started gathering “numerous and significant proofs for the benefit of the theory of descendent” (Zittel, 1876, p. 42). These had the form of morphological data, which could be used to illustrate phylogenic relationships. Zittel and others decided that the chief role for paleontology within biology was to document the morphological succession of forms predicted by Darwin’s theory, and not to advance its own theoretical interpretations (which might have conflicted with Darwin’s view) as, for example, was popular among paleontologists in North America (see Bowler, this volume). In other words, what Zittel advocated for was a handmaid’s role for paleontology which increased biologists’ dependence on paleontologists for empirical evidence, but which limited paleontologists’ theoretical autonomy. Hence, Darwin’s theory emancipated paleontology from the domain of geology, in effect assuming a pivotal role among biological sciences. As Zittel triumphantly announced at the sixth international geological congress in Zurich in 1894, Paleontology has long since desisted from putting itself into the service of geology as the science of index fossils. Gradually, it has grown into an autonomous branch of the biological sciences and takes part in all of its trends and movements. The most powerful revolution has caused the final victory of the descent theory in the descriptive natural sciences and it has completely influenced and reformed their research method (Zittel, 1894, p. 125). Hence, Zitteldand following him the majority of German paleontologists15dwelcomed the new cornerstone offered by the theory of evolution, for it finally promoted and established their discipline as a biological science. The collection of fossils assumed a much broader function beyond the mere depiction of a stratigraphic column of time; this work could be useful to gather data that prove Darwin’s theory. Indeed, according to Zittel, paleontology functioned as corroboration. In a letter to Ernst Haeckel in 1868, after having praised his Natürliche Schöpfungsgeschichte, Zittel asserted that “the phylogenetic trees [presented in Haeckel’s book]

14 On the explicit reference to Cuvier’s definition of paleontology see (Rudwick, 2008, pp. 47e48). 15 For example, Otto Jaekel, Josef Felix Pompeckj, Karl Heinrich Ernst Freiherr Stromer von Reichenbach, Melchior Neumayr, Friedrich Rolle, and Rudolf Hoernes followed Zittel’s lead. Also Karl Diener remarked that “it is probably beyond any doubt that paleontology [.] provides us with a time-ordered material for reconstructing the phylogeny of organisms. It deserves a major say in deciding on the relative value of theories tied with the problem of evolution, Darwin himself has openly admitted that his theory of natural selection cannot exist, if it is in accordance with the fossil documentation used to decipher the history of life.” (Diener, 1910, p. 29).

Please cite this article in press as: Tamborini, M., The reception of darwin in late nineteenth-century German paleontology as a case of pyrrhic victory, Studies in History and Philosophy of Biological and Biomedical Sciences (2017), https://doi.org/10.1016/j.shpsc.2017.09.004

4

M. Tamborini / Studies in History and Philosophy of Biological and Biomedical Sciences xxx (2017) 1e9

will definitely be contested and, as I fear, they will possibly reduce the effect of your work” (Zittel, 11.11.1868, p. 1). This may happen since, as Zittel stated in his Grundzüge der Palaeontologie, “phylogenetic trees can only be considered well justified if they are validated by paleontological facts” (Zittel, 1895b, p. 11). The irony, though, is that German-speaking paleontologists positively accepted Darwin not because they were convinced by the evolutionary mechanism he proposed (natural selection). Indeed, as soon as paleontology was recognized as an autonomous biological discipline, several German paleontologists rejected the Darwinian framework. For instance, paleontologist Edgar Dacqué (1878e1945) embraced orthogenetic and neo-Lamarckian theories which were easily reconcilable with a literal reading of the fossil record. German-speaking paleontologists welcomed therefore Darwin’s thought berceuse this called attention to the role of phylogenetic investigations and thus to the richness and value of paleontological data as means for corroborating evolutionary narration: “Darwin has for the first time highlighted that systematics needed to undertake a drastic change according to the new aspects opened by the theory of evolution and that this dry field would obtain an unexpected interest, if it gave expression to relationships of descent though genetic16 basis; that is valid for paleontology as well.” (Neumayr, 1874, p. 144) Thus, by embracing Darwin’s theory paleontologists were able to stake out a rich, stable, and innovative field of research. For instance, in the introduction to the first edition of the Handbuch der Palaeontologie (Palaeozoologie) (1876) Zittel wrote that paleontology aimed at the “realization of a natural systematics in accordance with morphological and phylogenetic experience”. Echoing Zittel over a decade later, German paleontologist Rudolf Hoernes asserted that “[z] oopaleontology is an independent science [.] which aims to clarify the evolutionary relationship between recent and fossil forms by examining the latter [through] comparative anatomical and embryological investigation” (R. Hoernes, 1884, p. 6). To convince both his students and the scientific community that fossils were positive evolutionary evidence and not merely imperfect data that only incompletely represent extinct organisms, Zittel needed a medium whereby he might systematically order, represent, and disseminate this notion. He came up with the Handbuch der Palaeotontologie (Palaeozoologie) and the Grundzüge der Palaeontologie,17 because textbooks and manuals could systematically create and easily broadcast such data among the scientific community.18 Both textbooks endeavored to provide a general survey of paleontological subject matter in harmony with the modern standpoint of zoology [.] throughout the entire work the primary object has been to point out the close relationships between paleontology and the other branches of biological science (zoology, comparative anatomy, botany, embryology) and to make application to paleontology of the data acquired by those sciences (Zittel, 1876, p. 380). Through the study of Handbuch and Grundzüge der Palaeontologie the student was immediately immersed in the complexity and the richness of paleozoological data: these books are in-depth systematics manuals for the major phyla that have appeared in the history of life. All the major fossilized phyla were thoroughly

