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From Candolle to croizat: Comments on the history of biogeography

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References

  1. A. P. de Candolle, “Géographie botanique,” in Dictionnaire des sciences naturelles, XVIII (Strasbourg and Paris, 1820), p. 359; reprinted as Essai élémentaire de géographie botanique (Strasbourg, 1820). This essay summarizes and expands ideas published earlier: “Explication de la carte botanique de la France,” in Lamarck and de Candolle, Flore française, II (Paris, 1815), p. v; “Rapport sur un voyage botanique et agronomique dans les départemens de l'ouest ... du sud-ouest ... du sud-est ... de l'est ... du nord-est ... du centre,” Mémoires d'agriculture, d'economie rurale et domestique, publiés par la Société d'Agriculture du département de la Seine, X (Paris, 1807), p. 228; XI (1808), p. 1; XII (1809), p. 210; XIII (1810), p. 203; XIV (1811), p. 213; XV (1812), p. 200; “Géographie agricole et botanique,” in Nouveau cours complet d'agriculture ou dictionnaire raisonné et universel d'agriculture, VI (Paris, 1809), p. 355, and reprinted in a slightly revised form in VII (Paris, 1822), p. 303; Mémoire sur la géographie des plantes de France, considérée dans ses rapports avec la hauteur absolue,” in Mémoires de Physique et de Chimie de la Société d'Arcueil, 3 (1817), 262, translated as “On the Effect of Elevation above the Level of the Sea upon the Geography of Plants in France,” in J. Sci. Arts, 4 (New York, 1818), 176; and in abbreviated form as “Memoir upon the Geography of the Plants of France, Considered More Especially with Regard to Their Height above the Level of the Sea,” in Annals of Philosophy, 6 (London, 1818), p. 408.

  2. C. Lyell, Principles of Geology, II (London, 1832). Excellent studies of the importance of Lyell and Prichard are in M. P. Kinch, “An Assessment of Rival British Theories of Biogeography, 1800–1859, master's thesis, Oregon State University, 1974, and R. A. Richardson, “The Development of the Theory of Geographical Race Formation: Buffon to Darwin, Ph.D. diss., University of Wisconsin, 1968.

  3. J. C. Prichard, Researches into the Physical History of Mankind, 3rd. ed., I (London, 1836), 4th ed., I (London, 1841). According to Stocking (G. W. Stocking, “Bibliography of James Cowles Prichard,” in J. C. Prichard, Researches in the Physical History of Man [Chicago, 1973], p. cxv), the fourth edition of this work is a reprint of the third edition (5 vols., [London: Sherwood, Gilbert, and Piper, 1836–1847]). As listed by Stocking, the fourth edition was reprinted by “Houlston and Stoneman, 1851.” The volume cited is apparently another “4th edition,” reprinted by the publisher of the third edition. No reference to Candolle appears in either the first or second editions of Prichard (London: John and Arthur Arch, 1813 and 1826). Prichard apparently learned of Candolle from Lyell's Principles.

  4. The same is true for many historians, according to Kinch, “Assessment,” p. 1. Botanists, however, usually begin with Linnaeus or even earlier writers.

  5. E. Mayr, “What is a Fauna,” in Evolution and the Diversity of Life: Selected Essays (Cambridge, Mass.: 1976), p. 552, slightly revised from an article of the same title in Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Tiere, 92 (1965), 473.

  6. Ibid.

  7. C. Darwin, Origin of Species (London, 1859), p. 347.

  8. Prichard, Researches, p. 50.

  9. Lyell, Principles, p. 68. Elsewhere, Lyell refers to Candolle's essay as “the most beautiful generalisation of a multitude of facts which I think was ever produced in natural history”; in K. M. Lyell, ed., Life, Letters and Journals of Sir Charles Lyell, Bart., I (London, 1881), p. 245. Further comments on the interaction between Candolle and Lyell are in L. G. Wilson, Charles Lyell, the Years to 1841: The Revolution in Geology (New Haven, 1972), noted also in A. de Candolle, ed., Mémoires et souvenirs de Augustin-Pyramus de Candolle (Geneva, 1862), p. 435. Relevant here are Egerton's comments and evaluation: “No one before de Candolle seems to have stated so clearly the factors involved in plant distribution,” F. N. Egerton, “Studies of Animal Populations from Lamarck to Darwin,” in J. Hist. Biol. 1 (1968), 232.

