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Chapter 2

HYBRID PRIMULAS.


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. The difference also between the number of seeds produced by the long-styled and short-styled flowers of the primrose, when both are illegitimately fertilised, is greater than that between the number produced under similar circumstances by the two forms of the cowslip. The long-styled flowers of the primrose when protected from the access of all insects, except such minute ones as Thrips, yield a considerable number of capsules containing on an average 19.2 seeds per capsule; whereas 18 plants of the long-styled cowslip similarly treated did not yield a single seed.

The primrose, as every one knows, flowers a little earlier in the spring than the cowslip, and inhabits slightly different stations and districts. The primrose generally grows on banks or in woods, whilst the cowslip is found in more open places. The geographical range of the two forms is different. Dr. Bromfield remarks that "the primrose is absent from all the interior region of northern Europe, where the cowslip is indigenous." (2/3. 'Phytologist' volume 3 page 694.) In Norway, however, both plants range to the same degree of north latitude. (2/4. H. Lecoq 'Geograph. Bot. de l'Europe' tome 8 1858 pages 141, 144. See also 'Annals and Magazine of Natural History' 9 1842 pages 156, 515. Also Boreau 'Flore du centre de la France' 1840 tome 2 page 376. With respect to the rarity of P. veris in western Scotland, see H.C. Watson 'Cybele Britannica' 2 page 293.)

The cowslip and primrose, when intercrossed, behave like distinct species, for they are far from being mutually fertile. Gartner crossed 27 flowers of P. vulgaris with pollen of P. veris, and obtained 16 capsules; but these did not contain any good seed. (2/5. 'Bastarderzeugung' 1849 page 721.) He also crossed 21 flowers of P. veris with pollen of P. vulgaris; and now he got only five capsules, containing seed in a still less perfect condition. Gartner knew nothing about heterostylism; and his complete failure may perhaps be accounted for by his having crossed together the same forms of the cowslip and primrose; for such crosses would have been of an illegitimate as well as of a hybrid nature, and this would have increased their sterility. My trials were rather more fortunate. Twenty-one flowers, consisting of both forms of the cowslip and primrose, were intercrossed legitimately, and yielded seven capsules (i.e. 33 per cent), containing on an average 42 seeds; some of these seeds, however, were so poor that they probably would not have germinated. Twenty-one flowers on the same cowslip and primrose plants were also intercrossed illegitimately, and they likewise yielded seven capsules (or 33 per cent), but these contained on an average only 13 good and bad seeds. I should, however, state that some of the above flowers of the primrose were fertilised with pollen from the polyanthus, which is certainly a variety of the cowslip, as may be inferred from the perfect fertility inter se of the crossed offspring from these two plants. (2/6. Mr. Scott has discussed the nature of the polyanthus ('Proceedings of the Linnean Society' 8 Botany 1864 page 103), and arrives at a different conclusion; but I do not think that his experiments were sufficiently numerous. The degree of infertility of a cross is liable to much fluctuation. Pollen from the cowslip at first appears rather more efficient on the primrose than that of the polyanthus; for 12 flowers of both forms of the primrose, fertilised legitimately and illegitimately with pollen of the cowslip gave five capsules, containing on an average 32.4 seeds; whilst 18 flowers similarly fertilised by polyanthus-pollen yielded only five capsules, containing only 22.6 seeds. On the other hand, the seeds produced by the polyanthus-pollen were much the finest of the whole lot, and were the only ones which germinated.) To show how sterile these hybrid unions were I may remind the reader that 90 per cent of the flowers of the primrose fertilised legitimately with primrose-pollen yielded capsules, containing on an average 66 seeds; and that 54 per cent of the flowers fertilised illegitimately yielded capsules containing on an average 35.5 seeds per capsule. The primrose, especially the short-styled form, when fertilised by the cowslip, is less sterile, as Gartner likewise observed, than is the cowslip when fertilised by the primrose. The above experiments also show that a cross between the same forms of the primrose and cowslip is much more sterile than that between different forms of these two species.

The seeds from the several foregoing crosses were sown, but none germinated except those from the short-styled primrose fertilised with pollen of the polyanthus; and these seeds were the finest of the whole lot. I thus raised six plants, and compared them with a group of wild oxlips which I had transplanted into my garden. One of these wild oxlips produced slightly larger flowers than the others, and this one was identical in every character (in foliage, flower- peduncle, and flowers) with my six plants, excepting that the flowers of the latter were tinged of a dingy red colour, from being descended from the polyanthus.

