Mendel in Darwin's Shadow

Abbreviations, notes, and works cited

Famous Biologists


Notes, works cited, and an abbreviation key for the article Mendel in Darwin's Shadow by David Allen appear on this page.


BM  Nägeli, C. (1866) Botanische Mittheilungen

LAC  Callender, L. A. (1988) 'Gregor Mendel: An Opponent of Descent with Modification'

OG  Origin of Genetics (1966) Stern & Sherwood (eds.),

VB  Gärtner, C. F. von (1849) Versuche und Beobachtungen über die Bastarderzeugung im Pflanzenreich


1. 'Dr. Confuser' is actually Charles H. Porter.

2. All subsequent references to the article 'Gregor Mendel: An Opponent of Descent with Modification' by L. A. Callender (1988) are referred to as LAC in the text.

3. See Robert Olby, 'Mendel No Mendelian?'. In: History of Science (1979), 17, pp. 55-72.

4. In Science vs. Religion, Fuller actually cites Olby as the source for his description of Mendelian inheritance as a 'mathematically rigorous version of special creationism'—Fuller 2007 p. 135.

5. Fuller has said that he is not personally an 'ID-er', but feels that the movement should have a 'fair run for its money'. See: 'Steve Fuller : Designer trouble', by Zoë Corbyn. In: The Guardian, 31 January, 2006 (last accessed 16 August, 2009). Yet he has published numerous books and articles advocating ID, so his claims for 'fellow traveller' status seems dubious.

6. Translated by the author. Unless otherwise specified, all translations are my own.

7. The exact date Mendel began his Pisum experiments is not known. He told Nägeli that they began in 1856 (OG p. 60); but it is likely that, prior to this, he tested the constancy of traits in his plant specimens over several generations.

8. All subsequent references to the book Origin of Genetics (1966), edited by Stern & Sherwood, are marked OG in the text.

9. In the 2nd German edition of The Origin of Species, published 1863 (the edition which Mendel read), Entwicklung was used to translate the word 'development'. In the context in which it appeared, it clearly implied development (e. g. embryo development) rather than 'evolution'. However, despite its associations with embryology and development, Haeckel used the term as early as 1863 to mean Darwinian evolution—for example, in his paper, 'Über die Entwicklungstheorie Darwins'.

Darwin introduced the word 'evolution' into the sixth edition of Origin (1872). Just to complicate matters, the German version of that edition (1876) translated the word sometimes as Entwicklung, and sometimes as Evolution.

10. Predictably, Haeckel suggested that Descendenztheorie 'shows us that ontogeny is nothing more than a short recapitulation of phylogeny'—1866, p. 7.

11. All subsequent references to the book Versuche und Beobachtungen über die Bastarderzeugung im Pflanzenreich by Gärtner (1849) are marked VB in the text.

12. Müller-Wille & Orel argue that 'Mendel’s open disregard for the species-variety distinction is rather troublesome for interpretations that see him as part of the hybridist tradition. According to these interpretations, Mendel was interested in the question whether hybridization could give rise to new species. Yet, if it was that question which interested Mendel, then the taxonomic status of his experimental plants should have mattered to him.' (2007, p. 173.) But it is clear that, like Darwin and Nägeli, Mendel believed that species and varieties were shifting entities. As I argue later in the article, he was interested in the creation of new stable forms through hybridisation; and these 'attain the status of new species' (OG p. 41).

Müller-Wille & Orel also make much of the fact that Mendel referred, in the main body of the Pisum paper, to the different pea plants he used as species rather than varieties. They infer from this that he was, in fact, applying the 'strictest definition' of species, based on constant characteristics. However, this ignores the fact that, while most of the plants Mendel was using were varieties of one species (Pisum sativum), he was also using plants from other 'species' (as they were defined at the time)—viz. P. quadratum, P. saccharatum, and P. umbellate. Thus, both species and varieties were mixed up in his experiments. It may have been simpler for him, then, to use just one word (Arten), as a general term for the different types of plants he was using, rather than continually trying to distinguish between species and varieties (especially as he said that the distinction was not important to him).

Müller-Wille & Orel object that English versions of the Pisum paper have generally (mis)translated Arten as 'varieties' rather than as 'species'. They have a point. However, as we now classify all the plants that Mendel was using as varieties rather than species, it could be argued that it would confuse modern readers if translations used the term 'species. '.