16

With the term “genetic” Neumayr meant “phylogenetic”. Zittel was not the only one who decided to write a textbook to consolidate his ideas; several of Zittel’s students and other Darwinian paleontologists took up this task as well. See for example (R. Hoernes, 1884; Neumayr, 1889; Stromer, 1909). 18 On the importance of textbook in the process of knowledge production and communication see (Lundgren & BensaudeeVincent, 2000; Simon, 2011; Vicedo, 2012a, 2012b). 17

described and meticulously illustrated according to their “morphological features and their historical development d in short according to their natural affinity” (Zittel, 1876, p.43). Hence, the paleozoological practice was based upon comparative and morphological work: As the Austrian paleontologist Othenio Abel put it, “the most important task of the paleozoologist is the proper detection of the anatomical structure of the fossil animals. It follows that the paleozoologist must be, first and foremost, a comparative anatomist.” (Abel, 1907, p. 70). Yet, although fossils could potentially be treated as valuable biological data that aid biologists in the discovery of evolutionary relationships, they were still incomplete and imperfect entities, since not all organisms of former epochs have been completely fossilized. This might have undermined the epistemic status of paleontology and its reliability.19 Zittel escaped this impasse by relying again on Darwin’s theory. He asserted that the Origin of Species gave paleontology “a hint on both how the still missing forms should approximately look like and where the paleontologists have to look for them” (Zittel, 1876, p. 42). As a result, the paleontologist could work on his findings and integrate the missing parts. He could create a complete morphological structure of the extinct organisms. Zittel took up this practice since, as he claimed, it had already been successfully adopted in other disciplines: “in chemistry new connections are often constructed through purely theoretical ways. Likewise, paleontology is capable of presupposing the existence of extinct forms and supposing in all probability their organization.” (Zittel, 1876, p. 42) The paleozoological practice was thus based on classificatory work on fossil remains and theoretical efforts to construct and hypothesize their organization even if these remains were only partially fossilized. In this way, the paleontologist “gained an accurate and safe knowledge of the fossils concerning their organizational relations, their mutual relationships, and affinity to the living forms” (Zittel, 1876).20 However, by putting a premium on “accurate and safe knowledge” for fear of losing the recently acquired independent biological space, Zittel’s practice carefully restrained from investigating and illustrating possible evolutionary relationships as well as their causes. He emphasized only the search of forms and the importance of fossils as genuine morphological data, avoiding any discussion about the nature of the evolutionary mechanisms. This would be the main reason behind Zittel’s Pyrrhic victory and would leave a mark on the future generations of German paleontologists that would persist for a century or more. For instance, in describing Ulrich Lehmann’s Fossils Invertebrate (1983), British paleontologist and author of the important textbook Principles of Paleoecology (1963) Derek Ager (1923e1993) expressed this sharp judgment about German scholarship: “this is a work of German scholarship; in fact it out-Zittels Zittel, though at much shorter length: [.] apart from a brief section on the origin of life, there is very little on evolution, or paleoecology, or functional morphology, or geographical distribution or any of the dozen other subjects that many of us find so fascinating. No doubt that makes it the more scientific in a purely descriptive sense; it alsodregrettablyd makes

19 This was indeed a crucial point, for several paleontologists gave a new ontological status to the allegedly incomplete and imperfect fossils so that these could be employed in a statistical framework. The quantitative approach used by those paleontologists generated, in turn, results openly in contrast with the Darwinian theory of evolution. See (Tamborini, 2015b). 20 This point clearly indicates the difference between Zittel’s approach and the late idealistic morphology school. Goethe only implicitly inspired Zittel’s practice: Zittel took up a morphological method only because it guaranteed an autonomous space of knowledge for paleontology; idealist morphologists, in contrast, explicitly appealed to Goethe, thus seeing their research entirely within the Goethian research program. See (G. S. H. Levit, Uwe; Olsson, Lennart, 2014).