We thus see that the cowslip and primrose cannot be crossed either way except with considerable difficulty, that they differ conspicuously in external appearance, that they differ in various physiological characters, that they inhabit slightly different stations and range differently. Hence those botanists who rank these plants as varieties ought to be able to prove that they are not as well fixed in character as are most species; and the evidence in favour of such instability of character appears at first sight very strong. It rests, first, on statements made by several competent observers that they have raised cowslips, primroses, and oxlips from seeds of the same plant; and, secondly, on the frequent occurrence in a state of nature of plants presenting every intermediate gradation between the cowslip and primrose.

The first statement, however, is of little value; for, heterostylism not being formerly understood, the seed-bearing plants were in no instance protected from the visits of insects (2/7. One author states in the 'Phytologist' volume 3 page 703 that he covered with bell-glasses some cowslips, primroses, etc., on which he experimented. He specifies all the details of his experiment, but does not say that he artificially fertilised his plants; yet he obtained an abundance of seed, which is simply impossible. Hence there must have been some strange error in these experiments, which may be passed over as valueless.); and there would be almost as much risk of an isolated cowslip, or of several cowslips if consisting of the same form, being crossed by a neighbouring primrose and producing oxlips, as of one sex of a dioecious plant, under similar circumstances, being crossed by the opposite sex of an allied and neighbouring species. Mr. H.C. Watson, a critical and most careful observer, made many experiments by sowing the seeds of cowslips and of various kinds of oxlips, and arrived at the following conclusion, namely, "that seeds of a cowslip can produce cowslips and oxlips, and that seeds of an oxlip can produce cowslips, oxlips, and primroses." (2/8. 'Phytologist' 2 pages 217, 852; 3 page 43.) This conclusion harmonises perfectly with the view that in all cases, when such results have been obtained, the unprotected cowslips have been crossed by primroses, and the unprotected oxlips by either cowslips or primroses; for in this latter case we might expect, by the aid of reversion, which notoriously comes into powerful action with hybrids, that the two parent-forms in appearance pure, as well as many intermediate gradations, would be occasionally produced. Nevertheless the two following statements offer considerable difficulty. The Reverend Professor Henslow raised from seeds of a cowslip growing in his garden, various kinds of oxlips and one perfect primrose; but a statement in the same paper perhaps throws light on this anomalous result. (2/9. Loudon's 'Magazine of Natural History' 3 1830 page 409.) Professor Henslow had previously transplanted into his garden a cowslip, which completely changed its appearance during the following year, and now resembled an oxlip. Next year again it changed its character, and produced, in addition to the ordinary umbels, a few single- flowered scapes, bearing flowers somewhat smaller and more deeply coloured than those of the common primrose. From what I have myself observed with oxlips, I cannot doubt that this plant was an oxlip in a highly variable condition, almost like that of the famous Cytisus adami. This presumed oxlip was propagated by offsets, which were planted in different parts of the garden; and if Professor Henslow took by mistake seeds from one of these plants, especially if it had been crossed by a primrose, the result would be quite intelligible. Another case is still more difficult to understand: Dr. Herbert raised, from the seeds of a highly cultivated red cowslip, cowslips, oxlips of various kinds, and a primrose. (2/10. 'Transactions of the Horticultural Society' 4 page 19.) This case, if accurately recorded, which I much doubt, is explicable only on the improbable assumption that the red cowslip was not of pure parentage. With species and varieties of many kinds, when intercrossed, one is sometimes strongly prepotent over the other; and instances are known of a variety crossed by another, producing offspring which in certain characters, as in colour, hairiness, etc., have proved identical with the pollen-bearing parent, and quite dissimilar to the mother-plant (2/11. I have given instances in my work 'On the Variation of Animals and Plants under Domestication' chapter 15 2nd edition volume 2 page 69.); but I do not know of any instance of the offspring of a cross perfectly resembling, in a considerable number of important characters, the father alone. It is, therefore, very improbable that a pure cowslip crossed by a primrose should ever produce a primrose in appearance pure. Although the facts given by Dr. Herbert and Professor Henslow are difficult to explain, yet until it can be shown that a cowslip or a primrose, carefully protected from insects, will give birth to at least oxlips, the cases hitherto recorded have little weight in leading us to admit that the cowslip and primrose are varieties of one and the same species.

Negative evidence is of little value; but the following facts may be worth giving:--Some cowslips which had been transplanted from the fields into a shrubbery were again transplanted into highly manured land. In the following year they were protected from insects, artificially fertilised, and the seed thus procured was sown in a hotbed. The young plants were afterwards planted out, some in very rich soil, some in stiff poor clay, some in old peat, and some in pots in the greenhouse; so that these plants, 765 in number, as well as their parents, were subjected to diversified and unnatural treatment; but not one of them presented the least variation except in size--those in the peat attaining almost gigantic dimensions, and those in the clay being much dwarfed.