  10. Mayr, “Fauna” (1976), p. 552, revised from 1965 version, p. 473.

  11. Ibid. (1976), p. 553; 1965 version, p. 474.

  12. Ibid. and A. R. Wallace, The Geographical Distribution of Animals, I (London, 1876), p. 5.

  13. Ibid.

  14. Ibid.

  15. Lyell, Principles, p. 66. With respect to plants, brief statements to a similar effect are made by Linnaeus, Philosophia Botanica (Stockholm, 1761), p. 264. His early achievements in geographical studies of plants are summarized in G. E. Du Rietz, “Linné Som Fjällväxtgeograf,” in Svenska Linné-Sällskapets Arsskrift, 25 (1942), 33. The earlier history of these ideas is given in an article, remarkable for its thoroughness, by N. von Hofsten, “Zur älteren Geschichte des Diskontinuitätsproblems in der Biogeographie,” Zoologische Annalen, Zeitschrift für Geschichte der Zoologie, 7 (1919), 197.

  16. Lyell, Principles, p. 87. Here Lyell quotes from an English translation of Buffon, Natural History, General and Particular, by the Count de Buffon, trans, W. Smellie, V (London: Strahan and Cadell, 1781), pp. 432–433: “We mean not to deny the possibility of equal climates, in both Continents, producing some animals of the precise same species. We formerly remarked, that the same temperature, in different parts of the globe, should produce the same beings, both in the animal and vegetable kingdoms, if all other circumstances were equal. We treat not here, however, of philosophical possibilities, which may be more or less probable, but of a very general fact, of which numberless examples may be given. It is certain, that, when America was discovered, none of the following animals existed in the New World.” The original reads: “Ce n'est pas qu'absolument parlant, et même raisonnant philosophiquement, il ne fût possible qu'il se trouvât dans les climats méridionaux des deux continens, quelques animaux qui seroient précisément de la même espéce; nous avous dit ailleurs (C), et nous le répétons ici, que la même température doit faire dans les différentes contrées du globe les mêmes effets sur la Nature organisée, et par conséquent produire les mêmes êtres, soit animaux, soit végétaux, si toutes les autres circonstances étoient comme la température, les mêmes à tous égards; mais il ne s'agit pas ici d'une possibilité philosophique, qu'on peut regarder comme plus ou moins probable; il s'agit d'un fait et d'un fait très-général, dont il est aisé de présenter les nombreux et très-nombreux examples. Il est certain qu'au temps de la découverte de l'Amérique, il n'existoit dans ce nouveau monde aucun des animaux que je vais nommer” (Buffon, Histoire naturelle générale et particulière, supplément, III [Paris, 1776], p. 270). In the translation, the words “philosophical possibilities” appear, but in the original they are in the singular, not the plural. Lyell's reference is to “Buffon, vol. v.-On the Virginian Opossum,” which is not a reference to any French edition of Buffon, but accords with the above and other English editions: “Second Edition,” V (London, 1785), pp. 432–433; “Third Edition,” V (London, 1791), pp. 432–433. Subsequent editions of Smellie's translation are slightly altered: “A New Edition,” VII (London: Cadell and Davies, 1812), p. 113; “A New Edition Corrected and Enlarged, I (London: Thomas Kelly, 1866), p. 529.

  17. Lyell, Principles, p. 87.

  18. The definitive birth of plant geography is often considered to be marked by the essay of A. von Humboldt and A. Bonpland, Essai sur la géographie des plantes (Paris, 1805) and its German equivalent, Ideen zu einer Geographie der Pflanzen (Tübingen, 1807, and Vienna, 1811). A reprint of the French edition was published by the Institut Panaméricain de Géographie et d'Histoire (Mexico City, 1955), another as “Sherborn Fund Facsimile No. 1” (London: Society for the Bibliography of Natural History, 1959), and still another in N. Zuniga, Humboldt y la Geográfia de las Plantas (Quito, 1964). A reprint of the German edition is in M. Dittrich, ed., Ideen zu einer Geographie der Pflanzen von Alexander von Humboldt (Leipzig, 1960). A collection of recent papers on Humboldt is in W. T. Stearn, ed., Humboldt, Bonpland Kunth, and Tropical American Botany (Lehre, 1968).

  19. Buffon, Histoire naturelle générale et particuliére, IX (Paris, 1761), p. 96 (translated).

  20. Ibid., p.118 (translated).

  21. A. de Humboldt, “Sur les lois que l'on observe dans la distribution des formes végétales. Extrait d'un mémoire lu à l'institut, dans la séance du 5 février 1816 [29, January according to Procés-verbaux des séances de l'Académie, VI (Hendaye, 1915), p. 11],” Annales de Chimie et de Physique, 2nd ser., I (1816), 234 (translated); translated as “On the Laws Observed in the Distribution of Vegetable Forms”, Philo. Mag. and J., 47 (1816), 446; “Ueber die Gesetze, welche Man in der Vertheilung der Pflanzenformen beobachtet”, Journal für Chemie und Physik, 18 (1916), 129; ‘Ueber die Gesetze in der Vertheilung der Pflanzenformen beobachtet”, in Isis oder Encyclopädische Zeitung von Oken, I (1817), col. 177.