Please cite this article in press as: Tamborini, M., The reception of darwin in late nineteenth-century German paleontology as a case of pyrrhic victory, Studies in History and Philosophy of Biological and Biomedical Sciences (2017), https://doi.org/10.1016/j.shpsc.2017.09.004

M. Tamborini / Studies in History and Philosophy of Biological and Biomedical Sciences xxx (2017) 1e9

it extremely dull.” (Ager, 1983, p. 470) Hence, Zittel set important rules to conceive paleontology as a biological science, following and embracing Darwin’s thought. However, these did not enable paleontologists to go beyond a mere collection and classification of morphological data.

2. Mode and cause of evolution Not all German-speaking paleontologists, however, accepted Zittel’s resistance to contribute to causal evolutionary explanations. For instance, German paleontologist Melchior Neumayr21 offers an outstanding case study to show how German-speaking paleontologists modeled their practice on the Darwinian framework. Unlike Zittel, he sought to use the fossil record to contribute to evolutionary theory. However, as I will show, he reached the same conclusion as his prominent colleague. Indeed, Neumayr won important battles in order to establish paleontology as an independent biological science, but these ultimately lost him the war. In 1889, Neumayr published the first part of his unfinished textbook, Die Stämme des Thierreiches (1889), in which he dealt with invertebrate fossils.22 These were allegedly the source of information for stratigraphic and geological studies. Nevertheless, in accordance with Darwin’s theory, he explicitly asserted that these records could be used “to demonstrate that all animals and plants evolved from one or a few basic forms by gradual transformation and to seek the causes which have led to this procedure.” (Neumayr, 1889, p. 2) As a result, “paleontology investigated the plants and animals, which had lived before the present time on Earth. In the investigation of animal remains, [.] paleontology follows the same principles used in zoology and it has an intimate connection with this science, so that a sharp distinction between the two areas is hardly possible.” (Neumayr, 1889). Morphological data were thought to be useful only if coupled with evolutionary investigations that show “the direct analytical proof of the variability of species” (Neumayr, 1889). This implied an “inductive treatment” of these data in order to illustrate evolutionary patterns. This treatment was possible by unifying in series the collected data so that potential patterns of development could emerge. This unification generated a series of forms which could prove Darwin’s theory, as paleontologists Moritz Hoernes (1815e 1868), Franz Hilgendorf (1839e1904), and Wilhelm Waagen (1841e 1900) had previously done (Hilgendorf, 1863; M.; Hoernes, 1856; Reif, 1983a; Waagen, 1869). Therefore, Neumayr centered his practice on the identification of historical trends in order to illustrate the mode of evolution. He unified into a straight line of development all the fossilized organisms, which were spread over different earth layers despite belonging to the same family or genus (see, for instance, Fig. 1). He used this and similar visualizations as a necessary basis “for further conclusions of greater importance” (Neumayr & Paul, 1875, p. 16) about Darwinian evolution. In fact, the series of forms could prove whether evolution occurred gradually and constantly, as Darwin proposed. By carefully investigating the morphological differences

21 Melchior Neumayr was born in Munich and studied natural sciences in the same city under Albert Oppel and Wilhelm von Gümbel (1823e1898). He studied afterwards at Heidelberg, where he obtained his doctorate and venia legendi. He moved then to Vienna anddafter working at the Australian Reichsanstalt under Suessdtook the professorship of Paleontology (the first chair in Austria, as above) at the University of Vienna. 22 Wolf-Ernst Reif described “the book’s 150-page introduction [as] probably one of the best treatises on Darwinism to be found anywhere in the literature of the nineteenth century’ (Reif, 1986, p. 89; 1983b). As Austrian paleontologist Franz Toula (1845e1920) asserted, “Neumay’s results in this field belong to the most solid supports for the theory of evolution” (Toula, 1890, p. 316).

