(b. Würzburg, Germany, 21 August 1826; d. Heidelberg, Germany, 14 June 1903),
comparative anatomy and morphology, zoology.
Beginning his academic career as a medical student, Gegenbaur quickly became a leading invertebrate zoologist and, soon thereafter, emerged as the foremost vertebrate morphologist of the Darwinian era. Assiduity, thoroughness, and a keen insight into formal relations characterize his scientific endeavors. Through his efforts in the classroom and laboratory and, above all, his series of closely reasoned and richly documented monographs, Gegenbaur largely established the priority of comparative anatomy in the task of phylogenetic reconstruction and determined the centrality of this science within the biological curriculum, a position it has lost only with the rise since 1940 of molecular biology.
Gegenbaur’s parents, Franz Joseph Gegenbaur and Elisabeth Karoline Roth, belonged to well-established Roman Catholic families of southern Germany. His father’s responsibilities as an official in the Bavarian tax administration led to numerous moves and varied educational experiences during Carl’s younger years. In 1838, however, rigorous training began, first at the Würzburg Lateinschule and then at that city’s Gymnasium. The severity and narrowness of spirit of these schools decided Gegenbaur’s lifelong aversion to ecclesiastical practices and personnel. Nonetheless, in these schools he received exemplary training in the classics and, no doubt, in mental discipline. His hours out of school were devoted to impassioned self-instruction in all aspects of natural history, based largely on walking tours of the Franconian countryside.
There followed a conventional sequence of university training and service. Gegenbaur entered the University of Würzburg in 1845, completed the preparatory biennium philosophicum in 1847, and received the medical diploma in 1851. He immediately began to travel, visiting Johannes Müller in Berlin and, upon his advice, staying briefly at Helgoland to study the North Sea fauna. “For the first time [I saw] the sea!”1 he later exclaimed, and that experience was decisive. The following year (1852), with Rudolf Albert von Koelliker and Heinrich Müller, he began an eighteen-month research period at Messina. Returning to Würzburg. he presented his Habilitationsschritr (1854) and became a Privatedozent in zoology. Gegenbaur was called to Jena in 1856 as extraordinary professor of zoology in the medical faculty. He remained there, occupying various chairs (ultimately that of anatomy) until 1873, when he accepted the chair of anatomy recently vacated by his father-in-law, Friedrich Arnold, at Heidelberg. There Gegenbaur found himself once more in the familiar landscape of south Germany and in regular communication with an intimate friend from his Jena days, the philosopher Kuno Fischer. The Heidelberg years were especially devoted to the teaching of human anatomy, direction and improvement of the university’s important Anatomy Institute, and continued investigation of vertebrate morphology, a subject begun in earnest at Jena.
Gegenbaur presents a marvelous example of the better qualities of the higher professoriate during the golden years of the German universities. Absolute integrity, vast strength of mind and will, and a deep sense of responsibility characterized him. He attracted numerous outstanding students, including Oscar and Richard Hertwig, Max Fürbringer, T. W. Engelmann, Hans Gadow, and Giovanni Battista Grassi. These students were immediately exposed to the intensive research training that had marked Gegenbaur’s own education at Würzburg. Intellectually, Gegenbaur sought breadth, but only insofar as he could personally master in depth the subject matter before him; it was partly for this reason that at Jena he sought—successfully—to remove instruction in physiology from the responsibilities of the chair of anatomy.
The emphasis within Gegenbaur’s scientific studies shifted during his career. His anatomical investigations may be distributed into three periods. At Würzburg and during the early years at Jena his attention was devoted primarily to the life cycles and morphology of different stages of various marine animals. About 1860 he began the intensive comparative examination of vertebrate musculature, osteology, and neural structures upon which his great contemporary reputation so largely rested. This work continued during the third period, that at Heidelberg, and included much effort directed toward specifying in detail the anatomical bases for regarding man as an animal. During this period he also published several general and critical essays on the state of the anatomical art.
The materials and instruments of research were always an intrinsic component of the anatomisi’s concern. Gegenbaur was responsible for the care and augmentation of prepared and preserved specimens at both Jena and Heidelberg, a duty assumed with utmost seriousness. The research on invertebrates was pursued, as far as possible, with living specimens, thus necessitating study periods by the sea. At Würzburg, Gegenbaur had had the extraordinary good fortune to have received his scientific training from perhaps the finest group of research biologists then active in Germany: Rudolf Virchow, Franz Leydig, and Koelliker. He early learned microscopical techniques and the significance of the cell for organic structure and development. This was a critical lesson, for the formal and temporal priority of the cell provided the base point for interpretation in literally the whole of his scientific work.