I do not, of course, doubt that cowslips exposed during SEVERAL successive generations to changed conditions would vary, and that this might occasionally occur in a state of nature. Moreover, from the law of analogical variation, the varieties of any one species of Primula would probably in some cases resemble other species of the genus. For instance I raised a red primrose from seed from a protected plant, and the flowers, though still resembling those of the primrose, were borne during one season in umbels on a long foot-stalk like that of a cowslip.

With regard to the second class of facts in support of the cowslip and primrose being ranked as mere varieties, namely, the well-ascertained existence in a state of nature of numerous linking forms (2/12. See an excellent article on this subject by Mr. H.C. Watson in the 'Phytologist' volume 3 page 43.):--If it can be shown that the common wild oxlip, which is intermediate in character between the cowslip and primrose, resembles in sterility and other essential respects a hybrid plant, and if it can further be shown that the oxlip, though in a high degree sterile, can be fertilised by either parent-species, thus giving rise to still finer gradational links, then the presence of such linking forms in a state of nature ceases to be an argument of any weight in favour of the cowslip and primrose being varieties, and becomes, in fact, an argument on the other side. The hybrid origin of a plant in a state of nature can be recognised by four tests: first, by its occurrence only where both presumed parent-species exist or have recently existed; and this holds good, as far as I can discover, with the oxlip; but the P. elatior of Jacq., which, as we shall presently see, constitutes a distinct species, must not be confounded with the common oxlip. Secondly, by the supposed hybrid plant being nearly intermediate in character between the two parent-species, and especially by its resembling hybrids artificially made between the same two species. Now the oxlip is intermediate in character, and resembles in every respect, except in the colour of the corolla, hybrids artificially produced between the primrose and the polyanthus, which latter is a variety of the cowslip. Thirdly, by the supposed hybrids being more or less sterile when crossed inter se: but to try this fairly two distinct plants of the same parentage, and not two flowers on the same plant, should be crossed; for many pure species are more or less sterile with pollen from the same individual plant; and in the case of hybrids from heterostyled species the opposite forms should be crossed. Fourthly and lastly, by the supposed hybrids being much more fertile when crossed with either pure parent-species than when crossed inter se, but still not as fully fertile as the parent-species.

For the sake of ascertaining the two latter points, I transplanted a group of wild oxlips into my garden. They consisted of one long-styled and three short- styled plants, which, except in the corolla of one being slightly larger, resembled each other closely. The trials which were made, and the results obtained, are shown in tables 2.14, 2.15, 2.16, 2.17 and 2.18. No less than twenty different crosses are necessary in order to ascertain fully the fertility of hybrid heterostyled plants, both inter se and with their two parent-species. In this instance 256 flowers were crossed in the course of four seasons. I may mention, as a mere curiosity, that if any one were to raise hybrids between two trimorphic heterostyled species, he would have to make 90 distinct unions in order to ascertain their fertility in all ways; and as he would have to try at least 10 flowers in each case, he would be compelled to fertilise 900 flowers and count their seeds. This would probably exhaust the patience of the most patient man.

TABLE 2.14. Crosses inter se between the two forms of the common Oxlip.

Column 1: Illegitimate union.

Short-styled oxlip, by pollen of short-styled oxlip: 20 flowers fertilised, did not produce one capsule.

Column 2: Legitimate union.

Short-styled oxlip, by pollen of long-styled oxlip: 10 flowers fertilised, did not produce one capsule.

Column 3: Illegitimate union.

Long-styled oxlip, by its own pollen: 24 flowers fertilised, produced five capsules, containing 6, 10, 20, 8, and 14 seeds. Average 11.6.

Column 4: Legitimate union.

Long-styled oxlip, by pollen of short-styled oxlip: 10 flowers fertilised, did not produce one capsule.

TABLE 2.15. Both forms of the Oxlip crossed with Pollen of both forms of the Cowslip, P. veris.

Column 1: Illegitimate union.

Short-styled oxlip, by pollen of short-styled cowslip: 18 flowers fertilised, did not produce one capsule.

Column 2: Legitimate union.

Short-styled oxlip, by pollen of long-styled cowslip: 18 flowers fertilised, produced three capsules, containing 7, 3, and 3 wretched seeds, apparently incapable of germination.

Column 3: Illegitimate union.

Long-styled oxlip, by pollen of long-styled cowslip: 11 flowers fertilised, produced one capsule, containing 13 wretched seeds.

Column 4: Legitimate union.

