From statements made by Vaucher I was led to think that Soldanella alpina was heterostyled, but it is impossible that Kerner, who has closely studied this plant, could have overlooked the fact. So again from other statements it appeared probable that Pyrola might be heterostyled, but H. Muller examined for me two species in North Germany, and found this not to be the case.



The oxlip a hybrid naturally produced between Primula veris and vulgaris. The differences in structure and function between the two parent-species. Effects of crossing long-styled and short-styled oxlips with one another and with the two forms of both parent-species. Character of the offspring from oxlips artificially self-fertilised and cross- fertilised in a state of nature. Primula elatior shown to be a distinct species. Hybrids between other heterostyled species of Primula. Supplementary note on spontaneously produced hybrids in the genus Verbascum.

The various species of Primula have produced in a state of nature throughout Europe an extraordinary number of hybrid forms. For instance, Professor Kerner has found no less than twenty-five such forms in the Alps. (2/1. "Die Primulaceen-Bastarten" 'Oesterr. Botanische Zeitschrift' Jahr 1875 Numbers 3, 4 and 5. See also Godron on hybrid Primulas in 'Bull. Soc. Bot. de France' tome 10 1853 page 178. Also in 'Revue des Sciences Nat.' 1875 page 331.) The frequent occurrence of hybrids in this genus no doubt has been favoured by most of the species being heterostyled, and consequently requiring cross-fertilisation by insects; yet in some other genera, species which are not heterostyled and which in some respects appear not well adapted for hybrid-fertilisation, have likewise been largely hybridised. In certain districts of England, the common oxlip--a hybrid between the cowslip (P. veris, vel officinalis) and the primrose (P. vulgaris, vel acaulis)--is frequently found, and it occurs occasionally almost everywhere. Owing to the frequency of this intermediate hybrid form, and to the existence of the Bardfield oxlip (P. elatior), which resembles to a certain extent the common oxlip, the claim of the three forms to rank as distinct species has been discussed oftener and at greater length than that of almost any other plant. Linnaeus considered P. veris, vulgaris and elatior to be varieties of the same species, as do some distinguished botanists at the present day; whilst others who have carefully studied these plants do not doubt that they are distinct species. The following observations prove, I think, that the latter view is correct; and they further show that the common oxlip is a hybrid between P. veris and vulgaris.

The cowslip differs so conspicuously in general appearance from the primrose, that nothing need here be said with respect to their external characters. (2/2. The Reverend W.A. Leighton has pointed out certain differences in the form of the capsules and seed in 'Annals and Magazine of Natural History' 2nd series volume 2 1848 page 164.) But some less obvious differences deserve notice. As both species are heterostyled, their complete fertilisation depends on insects. The cowslip is habitually visited during the day by the larger humble-bees (namely Bombus muscorum and hortorum), and at night by moths, as I have seen in the case of Cucullia. The primrose is never visited (and I speak after many years' observation) by the larger humble-bees, and only rarely by the smaller kinds; hence its fertilisation must depend almost exclusively on moths. There is nothing in the structure of the flowers of the two plants which can determine the visits of such widely different insects. But they emit a different odour, and perhaps their nectar may have a different taste. Both the long-styled and short-styled forms of the primrose, when legitimately and naturally fertilised, yield on an average many more seeds per capsule than the cowslip, namely, in the proportion of 100 to 55. When illegitimately fertilised they are likewise more fertile than the two forms of the cowslip, as shown by the larger proportion of their flowers which set capsules, and by the larger average number of seeds which the capsules contain.

Charles Darwin

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