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  22. Humboldt, “Distribution”, p. 235 (translated). Four years later Humboldt published a different article with the same title, in Dictionnaire des sciences naturelles, XVIII (Paris, 1820), p. 422; translated as “New Inquiries into the Laws Which are Observed in the Distribution of Vegetable Forms”, in Edinburgh Phil. J., 6 (1822), 273. A slightly expanded version was published as “Nouvelles recherches sur les lois que l'on observe dans la distribution des formes végétales. Extrait d'un mémoire lu à l'Academie des Sciences le 19 février 1821 [noted in Procés-verbaux, VII, 143]”, Annales de Chimie et de Physique, 2nd ser., 7 (1821), 267, with the added parts translated as a continuation in Edinburgh Phil. J., 7 (1822), 47; most of the entire expanded version was translated as “Neue Untersuchungen uber die Gesetze, welche Man in der Vertheilung der Pflanzenformen bemerkt”, in Isis von Oken, I (1821), col. 1033.

  23. Humboldt, “Distribution”, pp. 235–236 (translated). The 1820 article, immediately following Candolle's “Géographie”, makes no explicit mention of Buffon's law, but does inlcude the following: ”La distribution des êtres organisés sur le globe dépend non-seulement de circonstances climatériques très-compliqués; mais aussi de causes géologiques qui nous sont entièrement inconnues, parce qu'elles ont rapport au premier état de notre planète” (p. 429). Much the same statement as that of Humboldt (1816) is found in J. J. Virey, “Géographie naturelle des planets et des animaux”, Nouveau dictionnaire d'histoire naturelle, XIII (Paris, 1817), pp. 59–60: “Buffon and Zimmermann have observed that the quadrupeds and the birds of the tropics of the Old World — tropical Asia and Africa — are not native to America. In our time Australia has confirmed this fact in displaying terrestrial animals that all differ from those of other parts of the globe. Further researches by Cuvier have proved that the fact is also true for reptiles. If naturalists have sometimes cited boas (large, nonvenomous snakes) common to Africa and America, they appear to have confused boas with pythons — a very distinct genus. Caimans or alligators of Guiana and Florida are quite different from gavials of the Ganges and the Nile — as is true for other lizards” (translated). The reference to Zimmermann applies to mammals: E. U. W. Zimmermann, Specimen Zoologiae Geographicae, Quadrupedum Domicilia et Migrationes Sistens (Lugduni Batavorum, 1777), Geographische Geschichte des Menschen und der vierfüssigen Thiere, II (Leipzig, 1780), and Geographische Geschichte des Menschen, und der allgemein verbreiteten vierfüssigen Thiere, mit einer hiezu gehörigett zoologischen Weltcharte, III (Leipzig, 1783). The reference to Cuvier is not clear, but a summary is in Cuvier, Le régne animal, II (Paris, 1817).

  24. Buffon, Historire naturelle, XIV (Paris, 1766), pp. 316–317 (translated).

  25. Ibid., p. 373.

  26. The simplicity of Candolle's essay contrasts starkly with the diffuse complexity of what had been previously learned about plant and animal distribution, as may be appreciated in the compilation of G. R. Treviranus, Biologie, II (Göttingen, 1803). The importance of the essay was sensed not only by Lyell, but also, for a time, by Candolle's son, Alphonse de Candolle, whose general workds were important in the development of plant geography, particularly his Géographie botanique raisonée, 2 vols. (Paris, 1855).

  27. P. A. Latreille, “Introduction à la géographie générale des arachnides et des insectes”, in Mémoires du Muséum d'Histoire Naturelle, 3 (1817), 43 (translated); Reprinted in Mémoires sur divers sujets de l'histoire naturelle des insectes, de géographie ancienne et de chronologie (Paris, 1819), p. 172.