5

of numerous fossilized aquatic gastropod mollusks, for instance, Neumayr observed in 1875 that Vivipara neumayri (gastropod mollusks) did pass gradually through intermediate forms into Vivipara hoernesi (see Fig. 1). Hence, by representing evolution as series of fossilized forms, paleontologists were able to reveal “the mode of derivation of the newer from the older types” and therefore to demonstrate “that beyond all controversy the species with highly complicated ornamentation were variously derived by descent from the simple and unornamented Vivipare achationides” (ibid.). Secondly, this kind of visualization broadened the comprehension of the mechanisms of evolution. For instance, a careful analysis of the aligned forms proved that Moritz Wagner’s migrations theory was false: “in many cases, we can observe the gradual change of a phylum through several mutations within a single profile. This implies that a change happened without any kind of isolation of individuals taking place in accordance with the conditions of migration theory” (Neumayr & Paul, 1875, p. 100). Furthermore, by bringing into line the different forms found in several earth layers Neumayr indicated the directionality of morphological differences. He claimed that the series of forms demonstrated a particular and inevitable constancy in the mode and direction of its changes. From the first modification onwards, all the changes proceeded according to one direction. This directionality excluded the hypothesis that evolution could proceed backwards: “There is no single case in which series of forms returns to its roots; there are though socalled backwards lines [.] however, there is never an exact return to the starting points, but rather the result is always a new, unprecedented form of quite independent features” (Neumayr, 1889, p. 61). Based on the series of forms, Neumayr was consequently making an important point, which would be asserted again by Belgian paleontologist Louis Dollo (1857e1931) in 1893: evolution is an irreversible phenomenon.23 By embracing Darwin’s theory as a biological cornerstone for their practice, Neumayr, Rudolf Hoernes, Franz Hilgendorf, Wilhelm Waagen used the fossil record to draw series of forms and thereby study the mode and cause of evolution. Paleontologists’ capacity to investigate the tempo, mode, and causes of evolution was therefore publicly recognized. For instance, Darwin himself acknowledged the standing of Neumayr’s practice and the importance of his 1875 work. On the 9th of March 1877 he wrote to Neumayr his apologies “for not having formerly insisted more strongly on this head in my Origin of Species, as most of the best facts have been observed since its publication” (Darwin, 1969, p. 232) and praised Hilgendorf’s series of forms in the fifth edition of On the Origin of Species (1869). Neumayr’s invertebrate paleontology shared with Zittel the conviction that there was no separation between vertebrate and invertebrate paleontology, if fossils were no longer considered geological time-markers. This entailed a closer relation between paleontology and zoology and its consequent detachment from geology. Furthermore, he too based his investigations on Darwin’s theory of evolution to obtain an autonomous domain for paleontological investigations: the identification of series of forms over geological time. However, Neumayr, and the other paleontologists who developed series of forms, shared with Zittel also the same Pyrrhic victory. As I show in fact, they narrowed the paleontological practice so much so that it eventually lost its importance and efficacy after its initial enthusiastic reception.

23 In his 1884 textbook Elemente der Palaeontologie (Palaezoologie), also Rudolf Hoernes emphasized the importance of the series of forms for paleontology. This textbook was translated in French by Louis Dollo in 1886. See (R. Hoernes, 1866).

Please cite this article in press as: Tamborini, M., The reception of darwin in late nineteenth-century German paleontology as a case of pyrrhic victory, Studies in History and Philosophy of Biological and Biomedical Sciences (2017), https://doi.org/10.1016/j.shpsc.2017.09.004

6

M. Tamborini / Studies in History and Philosophy of Biological and Biomedical Sciences xxx (2017) 1e9

Fig. 1. Neumayr’s pictographic representation of a series of forms that indicates the gradual mode of evolution (Neumayr & Paul, 1875; plate IV).

Please cite this article in press as: Tamborini, M., The reception of darwin in late nineteenth-century German paleontology as a case of pyrrhic victory, Studies in History and Philosophy of Biological and Biomedical Sciences (2017), https://doi.org/10.1016/j.shpsc.2017.09.004

M. Tamborini / Studies in History and Philosophy of Biological and Biomedical Sciences xxx (2017) 1e9

3. Pyrrhic victory German-speaking paleontologists positively received Darwin’s theory because it emphasized the key role of phylogenetic investigations. Zittel established paleontological data as morphological entities which might aid phylogenetic reconstruction. By defining and using such epistemic entities, Zittel provided a great number of morphological data and a plenty of evidence supporting the theory of evolution. This enterprise was taken up in Germany and abroad. Bavaria, seeing a good opportunity to represent its intellectual power and autonomy worldwide,24 provided financial stimulus, consequently allowing Zittel to attract numerous students from Germany and abroad to Munich. Many of them would develop Zittel’s idea between the end of the nineteenth and the beginning of the twentieth centuries: Several invertebrate and vertebrate paleontologists accepted and promoted this pure morphological practice (Tamborini, 2016). However, Zittel scored only a Pyrrhic victory. His practice had too high epistemic costs. It limited the tasks and aims of paleontology and plunged it into an identity crisis. In order to avoid any possible metaphysical speculations about the cause of evolution, which would have undermined the status of paleontology, Zittel did not promote a theoretical-deductive practice to understand, corroborate, and eventually develop the theory of evolution. To be sure, the declared aim of many paleontologists, who favorably embraced Zittel’s morphological conception of data, was to use these data to understand evolution. They were unable to pursue this goal, however, because Zittel and his contemporaries did not explicitly determine how and to what extent these facts should be read in order to come up with biological processes and patterns. So anxious that subjective statements might emerge and in effect draw criticisms to paleontology, they delimited every possible misunderstanding. Zittel, as well as several important paleontologists in his wake, did not open up the theoretical process about evolutionary mechanisms: “they failed to provide one coherent theory.” (Reif, 1986, p. 121). This point is reflected in Zittel’s textbooks. They were powerful media for promoting and teaching the qualities of paleozoological data; however, they presented all the ambiguity of this notion as well. These textbooks did not teach how paleozoological data could properly be used as data, i.e. as starting points for further investigations. As a reviewer in Nature noticed: the title adopted for the English edition “Text-Book of Paleontology” neither correctly translates the German title nor