Long-styled oxlip, by pollen of short-styled cowslip: 5 flowers fertilised, produced two capsules, containing 21 and 28 very fine seeds.

TABLE 2.16. Both forms of the Oxlip crossed with Pollen of both forms of the Primrose, P. vulgaris.

Column 1: Illegitimate union.

Short-styled oxlip, by pollen of short-styled primrose: 34 flowers fertilised, produced two capsules, containing 5 and 12 seeds.

Column 2: Legitimate union.

Short-styled oxlip, by pollen of long-styled primrose: 26 flowers fertilised, produced six capsules, containing 16, 20, 5, 10, 19, and 24 seeds. Average 15.7. Many of the seeds very poor, some good.

Column 3: Illegitimate union.

Long-styled oxlip, by pollen of long-styled primrose: 11 flowers fertilised, produced four capsules, containing 10, 7, 5, and 6 wretched seeds. Average 7.0.

Column 4: Legitimate union.

Long-styled oxlip, by pollen of short-styled primrose: 5 flowers fertilised, produced five capsules, containing 26, 32, 23, 28, and 34 seeds. Average 28.6.

TABLE 2.17. Both forms of the Cowslip crossed with Pollen of both forms of the Oxlip.

Column 1: Illegitimate union.

Short-styled cowslip, by pollen of short-styled oxlip: 8 flowers fertilised, did not produce one capsule.

Column 2: Legitimate union.

Long-styled cowslip, by pollen of short-styled oxlip: 8 flowers fertilised, produced one capsule, containing 26 seeds.

Column 3: Illegitimate union.

Long-styled cowslip, by pollen of long-styled oxlip: 8 flowers fertilised, produced three capsules, containing 5, 6 and 14 seeds. Average 8.3.

Column 4: Legitimate union.

Short-styled cowslip, by pollen of long-styled oxlip: 8 flowers fertilised, produced 8 capsules, containing 58, 38, 31, 44, 23, 26, 37, and 66 seeds. Average 40.4.

TABLE 2.18. Both forms of the Primrose crossed with Pollen of both forms of the Oxlip.

Column 1: Illegitimate union.

Short-styled primrose, by pollen of short-styled oxlip: 8 flowers fertilised, did not produce one capsule.

Column 2: Legitimate union.

Long-styled primrose, by pollen of short-styled oxlip: 8 flowers fertilised, produced two capsules, containing 5 and 2 seeds.

Column 3: Illegitimate union.

Long-styled primrose, by pollen of long-styled oxlip: 8 flowers fertilised, produced 8 capsules, containing 15, 7, 12, 20, 22, 7, 16, and 13 seeds. Average 14.0.

Column 4: Legitimate union.

Short-styled primrose, by pollen of long-styled oxlip: 8 flowers fertilised, produced 4 capsules, containing 52, 52, 42, and 49 seeds, some good and some bad. Average 48.7.

We see in Tables 2/14 to 2/18 the number of capsules and of seeds produced, by crossing both forms of the oxlip in a legitimate and illegitimate manner with one another, and with the two forms of the primrose and cowslip. I may premise that the pollen of two of the short-styled oxlips consisted of nothing but minute aborted whitish cells; but in the third short-styled plant about one- fifth of the grains appeared in a sound condition. Hence it is not surprising that neither the short-styled nor the long-styled oxlip produced a single seed when fertilised with this pollen. Nor did the pure cowslips or primroses when illegitimately fertilised with it; but when thus legitimately fertilised they yielded a few good seeds. The female organs of the short-styled oxlips, though greatly deteriorated in power, were in a rather better condition than the male organs; for though the short-styled oxlips yielded no seed when fertilised by the long-styled oxlips, and hardly any when illegitimately fertilised by pure cowslips or primroses, yet when legitimately fertilised by these latter species, especially by the long-styled primrose, they yielded a moderate supply of good seed.

The long-styled oxlip was more fertile than the three short-styled oxlips, and about half its pollen-grains appeared sound. It bore no seed when legitimately fertilised by the short-styled oxlips; but this no doubt was due to the badness of the pollen of the latter; for when illegitimately fertilised (Table 2.14) by its own pollen it produced some good seeds, though much fewer than self- fertilised cowslips or primroses would have produced. The long-styled oxlip likewise yielded a very low average of seed, as may be seen in the third compartment of Tables 2.15 to 2.18, when illegitimately fertilised by, and when illegitimately fertilising, pure cowslips and primroses. The four corresponding legitimate unions, however, were moderately fertile, and one (namely that between a short-styled cowslip and the long-styled oxlip in Table 2.17) was nearly as fertile as if both parents had been pure. A short-styled primrose legitimately fertilised by the long-styled oxlip (Table 2.18) also yielded a moderately good average, namely 48.7 seeds; but if this short-styled primrose had been fertilised by a long-styled primrose it would have yielded an average of 65 seeds. If we take the ten legitimate unions together, and the ten illegitimate unions together, we shall find that 29 per cent of the flowers fertilised in a legitimate manner yielded capsules, these containing on an average 27.4 good and bad seeds; whilst only 15 per cent of the flowers fertilised in an illegitimate manner yielded capsules, these containing on an average only 11.0 good and bad seeds.