Besides materials and instruments, Gegenbaur devoted much care to means of publication. His approximately 160 publications include both books and articles. Already an active participant in the meetings and publications of the Physikalischmedizinische Gesellschaft in Würzburg, he assumed primary responsibility for issuing the Jenaische Zeirschrift für Medizin und Naturwissen- schaft in 1864. This responsibility was relinquished when Gegenbaur went to Heidelberg; but soon thereafter (1875) he began his own journal. Morphologisches Jahrbuch, the outstanding vehicle for publication in comparative anatomy over several generations.
Fürbringer provides a detailed commentary on the scope and value of Gegenbaur’s diverse contributions to zoology and morphology.2 This work often was both descriptive and interpretive. Especially notable were the tracing of life cycles of freeswimming marine invertebrates, particularly of heteropods and pteropods (1855), and three lengthy monographs on the homologies of the vertebrate appendages and of the cranial bones (1864, 1865, 1872). Gegenbaur’s interests and orientation are strikingly similar to those of his contemporary T. H. Huxley, who also turned from invertebrate to vertebrate morphology and utilized both in the evolutionary campaign. Better-known are Gegenbaur’s interpretive manuals and student textbooks, in which first principles and superbly marshaled evidence are combined to exhibit the achievements and promise of comparative anatomy. Of these works the most significant is doubtless the second, much revised edition (1870) of Grundzüge der vergleichenden Anatomie, in which Gegenbaur explains how he, and fellow comparative anatomists, may—and most certainly should—transfer their allegiance from the traditional, static type concept to the new evolutionary interpretation announced by Charles Darwin.
A Quest for Generality . Gegenbaur was well aware of the propensity of anatomists to accumulate facts and to designate the ensuing mass of incidental description a science. He condemned this practice. It is evident… that mere description of isolated observed facts cannot be the highest goal of science.’3 Indeed, the major goal of comparative anatomy in nineteenth-century Germany was to escape the apparent mindlessness of traditional anatomical practice. Generalization was sought; and that generalization could only be the product of judgment, of a positive act of the mind as it reflects on organic similarities and dissimilarities. Gegenbaur and his contemporaries, most notably his colleagues at Jena. Matthias Schleiden and Ernst Haeckel, were thoroughly imbued with the episternological demands of the post-Kantian generation. There had been diverse reactions to the speculative excesses of the Naturphilosophen, including not only banal empiricism and programmatic physicochemical reductionism but also an effort, based ultimately on Kant, to find a dependable manner of moving from seemingly bare fact or facts to increasingly broader generalization. The understanding itself would play a positive role in the creation of general propositions, but its lead was always subject to the rigorous control of empirical evidence.
The mind thus imposed its own order on nature, Gegenbaur held. But in doing so, neither willful imagination nor directionless intuition deserved serious consideration. At the heart of the matter, the guidance and control of the reflective mind, lay the generally accepted factual wealth of the science and, above all, comparison. The adjective “comparative” in the term “comparative anatomy” was most certainly a working word. Comparison, wrote J. V. Carus, is the “base point” of the science, not as mere procedure but “as the form of observation appropriate to morphological relationships.”4 Obviously these relationships themselves (as such, being the very essence of things, they could not, of course, be demonstrated) could well anticipate the comparative assessment by which they were revealed. Before 1859 those of organic form were expressed by the concept of animal types to which the perceived diversity of organisms could be reduced; after 1859 many morphologists, including Gegenbaur, explained all relationship by means of common descent.
But whatever interpretation was imposed, the objective of comparative anatomy as a science was the pursuit of such generalizations. “The combinatory power of the mind,” wrote Gegenbaur. “enters at precisely that point where sensory perception reaches its limit.”5 Man’s mind is truly an “ordering mind,”6 and it generates those categories—species, genera, types—that simple sensory experience alone would fail to disclose. Gegenbaur’s emphasis on generalization reached through comparison therefore served a double purpose: it exhibited structural similarity and dissimilarity among organisms, thereby revealing broader patterns of relationship, and it directed attention to the comparative procedure itself, the indispensable basis of a genuine science of form.