24 The flourishing of paleontology in Germany was highly supported by both the universities and the regional states, guaranteeing thus an indispensable condition for its existence. This entailed a deep difference between the spread and the development of paleontological thought in German and in English speaking countries between the nineteenth and twentieth centuries. In fact, while in Germany the prominent paleontologists were occupying outstanding academic and social positions (it is extremely difficult to separate the names of Heinrich Bronn, Alfred Zittel, Melchior Neumayr, Albert Oppel, and Othenio Abel from the fortune of the universities in which they grew up and taught), in American universities and curricula vertebrate paleontology were regarded only peripherally. In An Agenda for Antiquity, Rainger asserted that “many colleges maintained their collections and around the turn of the century a number of the new college museums were established. Yet it is questionable to what extent the collections in those museums were used by students. Courses on biology and geology included material on fossil vertebrate, but few students chose or were encouraged to choose that subject as a field of specialization. At Yale, where over fifty students obtained Ph.D. degrees in geology between 1900 and 1925, only three specialized in vertebrate paleontology. At Columbia University where 144 Ph.D. degrees were granted in zoology in the years 1895e1942, fourteen were in vertebrate paleontology. The story was the same at Princeton, the University of Chicago, the University of California, at Berkeley, and other schools” (Rainger, 1991, p. 21).

7

expresses the character of the book. Consisting as it does of a mass of facts with no indication either of methods of work or of the general theory of the subject, it is by itself of little use for the beginner, and requires to be supplemented in some way, either by a teacher or by some, as yet unwritten, introduction. (Watson, 1946, p. 681) The same can be said for the allegedly updated English versions of Zittel’s Grundzüge: Romer’s Vertebrate Paleontology (1933) or Shrock’s Principles of Invertebrate Paleontology (1953). In these manuals, students found various different useful taxonomical lists and entries, but these textbooks applied the same approach as Zittel. Both Romer and Shrock did not identify how far the data may be read and whether the paleontologist may take a step forward to seek the reasons behind the diversity of the gathered data. Neumayr’s series of forms shared the same fortune with Zittel’s. Although Neumayr, as well as Hoernes and Waagen, sought to illustrate how far data may be read to discover evolutionary mechanisms, they set criteria too severe for this reading. As a result, it was almost impossible to carry out this practice and remain a Darwinian paleontologist. First, Neumayr and Hoernes took for granted and even emphasized the incomplete nature of the fossil record in order to undermine its possible quantitative reading (Tamborini, 2015b). Second, Neumayr proposed to trace series of forms based on a literal reading of the fossil record “where good enough fossils are available” (Neumayr, 1889, p. 53). However, wellpreserved fossils were extremely difficult to be found: As Neumayr himself tellingly admitted, “for about 10 years, I have been mainly occupied with the transmutation of species and in all this time I found only a single example in which the production of series of forms was so easy that it would have been recognized by other observers without any difficulty” (Neumayr, 1889). As a result, after an initial enthusiastic recognition, only a few paleontologists decided to embrace and further develop series of forms. Second, several of Zittel’s students adopted the main results of Neumayr’s practice (i.e. directionality of the series of forms and pivotal role of external conditions) to support non-Darwinian theories much in line with Neumayr’s literal reading of the fossil record. At the beginning of the twentieth century, professorships for paleontology were indeed already established and the priority of the German paleontological community was rather to obtain enough national funding than to fight for a kind of biological autonomy (Tamborini, 2016). Without the necessity of being a Darwinian paleontologist in order to promote the disciplinary status and autonomy of this discipline, they took up these results and put them into another biological agenda much easier to support. They reframed Zittel’s and Neumayr’s method and results within a neo-Lamarckian, an orthogenetic or an idealistic framework, paving the way for the so-called eclipse of Darwinism (Bowler, 1983). The German paleontologists Edgar Dacqué (1878e 1945) perfectly epitomized this epistemic twist. Although he studied and obtained his doctorate under Zittel (1903), he became one of the champions of idealistic morphology in the early twentieth century (Rieppel, 2016). Against his teacher, he asserted that the Darwinian principle of descent with modification was neither the all-encompassing basis for illustrating and explaining the organic world nor a solid cornerstone for the paleontological practice. He proposed conversely to read the series of forms as forms connected by an immanent principle which unfolded an ideal type (Dacqué, 1935, p. 2). In his Organische Morphologie und Paläontologie (1935), Dacqué emphasized this epistemic turn by asserting that the series of forms proved that “mutations are fundamentally always jump-mutations [Sprungmutationen]” (Dacqué, 1935, p. 85). As a result, evolution did not follow the

Please cite this article in press as: Tamborini, M., The reception of darwin in late nineteenth-century German paleontology as a case of pyrrhic victory, Studies in History and Philosophy of Biological and Biomedical Sciences (2017), https://doi.org/10.1016/j.shpsc.2017.09.004