In a previous part of this chapter it was shown that illegitimate crosses between the long-styled form of the primrose and the long-styled cowslip, and between the short-styled primrose and short-styled cowslip, are more sterile than legitimate crosses between these two species; and we now see that the same rule holds good almost invariably with their hybrid offspring, whether these are crossed inter se, or with either parent-species; so that in this particular case, but not as we shall presently see in other cases, the same rule prevails with the pure unions between the two forms of the same heterostyled species, with crosses between two distinct heterostyled species, and with their hybrid offspring.

Seeds from the long-styled oxlip fertilised by its own pollen were sown, and three long-styled plants raised. The first of these was identical in every character with its parent. The second bore rather smaller flowers, of a paler colour, almost like those of the primrose; the scapes were at first single- flowered, but later in the season a tall thick scape, bearing many flowers, like that of the parent oxlip, was thrown up. The third plant likewise produced at first only single-flowered scapes, with the flowers rather small and of a darker yellow; but it perished early. The second plant also died in September; and the first plant, though all three grew under very favourable conditions, looked very sickly. Hence we may infer that seedlings from self-fertilised oxlips would hardly be able to exist in a state of nature. I was surprised to find that all the pollen-grains in the first of these seedling oxlips appeared sound; and in the second only a moderate number were bad. These two plants, however, had not the power of producing a proper number of seeds; for though left uncovered and surrounded by pure primroses and cowslips, the capsules were estimated to include an average of only from fifteen to twenty seeds.

From having many experiments in hand, I did not sow the seed obtained by crossing both forms of the primrose and cowslip with both forms of the oxlip, which I now regret; but I ascertained an interesting point, namely, the character of the offspring from oxlips growing in a state of nature near both primroses and cowslips. The oxlips were the same plants which, after their seeds had been collected, were transplanted and experimented on. From the seeds thus obtained eight plants were raised, which, when they flowered, might have been mistaken for pure primroses; but on close comparison the eye in the centre of the corolla was seen to be of a darker yellow, and the peduncles more elongated. As the season advanced, one of these plants threw up two naked scapes, 7 inches in height, which bore umbels of flowers of the same character as before. This fact led me to examine the other plants after they had flowered and were dug up; and I found that the flower-peduncles of all sprung from an extremely short common scape, of which no trace can be found in the pure primrose. Hence these plants are beautifully intermediate between the oxlip and the primrose, inclining rather towards the latter; and we may safely conclude that the parent oxlips had been fertilised by the surrounding primroses.

From the various facts now given, there can be no doubt that the common oxlip is a hybrid between the cowslip (P. veris, Brit. Fl.) and the primrose (P. vulgaris, Brit. Fl.), as has been surmised by several botanists. It is probable that oxlips may be produced either from the cowslip or the primrose as the seed- bearer, but oftenest from the latter, as I judge from the nature of the stations in which oxlips are generally found (2/13. See also on this head Hardwicke's 'Science Gossip' 1867 pages 114, 137.), and from the primrose when crossed by the cowslip being more fertile than, conversely, the cowslip by the primrose. The hybrids themselves are also rather more fertile when crossed with the primrose than with the cowslip. Whichever may be the seed-bearing plant, the cross is probably between different forms of the two species; for we have seen that legitimate hybrid unions are more fertile than illegitimate hybrid unions. Moreover a friend in Surrey found that 29 oxlips which grew in the neighbourhood of his house consisted of 13 long-styled and 16 short-styled plants; now, if the parent-plants had been illegitimately united, either the long- or short-styled form would have greatly preponderated, as we shall hereafter see good reason to believe. The case of the oxlip is interesting; for hardly any other instance is known of a hybrid spontaneously arising in such large numbers over so wide an extent of country. The common oxlip (not the P. elatior of Jacq.) is found almost everywhere throughout England, where both cowslips and primroses grow. In some districts, as I have seen near Hartfield in Sussex and in parts of Surrey, specimens may be found on the borders of almost every field and small wood. In other districts the oxlip is comparatively rare: near my own residence I have found, during the last twenty-five years, not more than five or six plants or groups of plants. It is difficult to conjecture what is the cause of this difference in their number. It is almost necessary that a plant, or several plants belonging to the same form, of one parent-species, should grow near the opposite form of the other parent-species; and it is further necessary that both species should be frequented by the same kind of insect, no doubt a moth. The cause of the rare appearance of the oxlip in certain districts may be the rarity of some moth, which in other districts habitually visits both the primrose and cowslip.

