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Darwin Correspondence Project

To Friedrich Ludwig   29 May 1878

Down | Beckenham, Kent (&c).

May 29th. 1878.

Dear Sir.

I thank you sincerely for the trouble which you have taken in sending me so long & interesting a letter, together with the specimens.1 Gradations are always very valuable, & you have been remarkably successful in discovering the stages by which the Plantago has become gyno-diœcious— Your view of its origin from being proterogynous, seems to me very probable especially as the females are generally the later flowering plants.2 If you can prove the reverse case with Thymus your view will manifestly be rendered still more probable— I have never felt satisfied with H. Müller’s view, though he is so careful & admirable an observer.3 It is more than 17 years since I attended to Plantago; & when nothing had been published on the subject, & in consequence I omitted to attend to several points,—& now after so long an interval, I cannot pretend to say to which of your forms, the English one belongs; I well remember that the anther of the females contained a good deal pollen, though not one sound grain.—4

Once again thanking you for your kindness, I remain with much respect— Dear Sir | Yours faithfully. | Ch. Darwin.

P.S. | Delpino5 is Professor of Botany in Genoa Italy; I have always found him a most obliging correspondent.


Ludwig’s letter to CD has not been found, but in Forms of flowers 2d ed., p. ix, CD wrote: I have received an additional account of Plantago lanceolata being gyno-diœcious in England; and Dr. F. Ludwig of Greiz has sent me a description of no less than five forms of this plant which graduate into one another; the intermediate forms being comparatively rare, whilst the hermaphrodite form is the commonest. Plantago lanceolata is ribwort plantain.
Ludwig later published a paper, ‘Ueber die Blütenformen von Plantago lanceolata L., und die Erscheinung der Gynodiöcie’ (On the flower forms of Plantago lanceolata L., and the appearance of gynodioecy; Ludwig 1879), in which he discussed the stages in the development of gynodioecy, or the existence of both hermaphrodite and female flowers. Proterogynous (now protogynous) flowers are hermaphrodite flowers in which the female sexual parts (carpels) mature before the male ones (stamens).
Ludwig noted that another gynodioecious species, Thymus serpyllum (Breckland thyme), differed from Plantago lanceolata in the ratio of hermaphrodite to female flowers during its flowering period. In T. serpyllum, whose hermaphrodite flowers are protandrous (i.e. stamens mature before carpels), female flowers predominated early in the flowering period, while hermaphrodite flowers became more frequent later. The opposite was the case in P. lanceolata. (Ludwig 1879, p. 448.) Hermann Müller had suggested that since insects tended to visit more conspicuous flowers first (where they were dusted with pollen), less conspicuous flowers of the same species, which were visited only later, might have gradually lost functional stamens and been converted to females (H. Müller 1873, pp. 319, 326; Forms of flowers, p. 304).
CD’s notes on dichogamy and dimorphism in Plantago lanceolata, dated 28 April 1863, are in DAR 109: A27–8.
Federico Delpino.


Forms of flowers 2d ed.: The different forms of flowers on plants of the same species. 2d edition. By Charles Darwin. London: John Murray. 1880.

Forms of flowers: The different forms of flowers on plants of the same species. By Charles Darwin. London: John Murray. 1877.

Ludwig, Friedrich. 1879. Ueber die Blütenformen von Plantago lanceolata L. und die Erscheinung der Gynodiöcie. Zeitschrift für die gesammten Naturwissenschaften 3d ser. 4: 441–9.

Müller, Hermann. 1873. Die Befruchtung der Blumen durch Insekten und die gegenseitigen Anpassungen beider. Ein Beitrag zur Erkenntniss des ursächlichen Zusammenhanges in der organischen Natur. Leipzig: Wilhelm Engelmann.


Thanks FL for the Plantago specimens. FL’s view of the stages by which the plant has become gynodioecious seems very probable.

Letter details

Letter no.
Charles Robert Darwin
Friedrich Ludwig
Sent from
Source of text
DAR 146: 137
Physical description

Please cite as

Darwin Correspondence Project, “Letter no. 11534,” accessed on 27 September 2021,