  28. Ibid., p. 44 (translated); in the reprint version, pp. 172–173.

  29. There is nothing comparable, nor even a section on plant geography, in Candolle's earlier general writings, Introduction à l'étude de la botanique (Paris, 1813), its German translation Theoretische Anfangsgründe der Botanik (Zurich, 1814), and its second edition, Théorie élémentaire de la botanique (Paris, 1819). Two chapters on distribution of plants, bearing no resemblance to Candolle's 1820 essay, were added by Sprengel to the German edition, A. P. de Candolle and K. Sprengel, Grundzüge der wissenschaftlichen Pflanzenkunde (Leipzig, 1820), translated as Elements of the Philosophy of Plants (Edinburgh, 1821). With reference to this work, and its English translation, Candolle writes: “Ouvrage entièrement rédigé par M. Sprengel, d'après ses propres idées, et auquel je suis étranger”; in Candolle, ed., Mémoires, p. 511. No chapter on plant distribution was added to the third edition, published posthumously by Candolle's son, A. de Candolle, Théorie élémentaire de la botanique par Aug. Pyr. de Candolle (Paris, 1844). But a revised version of the 1820 essay appears in his son's general text, A. de Candolle, Introduction à l'étude de la botanique (Brussels, 1837). Candolle the elder commented: “J'espérais donner un traité de géographie botanique ... Mon fils ayant mis un intéret spécial à cette branche de la botanique ... j'ai abandonné un travail pour lequel le temps et la force me manquaient, et j'ai lieu d'espérer qu'il le mènera à bien”; in Candolle, ed., Mémoires, pp. 394–395.

  30. Candolle, “Géographie”, p. 383 (translated).

  31. As noted by Egerton (“Studies”, p. 231; for Lyell, see Richardson, “Development”, pp.112–114), the concepts of station and habitation are important in Candolle's view, for they define two different sciences, which persist into the modern ear. Candolle attributes the concepts to Linnaeus: “Linnaeus est le premier qui ait pensé à les [habitations] indiquer dans les ouvrages généraux; il est le premier qui ait donné et le précepte et le modèle de la manière de rédiger les Flores; il est le premier surtout qui ait distingué avec soin les habitations, c'est-à-dire les pays dans lesquels les plantes croissent, et les stations, c'est-à-dire la nature particulière des localités dans lesquelles elles ont coutume de se développer. C'est donc de Linnaeus que sont réellement sorties les premières idées de géographie botanique” (Candolle, “Géographie”, p. 359). And again: “Linné l'a [habitation] introduite le premier dans les ouvrages généraux et en distinguant les stations et les habitations, il a indiqué (peut-etre sans le savoir) la base de la géographie botanique; cette science a commencé à naitre peu de temps après la liaison intime de la Botanique et de la physiologie sous la bannière de la méthode naturelle; comme nous en avons tracé ailleurs (Dict. des Sc. nat., art. Géographie Botanique) les progrès détaillés, nous nous bornons à faire remarquer ce fait historique digne d'attention” A.P. de Candolle, “Phytologie ou botanique”, Dictionnaire classique histoire naturelle, XII [Paris, 1828], p. 484. Candolle gives no further indication of what he means by “précepte” and “modèle”. A statement about “loca plantarum natalie” appears in paragraph 334, under “XI. Abumbrationes”, of Linnaeus's, Fundamenta Botanica (Amsterdam, 1736, and later editions), which is expanded to include a statement about “habitationes plantarum” and a list of stations (term not used) in the paragraph of the same number of his Philosophia Botanica (Stockholm, 1751, and later editions). Twenty-four stations are defined in the Linnean dissertation “Stationes Plantarum” in Amoenitates Academicae, IV 4 (Holm, 1759). These pedagogical writings perhaps constitute the “précepte” of Candolle. As for the “modèle”, that is probably found in Linnaeus's floristic and systematic works, in which for each species Linnaeus customarily gives a short statement pertaining to habitation and station, but without explicitly distinguishing between them. An example, for Salicornia europaea: “Habitat in Europae litoribus maritimis”, in Species Plantarum (Holm, 1753), p. 3. The Linnean model was followed by other botanists in their floristic and systematic works. The terms “station” and “habitation” are used in a sense different from Candolle's by J. C. Fabricius, Philosophia Entomologica (Hamburg, 1778), pp. 154–155. In combination they are used by Candolle in his Introduction (1813), p. 423, Théorie élémentaire (1819), p. 462, and Théorie élémentaire (1844), p. 383.