Any consideration of Gegenbaur’s general outlook on the sciences requires at least brief notice of his passionate allegiance to the Old Catholic position. In the midst of the many conflicts dividing nineteenth-century Germany (Gegenhaur, incidentally, favored unification and spoke approvingly of Bismarck), that provoked by religious dispute within the Roman Catholic Church was prominent. Pressed by growing unbelief and increasing secularism, after mid-century the papacy and its defenders mounted an ever more determined effort to reassert universal authority over matters both temporal and spiritual. The movement culminated in the definition of papal infallibility at the Vatican Council (1869-1870). A furious reaction, led by lgnaz von Döllinger, immediately arose in Germany, creating a small but autonomous assemblage of German and Dutch Roman Catholics known thenceforth as “Old Catholics.”
Virtually all commentators on Gegenhaur’s career remark upon his keen sympathies for the traditional German forms of church organization and practice. None, however—and Gegenbaur provides no more in his autobiography — offers even minimal exposition of his views. It is clear from the autobiography, nonetheless, that Gegenbaur held the church militant in total contempt. its spokesmen—the clergy and, above all, the Jesuits—he obviously loathed. Church control over the mind—— he had reported its effects upon himself in the Jesuit schools of Würzburg—was surely even a greater affront to him than its loudly proclaimed pretensions to social control and political power.
There is no witness to Gegenbaur’s adherence to or neglect of expected practices of worship (his second marriage was outside the church). Certainly, however, church doctrine—which, in contrast to that of various Protestant sects, was remarkably discrete— had absolutely no bearing upon Gegenbaur’s acceptance and propagation of the descent theory. That acceptance occurred probably early in the 1860’s: at least by the following decade he was insisting to his Heidelberg students that man was but another animal. Unlike his friend Haeckel, Gegenbaur did not shrilly preach his rejection of accepted Christian teaching regarding the uniqueness of man, but preach he did—and with powerful effect. The many editions of his Lehrbuch der Anatomie des Menschen conveyed above all the following single message: “The human organism is not isolated in nature, but is only one member of an endless series within which knowledge of the whole will illuminate that of the individual.”7 Gegenbaur’s proud assertion of Lehrfreiheit tolerated no opposition from ultramontane or other sources.
The Central Generalization: Descent Theory . Prior to the 1860’s Gegenbaur freely accepted the type concept as the basis for interpreting the facts of zoology. At Jena, however (presumably in 1861 or 1862), he discovered Darwin and the descent theory. Over the next several years the descent theory achieved full hegemony over Gegenbaur’s zoological and anatomical outlook. In this conversion Gegenbaur had the constant and enthusiastic support of his intimate friend Haeckel. The latter, attracted to study at Jena by Gegenbaur, soon joined the faculty there and in 1862 launched the first of his notorious series of polemical lectures on Darwinism. Throughout the 1860’s, while Haeckel was composing the text of the Generelle Morphologie (1866). his preeminent contribution to discussion of the general issues evoked by Darwin, Gegenbaur was preparing his own response to the new doctrine, first fully expressed in the second edition of Grundzüge der vergiechenden Anatomie (1870),
Despite the great differences in the emotional character and public style of the two men, they were joined by close personal and intellectual bonds. They shared a lively skepticism regarding the truth—or at least the wide application—of the doctrines of revealed religion, and harbored a splendid disdain for the clergy. They agreed that an explanation of the origin and interrelationship of organic forms — that is, of the structural patterns of animals and plants—taken both in particular and in general, was the foremost problem facing the zoologist or morphologist. By so defining the issue, they emphasized that morphology, the science of organic form, and not zoology, with its traditional emphasis upon description and classification, was of paramount concern to the post-Darwinian biologist. Morphology (itself built upon comparative anatomy and descriptive embryology) would not, obviously, altogether replace zoology; it was deemed, nonetheless, the more significant science because it promised to explain, at a seemingly more basic level, the otherwise incidental facts of zoology.
The type concept, the predominant morphological and zoological generalization of the first half of the nineteenth century, was the creation of a sizable group of Continental biologists, notably Goethe, Cuvier, and K. E. von Baer. According to these authors, the great diversity of form manifested in the animal world could be reduced to a small number of units or even a single unit, called types. Each type exhibited a unique set of characteristics. Its many members, however dissimilar they might at first glance appear to be, nonetheless presented common elements in the essential features of their organization; and each member, by sharing these same elements, stood absolutely apart from members of all other types.