8

M. Tamborini / Studies in History and Philosophy of Biological and Biomedical Sciences xxx (2017) 1e9

mechanisms proposed by Darwin. Dacqué concluded his campaign against the Darwinian framework, claiming that the more extinct genera were classified, the more it became evident that only a few types of fixed and ideal forms went through new adjustments and specializations: There was no proof of the gradual nature of evolution supported by Neumayer and a fortiori Darwin.25 The paleontological practices proposed by Zittel and Neumayr and the consequent narrow space of knowledge negotiated by them were mainly responsible for this epistemic shift. As Dacqué noticed, although paleontology obtained a biological status after Zittel’s and Neumayr’s reflections, its data endorsed a plurality of non-Darwinian theories, which cannot be bracketed out by using Zittel’s or Neumayr’s practice (Dacqué, 1928). As a main consequence, the biological Darwinian community ostracized paleontology from evolutionary biology. Indeed, during the last years of Zittel’s life, German paleontologists entered into a deep identity crisis as to their aims, practices, and role in society. This crisis culminated after Zittel’s death in 1904. As paleontologist Otto Jaekel, a former student of Zittel, remarked, Although excellent workers, such as Karl von Zittel in Munich, have prepared the ground for detailed paleontological studies and have wrested the success of German paleontology from foreign countries, this has remained well behind these countries in terms of its [financial] support. At that time we at least had a place in Germany that due to Zittel’s pure paleontological works was for the benefit to this science. Yet, since this place is now returned to geology, it seems that German paleontology has found on the part of the government a happy death d it ekes out a miserable existence as auxiliary disciplinary of geology. [italics mine] (Jaekel, 1910, p. 35) Hence, according to Jaekel, paleontology was dramatically returning to be a mere auxiliary science of geology.26 He vividly emphasized that the last sixty years of disciplinary emancipation had been quickly nullified. Zittel’s revolutionary move was in vain and his legacy dissipated: “this is a dishonorable situation of German Empire, not only because paleontology had been so fervently carried out [in the past] or because other countries have been incomparably better promoting it for many years, but especially because [paleontology] has a fundamental importance for estimating the organic remains on our earth.” (Jaekel, 1910, p. 35). To conclude, German paleontologists positively accepted Darwin during the second half of the nineteenth century because his theory guaranteed an autonomous space of knowledge for their discipline. Being a Darwinian paleontologist thus entailed taking part in the making of a scientific discipline, paleontology, which tried to go beyond a plurality of methods, and aiming towards a coherent and independent biological discipline. However, German paleontologists were not able to set the conditions under which knowledge could be gained and disseminated.27 Hence, through the analysis of German paleontologists’ practices at the end of the nineteenth century, this paper shows the importance of scientists’ ability to work, polish, and refine what they are willing to cede and concede in order to be recognized by the broader scientific community. In order to gain acceptance within the biological

25 On Dacqué see (G. S. M. Levit, 2006). Broadly on the anti-Darwinian evolutionary theories around 1900 (Bowler, 1983). 26 On Jaekel’s place in twentieth-century paleontology and German system of science see also (Nieuwland, 2017). 27 Indeed, as Nieuwland also noticed, “it was not until the first decade of the twentieth century that a more integrated view of the study of fossil life, that took in factors such as ecology, was considered by people such as Otto Jaekel and Othenio Abel” [italics mine] (Nieuwland, 2017, p. 17).