13. All subsequent references to the book Botanische Mittheilungen by Carl Nägeli (1866) are marked BM in the text.

14. Lönnig is a geneticist at the Max-Planck-Institut für Züchtungsforschung in Cologne. A scandal broke in 2003, when scientists (led by Ulrich Kutschera) protested that Lönnig was using the Institute's web server to post anti-evolutionary material. Lönnig responded that he was the victim of an inquisition by adherents of the 'totalitarian-materialistic theory of evolution'. See 'Die Affäre Max Planck. Über die fragwürdigen Diskursmethoden eines Evolutiongegners' by M. Neukamm and A. Beyer, in Kutschera (ed.) (2007), pp. 232-276.

15. It might be argued that Kölreuter and Gärtner were driven, in their work, by an opposition to Linnaeus. It is notable, however, that in his book, Versuche und Beobachtungen… (1849), Gärtner barely mentions Linnaeus; and makes no reference at all to the Linnaean theory of descent through hybridisation. In reviewing the history of debates about plant hybridisation, he actually cites Camerer as the first important theorist on the subject (p. 4). He records the work of early hybridists such as Morland, Blair, Fairchild and Kölreuter; but not Linnaeaus. The debate had moved on, it seems; other people, even for Gärtner, were now the main reference points, and the main opponents.

16. Some of Nägeli's ideas were, in fact, taken up by ant-Darwinists; nevertheless, the notion that he himself was, broadly, a supporter of Darwinism still stands. See Junker 2002.

17. Mendel's marking of this passage is noted by Richter (1943), p. 203.

18. Similarly, in a letter to Nägeli, Mendel noted that the Hieracium experiments had led him to conclude that, in this genus at least, 'fertilization with foreign pollen can occur only if self-fertilization fails'—OG p. 101.

19. See Peter's detailed descriptions of various hybrids: 1884 pp. 450-496; & 1885 pp. 111-134.

20. There are three Hieracium subgenera, and they differ in their mode of reproduction. It is believed that all Chionoracium species are sexual diploids; subg. Hieracium species are either polyploid obligate apomicts (of the diplosporous type), or diploid sexuals; and Pilosella 'is characterized by a mixture of sexual and facultatively apomictic taxa of the aposporous type'—Fehrer et al, 2005 p. 176.

Works Cited

Arnold, Michael Lynn (2006). Evolution Through Genetic Exchange (Oxford: OUP).

Bateson, William (1913). Mendel's Principles of Heredity, 3rd impression (Cambridge: CUP).

Bishop, B. E. (1996). 'Mendel's Opposition to Evolution and to Darwin'. In: Journal of Heredity 87 (3), pp. 205-213.

Blaringhem, Louis (1919). Les Problèmes de l'Hérédité Expérimentale (Paris: Ernst Flammarion).

Blaringhem, Louis (1920). ‘Biologie végétale. Stabilité et fertilité de l'hybridé Geum urbanum L. × G. rivale L. Note de M. L. BLARINGHEM, présentée par M. L. Guignard.’ In: Comptes rendus hebdomadaires des séances de l'académie des sciences, Vol. 170 (Paris: Gauthier-Villars), pp. 1284-1286.

Bowler, Peter J. (2003). Evolution: The History of an Idea, 3rd edition (Berkeley & Los Angeles / London: University of California Press).

Callender, L. A. (1988). 'Gregor Mendel: An Opponent of Descent with Modificaton'. In: Hist. Sci. xxvi, pp. 41-75.

Cetl, Ivo (2002/3). 'Mendel's Hybridization Experiments with Other Plants than Pisum'. In: Folia Mendeliana 37-38, pp. 5-36.

Clausen, J. (1954). 'Partial apomixis as an equilibrium system'. In: Caryologia, Supplement, pp. 469–479.

Darwin, Charles (1859). Origin of Species (London: John Murray).

Darwin, Charles (2005). The Correspondence of Charles Darwin, Vol. 15 (1867) (Cambridge: CUP).

Darwin, Charles (2007). The Effects of Cross and Self Fertilisation in the Vegetable Kingdom (Middlesex: Echo Library).

Endersby, Jim (2007). A Guinea Pig's History of Biology (Harvard: Harvard University Press).

Fairbanks, Daniel J. & Rytting, Bryce (2001). 'Mendelian Controversies: A Botanical and Historical Review'. In: American Journal of Botany 88(5), pp. 737–752.