Finally, as the cowslip and primrose differ in the various characters above specified,--as they are in a high degree sterile when intercrossed,--as there is no trustworthy evidence that either species, when uncrossed, has ever given birth to the other species or to any intermediate form,--and as the intermediate forms which are often found in a state of nature have been shown to be more or less sterile hybrids of the first or second generation,--we must for the future look at the cowslip and primrose as good and true species.

Primula elatior, Jacq., or the Bardfield Oxlip, is found in England only in two or three of the eastern counties. On the Continent it has a somewhat different range from that of the cowslip and primrose; and it inhabits some districts where neither of these species live. (2/14. For England, see Hewett C. Watson 'Cybele Britannica' volume 2 1849 page 292. For the Continent, see Lecoq 'Geograph. Botanique de l'Europe' tome 8 1858 page 142. For the Alps see 'Annals and Magazine of Natural History' volume 9 1842 pages 156 and 515.) In general appearance it differs so much from the common oxlip, that no one accustomed to see both forms in the living state could afterwards confound them; but there is scarcely more than a single character by which they can be distinctly defined, namely, their linear-oblong capsules equalling the calyx in length. (2/15. Babington 'Manual of British Botany' 1851 page 258.) The capsules when mature differ conspicuously, owing to their length, from those of the cowslip and primrose. With respect to the fertility of the two forms when these are united in the four possible methods, they behave like the other heterostyled species of the genus, but differ somewhat (see Tables 1.8 and 1.12.) in the smaller proportion of the illegitimately fertilised flowers which set capsules. That P. elatior is not a hybrid is certain, for when the two forms were legitimately united they yielded the large average of 47.1 seeds, and when illegitimately united 35.5 per capsule; whereas, of the four possible unions (Table 2.14) between the two forms of the common oxlip which we know to be a hybrid, one alone yielded any seed; and in this case the average number was only 11.6 per capsule. Moreover I could not detect a single bad pollen-grain in the anthers of the short-styled P. elatior; whilst in two short-styled plants of the common oxlip all the grains were bad, as were a large majority in a third plant. As the common oxlip is a hybrid between the primrose and cowslip, it is not surprising that eight long-styled flowers of the primrose, fertilised by pollen from the long-styled common oxlip, produced eight capsules (Table 1.18), containing, however, only a low average of seeds; whilst the same number of flowers of the primrose, similarly fertilised by the long-styled Bardfield oxlip, produced only a single capsule; this latter plant being an altogether distinct species from the primrose. Plants of P. elatior have been propagated by seed in a garden for twenty-five years, and have kept all this time quite constant, excepting that in some cases the flowers varied a little in size and tint. (2/16. See Mr. H. Doubleday in the 'Gardener's Chronicle' 1867 page 435, also Mr. W. Marshall ibid. page 462.) Nevertheless, according to Mr. H.C. Watson and Dr. Bromfield (2/17. 'Phytologist' volume 1 page 1001 and volume 3 page 695.), plants may be occasionally found in a state of nature, in which most of the characters by which this species can be distinguished from P. veris and vulgaris fail; but such intermediate forms are probably due to hybridisation; for Kerner states, in the paper before referred to, that hybrids sometimes, though rarely, arise in the Alps between P. elatior and veris.

Finally, although we may freely admit that Primula veris, vulgaris, and elatior, as well as all the other species of the genus, are descended from a common primordial form, yet from the facts above given, we must conclude that these three forms are now as fixed in character as are many others which are universally ranked as true species. Consequently they have as good a right to receive distinct specific names as have, for instance, the ass, quagga, and zebra.