community, Neumayr and Zittel, and following them the majority of German-speaking paleontologists, too narrowly limited the paleontological aim and practices. They were able to set their idea of science in the short term, but this eventually collapsed over the long term. References Abel, O. (1907). Die Aufgaben und Ziele der Paläozoologie. Verhandlungen der ZoologischeBotanischen Gesellschaft, 57, 67e78. Ager, D. V. (1963). Principles of paleoecology: An introduction to the study of how and where animals and plants lived in the past. New York: McGraw-Hill. Ager, D. V. (1983). Palaeontology by the telephone book. Nature, 304, 470. Barrande, J. (1872). Systême silurien du centre de la Bohême. Prague and Paris. Bowler, P. J. (1983). The eclipse of darwinism: Anti-darwinian evolutionary theories in the decades around 1900. Baltimore; London: Johns Hopkins University Press. Breidbach, O. (2003). Post-Haeckelian comparative biology d adolf Naef’s idealistic morphology. Theory in Biosciences, 122, 174e193. Coen, D. R. (2007). Vienna in the age of uncertainty. Science, liberalism, and private life. Chicago: Chicago University Press. Craw, R. (1992). Margins of cladistics: Identity, difference and place in the emergence of phylogenetic systematics. In P. Griffiths (Ed.), Trees of life. Dordrecht; London: Kluwer Academic Publishers. Dacqué, E. (1928). Das fossile Lebewesen. Eine Einführung in die Versteinerungskunde. Berllin, Heidelberg: Springer. Dacqué, E. (1935). Organische Morphologie und Paläontologie. Berlin: Gebrüder Borntraeger. Darwin, C. (1966). On the origin of species. Cambridge: Cambridge University Press. Darwin, C. (1969). Life and letters (vol. II). New York: Johnson Di Gregorio, M. A. (2005). From here to eternity: Ernst Haeckel and scientific faith. Göttingen: Vandenhoeck & Ruprecht. Di Gregorio, M. A. (2008). Under Darwin’s banner: Ernst Haeckel, Carl gegenbauer and evolutionary morphology. In E. M. G. Engels, & F. Thomas (Eds.), The reception of Charles Darwin in europe volume I. London: Continuum. Diener, C. (1910). Paläontologie und Abstammungslehre. Leipzig: Göschen. Fuchs, T. (1879). Ueber die praesumirte Unvollstaendigkeit der palaeontologischen Ueberlieferung. Verhandlungen der Kaiserlich-Königlichen Geologischen Reichsanstalt, 16. Gliboff, S. (2008). H.G. Bronn, Ernst Haeckel, and the origins of German darwinism. A Study in Translation and Transformation. Cambridge: The MIT Press. Hilgendorf, F. (1863). Beiträige zur Kenntnis des Süßwasserkalkes von Steinheim. University of Tübingen. Unpublished Ph.D. Thesis. Hoernes, M. (1856). Die fossilen Mollusken des Tertiär-Beckens von Wien. Vienna: Geologische Reichsanstalt. Hoernes, R. (1866). Manuel de paléontologie. Translated by L. Dollo. Paris: Savy. Hoernes, R. (1880). Die Unvollständigkeit der paläontologischen Ueberlieferung. Verhandlungen der Kaiserlich-Königlichen Geologischen Reichsanstalt, 2, 17e20. Hoernes, R. (1884). Elemente der Palaeontologie (Paleozoologie). Leipzig: Veit & Comp. Hoernes, R. (1911). Die Bedeutung der Palaeontologie fuer die Erdgeschichte. Scientia: Rivista Internazionale di Sintesi Scientifica, 10, 307e325. Jaekel, O. (1910). Über Geologie und Paläontologie an den deutschen Hochschulen. Naturwissenschaftliche Wochenschrift, IX, 33e35. Junker, T., & Backenköhler, D. (1999). ‘Vermittler dieses allgemeinen geistigen Handels’: Darwins deutsche Verleger und Übersetzer bis 1882. Verhandlungen zur Geschichte und Theorie der Biologie (3). In A. Geus, T. Junker, H.J. Rheinberger, C. Riedl-Dorn, & M. Weingarten (Eds.), Repräsentationsformen in den biologischen Wissenschaften. Berlin: Verlag für Wissenschaft und Bildung Levit, G. S. M., & Kay. (2006). The history of essentialism vs. Ernst Mayr’s "Essentialism Story": A case study of German idealistic morphology. Theory in Biosciences, 124, 281e307. Levit, G. S. H., Uwe, Olsson, & Lennart. (2014). The darwinian revolution in Germany: From evolutionary morphology to the modern synthesis. Endeavour, 30. Lundgren, A., & BensaudeeVincent, B. (Eds.). (2000). Communicating Chemistry: Textbooks and Their Audiences, 1789e1939. Canton: Watson Publishing International. Montgomery, W. M.. (2008). Germany. In T. F. Glick (Ed.), The comparative reception of Darwinism. Chicago: University of Chicago Press. Neumayr, M. (1874). Die Fauna der Schichten mit Aspidoceras acanthicum. Abhandlungen der Geologischen Bundesanstalt in Wien, 5, 141e257. Neumayr, M. (1889). Die Stämme des Thierreiches. Vienna; Prague: Tempsky. Neumayr, M., & Paul, C. M. (1875). Die Congeriene und Paludinenschichten Slavoniens und deren Faunen: Ein Beitrag zur DescendenzeTheorie. Abhandlungen der Kaiserlich-Königlichen geologischen Reichsanstalt, 7. Nieuwland, I. (2017). The colossal stranger. A cultural history of diplodocus carnegii, 1902-1913. University of Groningen (PhD thesis) Nyhart, L. K. (1987). The disciplinary breakdown of German morphology, 1870e 1900. Isis, 78, 365e389. Nyhart, L. K. (1995). Biology takes form. Animal morphology and the German universities 1800 e 1900. Chicago: University of Chicago Press. Oppel, A. (1856e1858). Die Juraformation Englands, Frankreichs und des südwestlichen Deutschlands, nach ihren einzelnen Gliedern eingetheilt und verglichen.