Fehrer J., Krahulcová A., Krahulec F., Chrtek J. Jr., Rosenbaumová R., Bräutigam S. (2007). 'Evolutionary aspects in Hieracium subgenus Pilosella'. In: Apomixis: Evolution, Mechanisms and Perspectives, ed. U. Grossniklaus, E. Hörandl, T. Sharbel, P. van Dijk. (Königstein: Koeltz. Series Regnum Vegetabile, No. 147), pp. 359-390.

Fehrer, J., Šimek, R., Krahulcová, A., Krahulec, F., Chrtek, J. Jr., Bräutigam, E., Bräutigam, S. (2005). 'Evolution, hybridisation, and clonal distribution of apo- and amphimictic species of Hieracium subgen. Pilosella (Asteraceae: Lactuceae) in a Central European mountain range.' In: Plant species-level systematics: new perspectives on pattern & process, ed. F. T. Bakker, L. W. Chatrou, B. Gravendeel & P. B. Pelser. (Königstein: Koeltz Scientific Books. Series Regnum Vegetabile No. 143), pp. 175-201.

Fisher, R. A. (1936). 'Has Mendel's work been rediscovered?'. In: Annals of Science 1, pp. 115-137.

Focke, Wilhelm Olbers (1881). Die Pflanzen-mischlinge: Ein Beitrag zur Biologie der Gewächse (Berlin: Gebrüder Borntraeger).

Fuller, Steve (2005). 'How Intelligent Design Came To Me', blog dated 14 November (last accessed 16 August, 2009).

Fuller, Steve (2006). 'Intelligent Design Theory: A Site for Contemporary Sociology of Knowledge'. In: Canadian Journal of Sociology/Cahiers canadiens de sociologie, 31(3), pp. 277-289.

Fuller, Steve (2007). Science vs. Religion? (Cambridge: Polity Press).

Gajewski, Wacław (1957). A Cytogenetic Study on the Genus 'Geum' L., Monographiae Botanicae, Vol. IV (Warsaw: PWN).

Gärtner, Carl Friedrich von (1849). Versuche und Beobachtungen über die Bastarderzeugung im Pflanzenreich (Stuttgart: Hering).

Gayon, Jean (1998). Darwinism's Struggle for Survival: Heredity and the Hypothesis of Natural Selection (Cambridge: CUP).

Glass, Bentley (1968). 'Heredity and Variation in the Eighteenth Century Concept of the Species'. In: Forerunners of Darwin 1745-1859, ed. Bentley Glass, Owsei Temkin & William L. Straus Jr. (Baltimore: John Hopkins Press), pp. 144-172.

Gliboff, Sander (1999). 'Gregor Mendel and the Laws of Evolution'. In: Hist. Sci., xxxvii, pp. 217-235.

Gliboff, Sander (2008). H. G. Bronn, Ernst Haeckel, and the Origins of German Darwinism: A Study in Translation and Transformation (Cambridge Mass. & London: MIT Press).

Haeckel, Ernst (1866). Generelle morphologie der organismen. Allgemeine Grundzüge der organischen formen-wissenschaft, mechanisch begründet durch die von Charles Darwin reformirte Descendenztheorie, vol. 2: Allgemeine Entwicklungsgeschichte der Organismen (Berlin: Georg Reimer).

Heimans, J. (1970). 'A Recently Discovered Note on Hybridization in Mendel's Handwriting'. In: Folia Mendeliana 5, pp. 13-24.

Henig, Robin Marantz (2001). A Monk and Two Peas: The Story of Gregor Mendel and the Discovery of Genetics (London: Phoenix).

Hoffmann, Hermann (1869). Untersuchungen zur Bestimmung des Werthes von Species und Varietät: ein Beitrag zur Kritik der Darwin'schen Hypothese (Gießen: J Richter).

Hoppe, B. (1971). 'Die Beziehung zwischen J. G. Mendel und C. W. Nägeli auf Grund neuer Dokumente'. In: Folia Mendeliana, 6, pp. 123–139.

Hornschuch, Christian (1848). 'Ueber Ausartung der Pflanzen'. In: Flora, No. 2 (pp. 17-28); No. 3 (pp. 33-44); No. 4 (pp. 50-64); No. 5 (pp. 66-86).

Hornung, Ernst (1832). Review of Beyträge zu einer kritischen Aufzählung der Schweizerpflanzen by Johannes Hegetschweiler. In: Literaturberichte zur allgemeinen botanischen Zeitung, Vol. 2, No. 13, pp. 193-208.

Iltis, Hugo (1932). The Life of Mendel (London: George Allen & Unwin).