  32. Lyell, Principles, p. 69.

  33. Wallace, Geographical Distribution, p. 4.

  34. Candolle, “Géographie”, p. 383 (translated).

  35. Ibid., p. 392.

  36. Ibid., p. 393.

  37. Ibid., pp. 393–400.

  38. Ibid., p. 401 (translated).

  39. Ibid., p. 402: “Il ne seroit peut-être pas difficile de trouver deux points dans les Etats-Unis et l'Europe, ou dans l'Amérique et l'Afrique équinoxiale, qui présentent toutes les mêmes circonstances, savoir, une même température, une même hauteur, un même sol, une dose égale d'humidité; cependant, presque tous, peut-être tous les végétaux seroient différens dans ces deux localités semblables: on pourrait bien trouver une certaine analogie d'aspect et même de structure entre les plantes de ces deux localités supposées; mais ce seroient en général des espèces différentes. Il semble donc que d'autres circonstances que celles qui déterminent aujourd'hui les stations, ont influé sur les habitations”. Egerton apparently misread this passage, commenting as follows: “The greatest mystery in biogeography was why species are found where they are but not in similar habitats elsewhere. De Candolle had replied to this question that environmental factors are never exactly the same in different places” (“Studies”, pp. 233–234). Actually, Candolle's 1820 statement is an expanded version of one made eleven years before: “ainsi, par exemple, quoiqu'il fûl sans doute possible de trouver dans l'Amérique septentrionale certains points tout-à-fait semblables à l'Europe par la nature du sol, sa hauteur, sa température, on n'y trouveroit pas [“indépendamment des modifications que l'homme y a apportées”] une seule des plantes qu'on trouveroit dans le lieu correspondant d'Europe. La position géographique de ces deux pays a donc influé sur le choix des végétaux qui croissent spotanément dans chacun d'eux” (“Géographie agricole” [1809], p. 356).

  40. Candolle, “Géographie”, p. 403 (translated).

  41. Ibid. (translated).

  42. Ibid., p. 410 (translated).

  43. Ibid., (translated).

  44. Lyell, Principles, p. 87.

  45. Candolle, “Géographie”, pp. 411–412 (translated). A few earlier authors had proposed schemes of biogeographical regions, but none was as lengthy a list as Candolle's.

  46. Ibid., p. 412 (translated).

  47. Ibid., and what may be the first use of the term “endemic” in biogeography: “je les appellerai, par analogie avec le langage médical, genres endémiques”

  48. Ibid., p. 413.

  49. Ibid., pp. 415–416 (translated).

  50. Ibid., p. 416: “centres d'où les plantes se seroient dispersées de tous cotés”. This concept was advocated by C. L. Willdenow, Grundriss der Kräuterkunde zu Vorlesungen entworfen (Berlin, 1792, 1798, 1802, 1805, 1810). Two editions were translated as Principles of Botany and of Vegetable Physiology (Edinburgh, 1805, 1811).

  51. Candolle, “Géographie”, p. 416: “Dans cette idée, les parties primitives du globe devroient être les centres des régions; mais, outre qu'il est difficile de reconnoître des traces de cette dispersion, il est très-douteux que les espèces de plantes qui végètent aujourd'hui soient les mêmes que celles qui ont dû exister avant les terrains secondaires, et dont nous trouvons des empreintes ou des débris dans ces terrains. Cette étude curieuse, commencée il y a peu de temps, au moins avec quelque exactitude, par M. de Sternberg, et que M. Adolphe Brongniart, tout jeune qu'il est, paroît déjà destiné à perfectionner; cette étude, dis-je, semble indiquer que nos espèces végétales sont différentes des espèces antédiluviennes, et que par conséquent il y a eu développement d'une nouvelle végétation depuis la formation des terrains secondaires”.

  52. Ibid., p. 417 (translated).

  53. Ibid., p. 419.

  54. In von Hofsten's terminology, the law is part of the “Diskontinuitätsproblem” which in his view, no doubt correct, has an older history than is presented here.

  55. The references to birds of Humboldt and Virey, quoted above, are probably based on Buffon's nine volumes on birds (1770–1783), which form vol. 16–24 of his Histoire naturelle. Apparently Buffon believed that birds were an exception to his law: “il est bien difficile, pour ne pas dire impossible, de reconnoître les oiseaux propres et naturels à chaque continent, et que la plupart doivent se trouver également dans tous deux” (Histoire naturelle, XVI [1770], p. xii). A consideration of bird distribution more in accord with Buffon's law is in D.F. Tiedeman, Anatomie und Naturgeschichte der Vögel, II (Heidelberg, 1814).

  56. Latreille, “Introduction”.

  57. Lyell, Principles, p. 71: “Decandolle has enumerated twenty great botanical provinces inhabited by indigenous or aboriginal plants; and although many of these contain a variety of species which are common to several others, and sometimes to places very remote, yet the lines of demarcation are, upon the whole, astonishingly well defined. Nor is it likely that the bearing of the evidence on which these general views are founded will ever be materially affected, since they are already confirmed by the examination of seventy or eighty thousand species of plants”.

  58. Ibid., pp. 142, 145, 156, 179.

  59. Ibid., pp. 84, 128–130, 141, 156, 166, 168–170, 175–176, 179.

  60. Ibid., pp. 124, 179, 182.

  61. Ibid., pp. 124, 182.

  62. Ibid., pp. 124, 126, 156, 166, 170, 176.

  63. Ibid., pp. 73, 81, 99, 110, 125.

  64. Kinch, “Assessment”, pp. 67–69, argues that “creation” for Lyell means creation according to natural law.

  65. Darwin, Origin, p. 350.

  66. Ibid., p. 408.

  67. Crucial to the Darwin-Wallace tradition is the idea that, if continental drift occurred, it occurred earlier than the history of the recent distributions of plant and animal life. Drift theory conflicts with this tradition only if drift develops simulcaneously with, and therefore directly causes, some significant part of the recent distributions. The conflict is between different causal principles.