Mr. Scott has arrived at some interesting results by crossing other heterostyled species of Primula. (2/18. 'Journal of the Linnean Society Botany' volume 8 1864 page 93 to end.) I have already alluded to his statement, that in four instances (not to mention others) a species when crossed with a distinct one yielded a larger number of seeds than the same species fertilised illegitimately with its own-form pollen, though taken from a distinct plant. It has long been known from the researches of Kolreuter and Gartner, that two species when crossed reciprocally sometimes differ as widely as is possible in their fertility: thus A when crossed with the pollen of B will yield a large number of seeds, whilst B may be crossed repeatedly with pollen of A, and will never yield a single seed. Now Mr. Scott shows in several cases that the same law holds good when two heterostyled species of Primula are intercrossed, or when one is crossed with a homostyled species. But the results are much more complicated than with ordinary plants, as two heterostyled dimorphic species can be intercrossed in eight different ways. I will give one instance from Mr. Scott. The long-styled P. hirsuta fertilised legitimately and illegitimately with pollen from the two forms of P. auricula, and reciprocally the long-styled P. auricula fertilised legitimately and illegitimately with pollen from the two forms of P. hirsuta, did not produce a single seed. Nor did the short-styled P. hirsuta when fertilised legitimately and illegitimately with the pollen of the two forms of P. auricula. On the other hand, the short-styled P. auricula fertilised with pollen from the long-styled P. hirsuta yielded capsules containing on an average no less than 56 seeds; and the short-styled P. auricula by pollen of the short- styled P. hirsuta yielded capsules containing on an average 42 seeds per capsule. So that out of the eight possible unions between the two forms of these two species, six were utterly barren, and two fairly fertile. We have seen also the same sort of extraordinary irregularity in the results of my twenty different crosses (Tables 2.14 to 2.18), between the two forms of the oxlip, primrose, and cowslip. Mr. Scott remarks, with respect to the results of his trials, that they are very surprising, as they show us that "the sexual forms of a species manifest in their respective powers for conjunction with those of another species, physiological peculiarities which might well entitle them, by the criterion of fertility, to specific distinction."

Finally, although P. veris and vulgaris, when crossed legitimately, and especially when their hybrid offspring are crossed in this manner with both parent-species, were decidedly more fertile, than when crossed in an illegitimate manner, and although the legitimate cross effected by Mr. Scott between P. auricula and hirsuta was more fertile, in the ratio of 56 to 42, than the illegitimate cross, nevertheless it is very doubtful, from the extreme irregularity of the results in the various other hybrid crosses made by Mr. Scott, whether it can be predicted that two heterostyled species are generally more fertile if crossed legitimately (i.e. when opposite forms are united) than when crossed illegitimately.

SUPPLEMENTARY NOTE ON SOME WILD HYBRID VERBASCUMS.

In an early part of this chapter I remarked that few other instances could be given of a hybrid spontaneously arising in such large numbers, and over so wide an extent of country, as that of the common oxlip; but perhaps the number of well-ascertained cases of naturally produced hybrid willows is equally great. (2/19. Max Wichura 'Die Bastardbefruchtung etc. der Weiden' 1865.) Numerous spontaneous hybrids between several species of Cistus, found near Narbonne, have been carefully described by M. Timbal-Lagrave (2/20. 'Mem. de l'Acad. des Sciences de Toulouse' 5e serie tome 5 page 28.), and many hybrids between an Aceras and Orchis have been observed by Dr. Weddell. (2/21. 'Annales des Sc. Nat.' 3e serie Bot. tome 18 page 6.) In the genus Verbascum, hybrids are supposed to have often originated in a state of nature (2/22. See for instance the 'English Flora' by Sir J.E. Smith 1824 volume 1 page 307.); some of these undoubtedly are hybrids, and several hybrids have originated in gardens; but most of these cases require, as Gartner remarks, verification. (2/23. See Gartner 'Bastarderzeugung' 1849 page 590.) Hence the following case is worth recording, more especially as the two species in question, V. thapsus and lychnitis, are perfectly fertile when insects are excluded, showing that the stigma of each flower receives its own pollen. Moreover the flowers offer only pollen to insects, and have not been rendered attractive to them by secreting nectar.