Iltis, Hugo (1951). 'Gregor Mendel's Life and Heritage'. In: Genetics in the 20th Century: Essays on the Progress of Genetics During its First 50 Years, ed. L. C. Dunn (New York: MacMillan), pp. 25-34.

Junker, Thomas (1989). Darwinismus und Botanik: Rezeption, Kritik und theoretic Alternatien im Deutschland des 19. Jahrhunderts (Stuttgart: Deutscher Apotheker Verlag).

Junker, Thomas (2002). 'Carl Nägeli und der anti-Darwinismus—von der Vervollkommungstheorie zur Makroevolution'. In: Pratum Floridum: Festschrift für Brigitte Hoppe (Augsburg: Rauner), pp. 205-219.

Kelly, Alfred (1981). The Descent of Darwin: The Popularization of Darwinism in Germany, 1860-1914 (Chapel Hill: University of North Carolina Press).

Kerner, Anton [Anton Kerner von Marilaun] (1903). The Natural History of Plants, translated by F. W. Oliver. Vol. II (London: Gresham).

Kölliker, Albert (1879). Entwicklungsgeschichte des Menschen und der höheren Thiere, 2nd edition (Leipzig: Wilhelm Engelmann).

Krahulcovà, Anna; Krahulec, František; Chapman, Hazel M. (2000). 'Variation in Hieracium Subgen. Philomela (Asteraceae): What Do We Know About its Sources?'. In: Folia Geobotanica 35, pp. 319-338.

Kříženecký, Jaroslav (1965). Gregor Johann Mendel 1822-1884: Texte und Quellen zu seinem Wirken und Leben (Leipzig: Johann Ambrosius Barth).

Kutschera, Ulrich, ed. (2007). Kreationismus in Deutschland: Fakten und Analysen (Münster-Hamburg-Berlin-Vienna-London-Zurich: Lit Verlag).

Larson, James L. (1971). Reason and Experience: The Representation of Natural Order in the Work of Carl von Linné (Berkeley & Los Angeles: University of California Press).

Lönnig, Wolf-Ekkehard (1998, 2000, 2001). Johann Gregor Mendel: Warum seine Entdeckungen 35 (72) Jahre ignoriert wurden (last accessed 18 Sept. 2009).

Macfarlane, John Muirhead (1892). A Comparison of the Minute Structure of Plant Hybrids with that of their Parents, and its Bearing on Biological Problems (Edinburgh: Robert Grant & Son).

Margulis, Lynn (1996). 'Gaia Is a Tough Bitch'. In: The Third Culture: Beyond the Scientific Revolution, John Brockman (ed.) (New York: Touchstone). (Last accessed 27. 8. 09).

Mayr, Ernst (1982). The Growth of Biological Thought: Diversity, Evolution, and Inheritance (Harvard: Harvard University Press).

Mayr, Ernst (1997). Evolution and the Diversity of Life: Selected Essays (Harvard: Harvard University Press).

McCarthy, Eugene M. (2008). On the Origin of New Forms of Life: A New Theory.

Mertens, Franz Karl; Koch, Wilhelm Daniel Joseph (eds.) (1832). J. C. Röhlings Deutschlands Flora (Frankfurt-am-Main: Friedrich Wilmans).

Müller-Wille, S. & Orel, V. (2007). 'From Linnaean Species to Mendelian Factors: Elements of Hybridism, 1751-1870'. In: Annals of Science 64:2, pp. 171-215.

Nägeli, Carl (1865). Entsehung und Begriff der naturhistorischen Art (Munich: Verlag der königl. Akademie).

Nägeli, Carl (1866). Botanische Mittheilungen, Vol. II (Munich: F. Straub).

Nägeli, Carl & Peter, Albert (1885). Die Hieracien Mittel-Europas: Monographische Bearabeitung der Piloselloiden mit besonderer Berücksichtigung der mitteleuropäischen Sippen (Munich: R. Oldenbourg).

Nogler, Gian A. (2006). 'The Lesser-Known Mendel: His Experiments on Hieracium'. In: Genetics, No. 172 (January 2006), pp. 1-6.

Olby, Robert C. (1979). 'Mendel No Mendelian?'. In: History of Science, 17, pp. 55-72.

Olby, Robert C. (1985). Origins of Mendelism, 2nd edition (Chicago & London: University of Chicago Press).

Olby, Robert C. (1997). 'Mendel, Mendelism and Genetics' (last accessed 16, August 2009).

Orel, Vitěslav (ed.) (1984). 'Mendels Hausarbeit in Naturgeschichte von 1850'. In: Folia Mendeliana 18 (Special edition).