  68. Mayr, “Fauna” (1976) p. 553; 1965 version, p. 474.

  69. By the time of Wallace, Candolle's original list of regions, as augmented by his son Alphonse, had grown from 20 to 27, with mention of several others (Monographie des campanulées [Paris, 1830]), and then from 27 to 45 (Introduction, 1837). In apparently his last consideration of the problem, Candolle the elder listed 40 regions exhibited by a particular group of plants, A. P. de Candolle, Statistique de la famille des composées (Paris, 1838). By 1855 Candolle the younger rejected such schemes of regions: “Je tiens donc les divisions du globe par régions, proposées jusqu'à présent, pour des systèmes artificiels ... Elles ont nui à la science” (Géographie, II, 1304–1305). Later he states, with reference to his Géographie: “ouvrage du reste complètement différent de celui auquel mon père pensait, car les documents étaient devenus plus nombreus, et mes idées s'étaient singulièrement éloignées de celles qui régnaient dans la science depuis vingt ans” (Candolle, ed., Mémoires, p. 395).

  70. Wallace, Geographical Distribution, p. 54.

  71. Ibid., pp. 57–58. See also M. Fichman, “Wallace: Zoogeography and the Problem of Land Bridges,” J. Hist. Bio., 10 (1977), 45.

  72. A. R.Wallace, “What Are Zoological Regions?” Nature (London) 49 (1894), 612.

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  73. Ibid.

  74. P. L.Sclater, “On the General Geographical Distribution of the Members of the Class Aves,” J. Proc. Linnean Soc. London (Zool.) 2 (1858), 130.

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  75. Ibid., p. 131.

  76. P. L.Sclater, “On the General Geographical Distribution of the Members of the Class Aves,” J. Proc. Linnean Soc. London (Zool.) 2 (1858) pp. 131–132.

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  77. W. D.Matthew, “Climate and Evolution” Ann. N.Y. Acad. Sci., 24 (1915), 171; revised edition, Special Publication of the New York Academy of Sciences (1939).

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  78. G. G. Simpson, Evolution and Geography (Eugene, Ore., 1953); The Geography of Evolution (Philadelphia and New York, 1965).

  79. P. J. Darlington, Jr., Zoogeography: The Geographical Distribution of Animals (New York, 1957), Biogeography of the Southern End of the World (Cambridge, Mass., 1965).

  80. P. J.DarlingtonJr., “Darwin and Zoogeography,” Proc. Amer. Phil. Soc., 103 (1959), 307.

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  81. Darlington, Zoogeography, p. 422.

  82. Ibid.

  83. A. R.Wallace, “Letter from Mr. Wallace [to P. L. Sclater] concerning the Geographical Distribution of Birds,” Ibis, 1 (1859), 449.

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  84. L. Croizat, Manual of Phytogeography (The Hague, 1952); Panbiogeography, 3 vols. (Caracas, 1958); Principia Botanica, 2 vols. (Caracas, 1960); Space, Time, Form: The Biological Synthesis (Caracas, 1962); L. Croizat-Chaley, “Biogeografía Analítica y Sintética (Panbiogeografía) de las Américas,” Biblioteca de la Academia de Ciencias Físicas, Matemáticas y Naturales vols. 15–16 (Caracas, 1976). A complete bibliography of Croizat's writings on biogeography, during the period 1947–1973, is in G. Nelson, “Comments on Leon Croizat's Biogeography,” Syst. Zool. 22 (1973), 312. Relevant comments are in L. Croizat, G. Nelson, and D. E. Rosen, “Centeŕs of Origin and Related Concepts,” Syst. Zool., 23 (1974), 265.

  85. W. Hennig, Phylogenetic Systematics (Urbana, 1966). Although Hennig does not refer to Croizat, the nature of the relation between the ideas of these two writers is best appreciated, so far, in D. E. Rosen, “A Vicariance Model of Caribbean Biogeography,” Syst. Zool., 24 (1975), 431. and “Vicariant Patterns and Historical Explanation in Biogeography” Syst. Zool., 27 (1978), 159.

  86. The concept of phenetic relationship is extensively discussed in R.R. Sokal and P. H. A. Sneath, Principles of Numerical Taxonomy (San Francisco and London, 1963), and in P. H. A. Sneath and R. R. Sokal, Numerical Taxonomy (San Francisco, 1973).