The mature type concept of the I 840’s and 1850’s assured the morphologist and zoologist that the animal kingdom was neither an uncontrollable chaos of altogether separate (and therefore unclassifiable) species—the view entertained by the taxonomic nominalist—nor were animals to be gathered together as simple permutations on a single structural theme—the opinion voiced by the still lively school of the Naturphilosophen and against the claims of which the morphological typologists were most firmly decided. It was in this milieu that Gegenbaur received his intellectual formation. Until the 1860’s he expressed full agreement with the premise of many typologists that the recognition and ever closer specification of the formal limits of the individual types was the principal objective of the science. Speculation regarding the putative origin of these types and their genealogical connections (the latter an inadmissible hypothesis) was excluded from consideration: morphology must attend to formal relationships only.
But the relationships between those types (Gegenbaur recognized seven: Protozoa, Coelenterata, Echinodermata, Vermes, Arthropoda, Mollusca, and Vertebrata) were indeed suggestive and lent themselves easily, when that option became clearly visible, to an alternative explanation. Gegenbaur’s acceptance of Darwinian descent theory was very carefully circumscribed. The theory of natural selection and its associated themes, particularly the laws of variation and heredity and the fact of adaptation, lay beyond his concern. Analysis of the dynamics of the evolutionary process he altogether ignored. It was, rather, the evolutionary product, the great diversity of organisms produced in time, that commanded attention. Gegenbaur sought to explore the pertinence of the vast fund of morphological fact for the demonstration of the likelihood of the occurrence of evolutionary change and to expand that fund as far as possible in support of the new interpretation. His turn to descent theory was, as he recognized, an act of judgment, the deliberate adoption of a new interpretive stance to obtain a better grasp of the order of organic nature. Unlike Haeckel, Gegenbaur refused to assert that organic descent was a proven or even a demonstrable fact. It was, instead, a leading idea, a guide for the reason in its encounter with the myriad and apparently unrelated facts of natural history. As such, it mediated between pure reason and bare empiricism and, in so doing, enabled morphologist and zoologist better to apprehend the overall temporal and formal patterns of nature.
Concretely expressed. Gegenbaur’s adherence to descent theory led in 1870 to an explicitly genealogical formulation of the relationship between the types. From the lowly Protozoa had arisen, in time, the entire array of living and extinct animal forms. Man stood at the apex of this ascent; other groups of animals (still designated “types”) were at lower levels along the main evolutionary trunk or, more commonly, were branches from that trunk. “We recognize,” Gegenbaur wrote, “that there exists a connection between the separate lines of descent…. [W]e can deal with the relations between types in a manner no different from that with which we treat of divisions within the types, that is, [treat them as expressions of] genealogical diversification.”8 The revised edition of the Grundzuge, as well as both editions (1874, 1878) of its simplified offspring, Grundriss der vergleichenden Anatomie, described the comparative morphology and presumed evolutionary connections of the invertebrate as well as vertebrate types. Gegenbaur’s final synthetic work, Vergleichende Anatomie der Wirbelthiere (1898 — 1901), however, placed overwhelming emphasis upon the vertebrates. This work thus is a fair representation of the emphasis of the morphologist’s later years. Through Gegenbaur’s influence and that of contemporary morphologists of like mind, such as Robert Wiedersheim, comparative vertebrate anatomy began increasingly to represent the summa of post-Darwinian investigation of the major lines of evolutionary descent and to provide the indispensable foundation for the introduction of the serious student to biological inquiry, a movement well advanced by the 1890’s.
An important reason for Gegenbaur’s overriding concern with the priority of comparative anatomy in evolutionary reconstruction (phylogeny) was his doubts concerning the scope and accuracy of alternative approaches. Paleontology, he conceded, provided direct access to the history of life. Regrettably, however, the fossil record seemed altogether too fragmentary to allow the re-creation of what was, above all else, a quite literal continuity: the course and changes of living forms over vast periods of time. He appears to have expected more from what Haeckel had in 1866 dramatized as the biogenetic law (“Ontogeny is the summary and rapid recapitulation of Phylogeny”) and that over the several preceding decades had received much sympathetic attention. If the developing embryo of a presumably higher form—a vertebrate, for example—did indeed display, in strictly comparable temporal and spatial order, the evolutionary history that lay behind its own existence, then an altogether extraordinary instrument of study was at hand. One need only carefully observe the emerging complexity of such an embryo in order to discover facts (“stages” of development) that would allow one to fill in those gaps inevitably present in the known fossil record.