I transplanted a young wild plant into my garden for experimental purposes, and when it flowered it plainly differed from the two species just mentioned and from a third which grows in this neighbourhood. I thought that it was a strange variety of V. thapsus. It attained the height (by measurement) of 8 feet! It was covered with a net, and ten flowers were fertilised with pollen from the same plant; later in the season, when uncovered, the flowers were freely visited by pollen-collecting bees; nevertheless, although many capsules were produced, not one contained a single seed. During the following year this same plant was left uncovered near plants of V. thapsus and lychnitis; but again it did not produce a single seed. Four flowers, however, which were repeatedly fertilised with pollen of V. lychnitis, whilst the plant was temporarily kept under a net, produced four capsules, which contained five, one, two, and two seeds; at the same time three flowers were fertilised with pollen of V. thapsus, and these produced two, two, and three seeds. To show how unproductive these seven capsules were, I may state that a fine capsule from a plant of V. thapsus growing close by contained above 700 seeds. These facts led me to search the moderately-sized field whence my plant had been removed, and I found in it many plants of V. thapsus and lychnitis as well as thirty-three plants intermediate in character between these two species. These thirty-three plants differed much from one another. In the branching of the stem they more closely resembled V. lychnitis than V. thapsus, but in height the latter species. In the shape of their leaves they often closely approached V. lychnitis, but some had leaves extremely woolly on the upper surface and decurrent like those of V. thapsus; yet the degree of woolliness and of decurrency did not always go together. In the petals being flat and remaining open, and in the manner in which the anthers of the longer stamens were attached to the filaments, these plants all took more after V. lychnitis than V. thapsus. In the yellow colour of the corolla they all resembled the latter species. On the whole, these plants appeared to take rather more after V. lychnitis than V. thapsus. On the supposition that they were hybrids, it is not an anomalous circumstance that they should all have produced yellow flowers; for Gartner crossed white and yellow-flowered varieties of Verbascum, and the offspring thus produced never bore flowers of an intermediate tint, but either pure white or pure yellow flowers, generally of the latter colour. (2/24. 'Bastardzeugung' page 307.)

My observations were made in the autumn; so that I was able to collect some half-matured capsules from twenty of the thirty-three intermediate plants, and likewise capsules of the pure V. lychnitis and thapsus growing in the same field. All the latter were filled with perfect but immature seeds, whilst the capsules of the twenty intermediate plants did not contain one single perfect seed. These plants, consequently, were absolutely barren. From this fact,--from the one plant which was transplanted into my garden yielding when artificially fertilised with pollen from V. lychnitis and thapsus some seeds, though extremely few in number,--from the circumstance of the two pure species growing in the same field,--and from the intermediate character of the sterile plants, there can be no doubt that they were hybrids. Judging from the position in which they were chiefly found, I am inclined to believe they were descended from V. thapsus as the seed-bearer, and V. lychnitis as the pollen-bearer.

It is known that many species of Verbascum, when the stem is jarred or struck by a stick, cast off their flowers. (2/25. This was first observed by Correa de Serra: see Sir J.E. Smith's 'English Flora' 1824 volume 1 page 311; also 'Life of Sir J.E. Smith' volume 2 page 210. I was guided to these references by the Reverend W.A. Leighton, who observed this same phenomenon with V. virgatum.) This occurs with V. thapsus, as I have repeatedly observed. The corolla first separates from its attachment, and then the sepals spontaneously bend inwards so as to clasp the ovarium, pushing off the corolla by their movement, in the course of two or three minutes. Nothing of this kind takes place with young barely expanded flowers. With Verbascum lychnitis and, as I believe, V. phoeniceum the corolla is not cast off, however often and severely the stem may be struck. In this curious property the above-described hybrids took after V. thapsus; for I observed, to my surprise, that when I pulled off the flower-buds round the flowers which I wished to mark with a thread, the slight jar invariably caused the corollas to fall off.

These hybrids are interesting under several points of view. First, from the number found in various parts of the same moderately-sized field. That they owed their origin to insects flying from flower to flower, whilst collecting pollen, there can be no doubt. Although insects thus rob the flowers of a most precious substance, yet they do great good; for, as I have elsewhere shown, the seedlings of V. thapsus raised from flowers fertilised with pollen from another plant, are more vigorous than those raised from self-fertilised flowers. (2/26. 'The Effects of Cross and Self-fertilisation' 1876 page 89.) But in this particular instance the insects did great harm, as they led to the production of utterly barren plants. Secondly, these hybrids are remarkable from differing much from one another in many of their characters; for hybrids of the first generation, if raised from uncultivated plants, are generally uniform in character. That these hybrids belonged to the first generation we may safely conclude, from the absolute sterility of all those observed by me in a state of nature and of the one plant in my garden, excepting when artificially and repeatedly fertilised with pure pollen, and then the number of seeds produced was extremely small. As these hybrids varied so much, an almost perfectly graduated series of forms, connecting together the two widely distinct parent-species, could easily have been selected. This case, like that of the common oxlip, shows that botanists ought to be cautious in inferring the specific identity of two forms from the presence of intermediate gradations; nor would it be easy in the many cases in which hybrids are moderately fertile to detect a slight degree of sterility in such plants growing in a state of nature and liable to be fertilised by either parent-species. Thirdly and lastly, these hybrids offer an excellent illustration of a statement made by that admirable observer Gartner, namely, that although plants which can be crossed with ease generally produce fairly fertile offspring, yet well-pronounced exceptions to this rule occur; and here we have two species of Verbascum which evidently cross with the greatest ease, but produce hybrids which are excessively sterile.


Charles Darwin

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