  87. By “Cladistically,” I mean that Buffon's law, in a particular case, involving areas of endemism A, B, C, etc., may be stated in the form of a cladogram, which serves as a statement of the interrelationships of the areas, and which is, or may be, true for all of the organisms endemic in the areas — for all of the species that exemplify the law. Croizat does not refer to “cladograms,” but the cladistic sense of relationship is implied by his approach; for example: “The patterns of geographic distribution of plants and animals — whatever their ‘means’ [of dispersal] — are absolutely congruent, as a fact of nature, within a minimum of fundamental tracks and centers” (Space, p. 712).

  88. For example: “‘Means of dispersal’ are hardly worthy of discussion, much as the contrary has always been taken for certain during the last century” (Croizat, Space, p. 213). “At least nine tenths of the literature of zoogeography and phytogeography from 1859 to 1962 has had for its purpose hopefully to clarify the nexus between geographic distribution ... and migration and its ‘means’ ... ” (ibid., p. 210). “I have no wish to oppose academic notions of ‘means of dispersal’ because I dislike what some say of ... [them] at this hour. I intend to destroy these notions because they positively interfere with the advance of knowledge” (ibid., p. vi). “Tectonics, not stratigraphy, are the proper standard ... by which to dovetail geology and biogeography” (ibid., p. 259). “Stratigraphy — rightfully a major preoccupation with the paleontologist — is of rather secondary interest for the biogeographer. What does primarily count for him are tectonics (structural geology ...), that is, major features of the earth past and present fit to influence the course of life by fixing biogeographic boundaries, primary and secondary centers of form-making, etc.” (ibid., pp. 115–116). “Earth and life evolved together: the contents are themselves modified and shifted whenever the container alters” (ibid., p. 46). “The deployment of life is but biological function of the geological history of the earth” (ibid., pp. 48–49). “[The distribution of plants and animals] ... was cast on maps other than the ones of the current world, and it is these maps, not the one of this hour, which dominate in the equations of life. The map of this day keeps the records; the maps of the past rationally account for them” (ibid., p. 85).

  89. I am unaware that Croizat has made a succinct statement of this sort, which may be my own interpretation. The concept nevertheless appears, it seems to me, in many statements by Croizat. For example: “The Atlantic Ocean (of the current map) was ‘crossed’ at many points, as we have seen already, but ... this body of waters itself marks a major axis of ... [distribution] north to south, west to east (and the other way around) as if land all over. What better ‘landbridge’ could be ever found anywhere than an ocean behaving in this manner?” (ibid., p. 56). ‘A formerly complex set-up of geography was shattered in Atlantic ranges of which the Mid-Atlantic Ridge does somehow mark today the backbone” (ibid., p. 42). “There is ... a general process of evolution over space, in time, by form which has for its effect that the same genus, even species, is today located on both sides of the Atlantic, Pacific, Indian oceans, etc., etc.” (ibid., p. 32).

  90. W. George, Biologist Philosopher: A Study of the Life and Writings of Alfred Russel Wallace (London, 1964), pp. 154–155.

  91. A point of confusion in recognizing areas of endemism was created early in the history of plant geography by J. F. Schouw, Grundzüge einer allgemeine Pflanzengeographie (Berlin, 1823). To qualify as botanical regions according to Schouw, areas must exhibit certain qualities beyond the elder Candolle's concept: “des espaces quelconques qui, si l'on fait exception des espèces introduites, offrent un certain nombre de plantes qui leur sont particulières et qu'on pourrait nommer véritablement aborigènes” (see note 43). Briefly, Schouw's concept requires, with some qualifications, that an area have at least one-half of its species endemic, one-quarter of its genera endemic, and have also endemic families, in Grundzüge, p. 505, and in an essay in Danish, part of which was translated into English as “Schow's Essay on Botanical Geography. Copenhagen, 1823,” Edinburgh J. Sci., 4 (1826), 161; and the English translated into French as “Essai sur la géographie botanique, par Schouw,” Bulletin de la Société de Géographie, 5 (1826), 127. The first of Schouw's requirements was soon adopted, without attribution, by the younger Candolle (Monographie des campanulées, p. 70), and the others were later discussed at some length (Introduction [1837], pp. 387–389), with added consideration of physical geography, climate, natural boundaries, and average size. The elder Candolle later replied to Schouw (and implicity to the younger Candolle) with an expansion of his original point of view (Statistique, pp. 8–15). The younger Candolle later explained why he rejected his father's simple concept in favor of Schouw's, which he felt was better but still unsatisfactory, and then declared that the whole matter was beside the point (Géographie, pp. 1298–1305).

  92. J. D. Hooker, “On Geographical Distribution,” in Raport of the Fifty-First Meeting of the British Association for the Advancement of Science (London, 1882), pp. 737–738. The contrast in the views of the elder and younger Candolles is repeated by the elder and younger Hookers. The plan of Candolle's 1820 essay is adopted in the general exposition by W. J. Hooker, “Geography in Relation to the Distribution of Plants,” in H. Murray (ed.), An Encyclopaedia of Geography, 1 (London, 1834), 227, and (Philadelphia, 1837), 236.