Originally agreeable to the idea, by the 1880’s Gegenbaur was vigorous in pointing out the pitfalls facing the unrestrained recapitulation theorist. Most important, he emphasized that caenogenesis (another of Haeckel’s neologisms) was a common and perhaps ubiquitous phenomenon. Caenogenesis referred to new evolutionary acquisitions by the embryo itself and thus dealt with parts or processes that, when manifest in the embryo, could not be accepted as reliable indicators of the ancestral condition or conditions from which the parent stock of the embryo had arisen. Basically, Gegenbaur’s criticism cast all in doubt, for how was one to decide whether a given part or process was caenogenetic or palingenetic (that is, truly indicative of ancestral conditions)? Here, once again, was a problem demanding the most exquisite exercise of judgment. Gegenbaur suggested that probably the best standard for venturing such judgments was provided by comparative anatomy, the science that could at least provide some reliable representation of the forms and interrelations of form toward which any known embryo was developing. The damage, however, had been done, for Gegenbaur was persuaded that ontogeny presents “no true likeness” of phytogeny.9
This negative assessment of the recapitulation theory led to a fürther conclusion. Just as paleontology had displayed shortcomings in its fitness for phylogenetic studies, so did the great expectations placed on embryology fail. From this fact Gegenbaur drew great comfort: surely comparative anatomy must assume leadership in the high function of re-creating the history of life. Comparative anatomy therefore commanded priority of place among all the special disciplines dealing with the descent theory. In 1858 the chair of zoology at Jena (which included anatomy among its responsibilities) was divided. Gegenbaur, who had demanded this action, retained the chair of zoology and comparative anatomy; a new chair devoted exclusively to physiology was occupied by Albert von Bezold. This event marked an important moment in the development of the natural sciences within the German university organization. Physiology, long tied to anatomy as anatomia animata). was given independence and could develop according to its own needs and opportunities. Upon the death of Johannes Müller (1858), the most important anatomical chair in Germany, that at Berlin, was split following the Jena example. Anatomical responsibilities fell to Karl Bogislaus Reichert and physiological ones to Emil du Bois-Reymond.
But, to Gegenbaur, anatomy also had been liberated. On diverse grounds he had been seeking, and throughout his career continued to seek, a rationale and appropriate institutional foundations for the autonomy of anatomical investigation. Physiology, increasingly an experimental science, had now received its rights. Descent theory presented a host of novel possibilities for investigation; and anatomy, especially comparative anatomy, appeared to be uniquely suited to this task. Gegenbaur continued the abiding interest of German morphologists in form itself—that is, in the interrelations between the purely spatial disposition of the parts of organisms. These several motives lay behind Gegenbaur’s unrelenting defense of the rights of anatomy and his claims for its central role in biological research and generalization.
1. Gegenbaur. Erlebtes und Erstrebtes,. p. 58.
2. M. Fürbringer. “Carl Gegenbaur” in Heidelberger Professoren,, 426-450; also in Gesammmelte Abhandlungen. I, iii-xxii.
3. Gegenbaur. “Condition and Significance of Morphology.” 40.
4. J. V. Cams. Svstetn der thierischen Morphologie (Leipzig, 1853), 30.
5. Gegenbaur, Grundzüge, 2nd ed.. 76.
7. Gegenbaur, Lehrbuch. 4th ed. (1890), 2.
8. Gegenbaur, Grundzüge. 2nd ed.. 77.
9. Gegenbaur, “Cänogenese.” 496.
I. Original Works An exemplary bibliography (with subject divisions) of Gegenbaur’s publications is M. Fürbringer, “Systematisches Verzeiehnis der Ver-öffentlichungen von Carl Gegenbaur” in HeideWerger Professoren atts dem 19. Jahrlmnderi,II (Heidelberg, 1903), 455-466; it reappears in modified form in Gegenbaufs Gesammelie Ahhandlttngen, III . 575-597. The anatomist’s principal monographs are Unterstichungen über Fteropodcn and Hctcropodcn. Ein Beitrag zur Anatomie und ErtfwicfdungSgeschichte dieser Thiere (Leipzig, 1855); GrundzUge der vergleichenden Anatomic (Leipzig, 1859; 2nd ed., rev., 1870); Untersuchungen der verlekhenden Anatomic der Wirbeithiere: Erstes Heft. Carpus und Tarsus (Leipzig. 1864). Zweites Heft. Schultergürtel der Wirbeithiere und Brustfiosse der Fische (Leipzig, 1865), Drittes Heft. Das Kopfskelet der Selachier, Ein Beitrag zur Erkenntniss der Genese des Kopfskeletcs der Wirbelthicres (Leipzig. 1872); Grandriss der verleichcndcn Anatomic (Leipzig, 1874; 2nd ed., 1878): Lehrhuch der Anatomic des Menschen (Leipzig, 1883: 7th ed., 2 vols.. 1898-1899): andVcruleichende Anatomic der Wirhelhicre mit Berücksichtigung der Wirbeltoscru 2 vols. (Leipzig, 1898 - 1901).