Please cite this article in press as: Tamborini, M., The reception of darwin in late nineteenth-century German paleontology as a case of pyrrhic victory, Studies in History and Philosophy of Biological and Biomedical Sciences (2017), https://doi.org/10.1016/j.shpsc.2017.09.004

M. Tamborini / Studies in History and Philosophy of Biological and Biomedical Sciences xxx (2017) 1e9 Jahreshefte des Vereins für vaterländische Naturkunde in Württemberg,; 12, 1856, 121-556, 13, 1857, 141-396, 14, 1858, 129-291. Osborn, H. F. (1904). Karl Alfred von Zittel. Science, 19, 186e188. Penny, G. H. (2001). Objects of Culture: Ethnology and ethnographic museums in imperial Germany. Chapel Hill: University of North Carolina Press. Penny, G. H. (2012). Reflections. German polycentrism and the writing of history. German History, 30, 265e282. Quenstedt, F. A. (1858). Der jura. Tübingen: Laupp. Rainger, R. (1991). An agenda for Antiquity: Henry Fairfield Osborn & vertebrate paleontology at the american museum of natural history, 1890e1935. Tuscaloosa: University of Alabama Press. Reif, W. E. (1983a). Hilgendorf’s (1863) dissertation on the steinheim planorbids (gastropoda; Miocene): The development of a phylogenetic research program for paleontology. Paläontologische Zeitschrift, 57, 7e20. Reif, W. E. (1983b). Evolutionary theory in German paleontology. In M. Grene (Ed.), Dimensions of Darwinism: Themes and counterthemes in twentiethecentury evolutionary theory. Cambridge: Cambridge University Press. Reif, W. E. (1986). The search for a macroevolutionary theory in German paleontology. Journal of the History of Biology, 19, 79e130. Richards, R. J. (2008). The tragic sense of life. Ernst Haeckel and the struggle over evolutionary thought. Chicago: University of Chicago Press. Rieppel, O. (2016). Phylogenetic Systematics: Haeckel to Hennig. Boca Raton: CRC Press, Taylor & Francis. Rudwick, M. J. S. (2008). Worlds before Adam: The reconstruction of geohistory in the age of reform. Chicago: University of Chicago Press. Sepkoski, D. & Tamborini M. (forthcoming). “An image of Science”: Cameralism, statistics, and the visual language of natural history in the nineteenth century, Historical Studies in the Natural Sciences. Simon, J. (2011). Communicating Physics: The production, circulation and appropriation of Ganot’s textbooks in France and england, 1851e1887. London: Pickering & Chatto. Stromer, K. H. E. F. . v. R. (1909). Lehrbuch der Paläozoologie. Leipzig und Berlin: Teubner.

9

Suess, E. (1901). Abschiedsvorlesung, gehalten bei seinem Rücktritt vom Lehramte am 13. Juli 1901. Beitr. Paläont. U. Geol. Österreicheungarns U. D. Orients, 16, 1e8. Tamborini, M. (2015a). Die Wurzeln der ideographischen Paläontologie: Karl Alfred von Zittels Praxis und sein Begriff des Fossils. NTM Zeitschrift für Geschichte der Wissenschaften, Technik und Medizin, 23, 117e142. Tamborini, M. (2015b). Paleontology and Darwin’s theory of Evolution: The subversive role of statistics at the end of the 19th century. Journal of the History of Biology, 48, 575e612. Tamborini, M. (2016). “If the americans can do it, so can we”: How dinosaur bones shaped German paleontology. History of Science, 54, 225e256. Toula, F. (1890). Zur Erinnerung an melchior Neumayr. Nachruf, gehalten den 12. Februar 1890. In Verein zur Verbreitung Naturwissenschaftlicher Kenntnisse (Vorträge,. Wien) Vicedo, M. (2012a). Introduction: The secret lives of textbooks. Isis, 103, 83e87. Vicedo, M. (2012b). Playing the game psychology textbooks speak out about love. Isis, 133. Waagen, W. H. (1869). Die Formenreihe des Ammonites subradiatus: Versuch einer paläontologischen Monographie. Geognostisch-Paläontologische Beiträge, 2, 179e256. Watson, D. M. S. (1946). Vertebrate paleontology. Nature, 157, 681e681. Zittel, K. A. v. (01.11.1868). Zittel an Ernst Haeckel. In: Archiv des Ernst-HaeckelHauses in Jena. 13028 Zittel, K. A. . v. (1876). Handbuch der Palaeontologie (Palaeozoologie). Munich; Leipzig: Rudolph Oldenbourg. Zittel, K. A. . v. (1895a). Geologie an der deutschen Universitäten. Zeitschrift für praktische Geologie, 13e16. Zittel, K. A. . v. (1895b). Grundzüge der Palaeontologie (Palaeozoologie). München: Rudolph Oldenbourg. Zittel, K. A. . v (1901). History of geology and palaeontology. To the end of the nineteenth century. London: Walter Scott. Zittel, K. A.v. (1894). Ontogenie, Phylogenie und Systematik. Congrès gèologique International. Compte-Rendu VI. Session, 125e136.

Please cite this article in press as: Tamborini, M., The reception of darwin in late nineteenth-century German paleontology as a case of pyrrhic victory, Studies in History and Philosophy of Biological and Biomedical Sciences (2017), https://doi.org/10.1016/j.shpsc.2017.09.004