  93. Ibid., p. 733. Hooker and many other early botanists were impressed with Humboldt's “Arithmeticae botanices,” which in Hooker's words “consist in determining the proportion which the species of certain large families or groups of families bear to the whole number of species composing the floras in advancing from the Equator to the Poles, and in ascending mountains ... The importance of this method of analysing the vegetation of a country in researches in geographical botany is obvious, for it affords the most instructive method of setting forth the relations that exist between a flora and its geographical position and climatal conditions.” Such matters were thought reducible to laws (for example, by the younger Candolle in Introduction [1837], pp. 374–378), who later reviewed the whole matter and, seemingly, disposed of it (Géographie [1855]). The reference to Forbes is primarily to E. Forbes, “On the Connexion between the Distribution of the Existing Fauna and Flora of the British Isles,” in Memoirs of the Geological Survey of Great Britain, 1 (1846), 336.

  94. Wallace, “Regions,” p. 612.

  95. Richardson, “Development,” pp. 193–194. His reference is to Darlington's Zoogeography: “an excellent introduction to the modern science of zoogeography. The reader wishing to make his own comparisons of Darwin's treatment of the problems of geographical distributions, with those of recent scholars, will find this book a very useful reference” (p. 233). A complementary evaluation of the same book is in L. Brundin, “Transantarctic Relationships and Their Significance as Evidenced by Chironomid Midges,” in Kungl. Svenska Vetenskapsakademiens Handlingar, 4th ser., 11 (1966).

  96. Fichman, “Wallace,” p. 63.

  97. For example, Aubréville comments on the work of Croizat: En dépit de son oeuvre impressionnante, Léon Croizat est peu connu. La plupart de biogéographes semblent l'ignorer — y compris les biogéographes américains et anglais pour lesquels ne se présente en l'occurrence aucune difficulté de lecture —; ils ne citent généralement pas ses ouvrages dans leur bibliographie. On croirait qu'une consigne de silence empêche la diffusion de ces oeuvres, et que la méconnaissance de l'auteur soit voulue; A. Aubréville, “A propos de l'”Introduction raisonée à la biogéographie de l'Afrique” de Léon Croizat,” Adansonia, 2nd ser., 9 (1969), 489.

  98. L. Germain, La vie des animaux à la surface des continents (Paris, 1924), pp. 2–5.

  99. T.H. Huxley (in a letter to Hooker, Sept. 5, 1858), in L. Huxley, Life and Letters of Thomas Henry Huxley, I (New York, 1900), 171.

  100. T. Thiselton-Dyer, “Distribution of Vegetable Life,” in A. R. Wallace and W. T. Thiselton-Dyer, “Distribution,” in Encyclopedia Britannica, 9th ed., VII (New York, 1878), p. 267; A.R. Wallace and W. T. Thiselton-Dyer, “The Distribution of Life, Animal and Vegetable, in Space and Time,” in Humboldt Library of Popular Science Literature, 6, no. 64 (1885), p. 227.

  101. W. Thiselton-Dyer, “Geographical Distribution of Plants,” in A. C. Seward, ed., Darwin and Modern Science (Cambridge, 1909), p. 308.

  102. Ibid., p. 316.

  103. “Improbable dispersal” refers to what Darwin termed dispersal by “occasional means of transport” — a concept that played an original, but relatively minor, role in Darwin's arguments (see e.g., Origin, p. 410). The concept was emphasized more and more by later writers such as Wallace, Matthew (“Climate,” pp. 206–209), and especially Simpson (“Probabilities of Dispersal in Geologic Time,” Bull. Amer. Mu. Nat. Hist., 99 [1952], 163). The end result was that, given enough time, the most improbable dispersal becomes not only probable but certain: “If the probability that some member of a population will cross a barrier is .000001 in any one year, in a large population this means that the probability for any one designated individual is almost infinitesimally small, so much so that it would seem absolutely impossible to even the best qualified observer in the field. Yet during the course of a million years the event would be probable, p=.63, again. In the course of 10 million years the event would become so extremely probable as to be, for most practical purposes, certain, p=.99995” (Simpson, “Probabilities,” p. 171).

  104. Darwin, Origin, pp. 350–351.

  105. Ibid., p. 382.

  106. R. H. MacArthur and E. O. Wilson, The Theory of Island Biogeography (Princeton, 1967).

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    Gareth Nelson

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Nelson, G. From Candolle to croizat: Comments on the history of biogeography. Journal of the History of Biology 11, 269–305 (1978). https://doi.org/10.1007/BF00389302

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