Gegenbaur’s articles have been collected and published as Gesammelte Abhandlungen von Carl Gegenbaur, M. Fürbringer and H. Bluntschli, eds., 3 vols. (Leipzig, 1912). Those dealing with matters of general biological interest include De animalium plantarnmque regni terminis et differentiis (Leipzig, 1860), a rare 16 page oration also in Gesammelte AhhandJunigen, II, I- 13; “Die Stellung und Bedeutung der Morphologic” in Morphologisvhes Jahrbuclu 1 (1875). 1-19, trans, as “The Condition and Significance of Morphology.” in The Interpretation of Animal Form. W. Coleman, ed. and trans. (New York, 1967), 39-54; “Einige Bemerkungen zu Gotte’s” “Entwieklungsgcschiehte der Unke als Grundlage einer vergleichenden Morphologie der Wirbelthiere.” In Morphologisvhes Jahrbuclu 1 (1875). 299-345: “Canogenese.” in Anatomischer Anzeiger, 3 (1888), 493-499; and “Ontogenie und Anatomie in ihren Wechsclbeziehungen betrachtet.” in Morphologisvhes Jahrbuclu 15(1889). I-9.
II. Secondary Literature There is no major biography of Gegenbaur. His autobiography, Erlebtes und Erstrebtes (Leipzig, 1901), provides an invaluable account of the Würzburg and Jena years. Dependent upon the autobiography, but also presenting much additional information, are two accounts by M. Fürbringer: “Carl Gegenbaur,” in Heidelberger ProfessorenII , 389-466, and, in briefer form, “Carl Gegenbaur,” in Anatomisch—er Anzeier, 23 (1903), 589-608. Also useful is E. Goppert, “Gegenbaur, Karl,” in Biographisches Jahr—buch und deutscher Nekrolog, 8 (1903). 324-339 (Gegenbaur did not adopt the spelling Karl).
On Gegenbaur as investigator and teacher, see Fried—rich Maurer, “Carl Gegenbaur, Rede zum Gedäehtnis seines 100. Geburtsjahres,” in Jenaische Zeitschrift für Medizin und Naturmssenschaft55 (1926), 501-518; and E. Goppert, “Karl Gegenbaurs genetische Methode im anatomischen Unterricht,” in Anatomischer Anzeiger,63 (1927); and “Friedrich Maurer und der Kreis um Carl Gegenbaur,” in Anatomischer Anzeiger,85 (1938), 313 —331. E. S. Russell offers extended discussion of nineteenth—century morphological interpretations in Form and Function (London. 1916); on Gegenbaur and Haeckel see esp. 246-267; see also the brief comment by William Coleman, Interpretation of Animal Form (New York, 1967), xi-xxx. For a description and assessment of why and how Gegenbaur converted to the descent theory in the 1860’s, consult William Coleman, “Morphology Between Type Concept and Descent Theory,” in Journal of the History of Medicine,31 (1976), 149-175.
On comparative anatomy since Cuvier, the basic source remains Wilhelm Lubosch, “Geschichte der ver—gleichenden Anatomie,” in Bolk et al., eds., Handbuch der Anatomie der Wirbelthiere,I (Berlin, 1931). 3-76: Lübosch must he approached with care, however, for he presents an unannounced apology for idealistic or “pure” morphology as well as an ill—disguised nationalistic bias. A classic article on problems central to Gegen—baur’s inquiry that also provides valuable histórical insight is Hans Spemann, “Zur Geschichte und Kritik des Begriffs der Homologie.” in C. Chun and W. Johannsen. eds., Aligemeine Btologie, in the series Kultur der Gegenwart (Leipzig, 1915), 63-86. Georg Uschmann, Geschichte der Zoologie und der zoologischen Anstaiten in Jena. 1779- 1919 (Jena, 1959), 27-62, describes Gegenbaur’s work at Jena in great detail and provides an insight into how and when Gegenbaur and Haeckel discovered Darwin. On descent théories in Germany before The Origin of Species, see the indispensable article by Owsei Temkin: “The Idea of Descent in Post-Romantic German Biology: 1848-1858,” in Bentley Glass et al., Forerunners of Darwin: 1745-1859 (Baltimore, 1959), 323-355.
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