Orel, Vitěslav (1996). Gregor Mendel: The First Geneticist (Oxford: OUP).

Orel, Vitěslav (1998). 'Constant Hybrids in Mendel's Research'. In: History and Philosophy of the Life Sciences, 1998 20:3, pp. 291-299.

Peter, Albert (1884) 'Über spontane und künstliche Gartenbastarde der Gattung Hieracium sect. Piloselloidea'. In: Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie, ed. A. Engler (Leipzig: Wilhelm Engelmann). Vol. 5, pp. 203-286 & pp. 448-496.

Peter, Albert (1885) 'Über spontane und künstliche Gartenbastarde der Gattung Hieracium sect. Piloselloidea' [continuation of article from Peter, 1884]. In: Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie, ed. A. Engler. Vol. 6, pp. 111-136.

Potter, Andrew (2001). 'Gott würfelt nicht: Eine Filmbesprechung von Andrew Potter'. In: Studiengemeinschaft Wort und Wissen, 4/01 (December 2001)— A Review of the film Gott würfelt nicht (last accessed 28. 9. 09).

Prywer, Czesława (1932). 'Genetische Studien über die bastarde zwischen Geum urbanum L. und Geum rivale L. '. In: Acta Societatis Botanicorum Poloniae, Vol. 9, pp. 87-98.

Punnett, R. C. (1909). Mendelism, American edition (New York: Wilshire Book Co.)

Radl, Emanuel (1930). History of Biological Theories, translated and adapted by E. J. Hatfield (London: Humphrey Milford / OUP).

Richter, Oswald (1943). Johann Gregor Mendel wie er wirklich war: Neue Beiträge zur Biographie des berühmten Biologen aus Brünns Archiven (Brünn: Verhandlungen des Naturforschenden Vereins in Brünn).

Sachs, Julius (1882). Text-book of Botany: Morphological and Physiological, 2nd. edition, ed. Sydney H. Vines (Oxford: Clarendon Press).

Salter, Thomas Bell (1852). 'On the Fertility of Certain Hybrids'. In: The Phytologist: A Popular Botanical Miscellany, ed. Edward Newman, Vol IV, Part 2 (London: John Van Voorst), pp. 737-742.

Schleiden, Matthias (1842). Grundzüge der Wissenschaftlichen Botanik nebst einer Methodologischen Einleitung als Anleitung zum Studium der Pflanze (Leipzig: Wilhelm Engelmann).

Schleiden, Matthias (1853). The Plant: A Biography in a Series of Thirteen Popular Lectures, translated Arthur Henfrey. 2nd edition (London: Hipployte Bailliere).

Schmidt, Oscar (1873). Descendenzlehre und Darwinismus (Leipzig: F. A. Brockhaus).

Sermonti, Giuseppe (2005). Why is a Fly Not a Horse? (Seattle: Dsicovery Institute Press).

Stern, Curt & Sherwood, Eva R. (1966). The Origin of Genetics; A Mendel Source Book (San Francisco & London: W. H. Freeman).

Page 1 >>

Page 2 >>

Famous Biologists >>

Sturtevant, Alfred Henry (2001). A History of Genetics (Cold Spring Harbor: CSHL Press).

Syme, John T. Boswell (ed.) (1864). English Botany Coloured Figures of British Plants, Vol. III, 3rd ed. (London: Robert Hardwicke).

Unger, Franz (1852). Botanische Briefe (Vienna: Carl Gerold & Son).

Voipio, Paavo (1987). 'What Did Mendel Say About Evolution?'. In: Hereditas 107, pp. 103-105.

Waller, John (2002). Fabulous Science: Fact and Fiction in the History of Scientific Discovery (Oxford: OUP).

Weinstein, Eric W. (1996-2007) Nägeli, Karl von (1817-1891) (last accessed 20 August, 2009).

Zahn, K. H. (1921). Compositae-Hieracium: Sect. I. Glauca—Sect. VII. Vulgata (Anfang). Part of the series, Das Pflanzenreich: Regni vegetablilis conspectus, (ed.) A. Engler. (Leipzig: Wilhelm Engelmann).

Most shared on

Human Origins: Are we hybrids?

On the Origins of New Forms of Life

Mammalian Hybrids

Cat-rabbit Hybrids: Fact or fiction?

Famous Biologists

Dog-cow Hybrids

Prothero: A Rebuttal

Branches of Biology

Dog-fox Hybrids