THE NATURAL HYBRID BETWEEN THE COWSLIP AND OXLIP, BY ARTHUR W. HILL, M.A.

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1 i62 Arthur W. Hill. THE NATURAL HYBRID BETWEEN THE COWSLIP AND OXLIP, BY ARTHUR W. HILL, M.A. [WITH PLATE V. AND TEXT-FIGS. 5 AND 6.] ONE of the most interesting features of the flora of the country around Cambridge is the occurrence of the true Oxlipr Primula elatior L. Jacquin's or the Bardfleld Oxiip in certain areas of woodland on the boulder :lay, and the ahsence of the Primrose P. acaulis L. from such woods as are well within the Oxlip area. The exact distribution of the Oxlip has been so beautifully worked out by Miller Christy' that it would be superfluous to enter into the question in any further detail in t^is brief note. Suffice it io say that the main Oxlip area occupies an irregular tract of country on the borders of Cambridgeshire,. Essex and SufFoIk, with large extensions into Suffolk and Essex; whilst there is a small triangular area to the west of Cambridge, roughly bisected by the old- North Road. In addition to this there are two small outliers, one in the neighbourhood of Great Livermere to the north of Bury St, Edmunds, and the other near Diss, in Norfolk. The primrose occurs round the margins of these areas, and, as is well known, hybridises freely with the Oxlip. There seems little reason to doubt that the restricted and peculiarly outlined habitat cf the Oxlip in England is due to the fact that it is being gradually ousted by the more successful Primrose. On the Continent the Oxlip appears to have as extensive a. range as both Primrose and Cowslip (P. officinalis L.), being found all over Western and Central Europe, from Holland and France to S. Russia and from S. Sweden and Denmark to the Northern Alps.- Althougli the Primrose and Oxlip appear to be mutually exclusive in England, the Cowslip, on the other hand, grows freely throughout the Oxlip country, but hybrids between the two species are very uncommon. A few records of the cross elatior X officinalis aregiven by Pax' and are sometimes known as P. media ; and Christy' mentions the finding of three probable Cowslip X Oxlip hybrids during the last eighteen years. The rarity of this cross is- ' Miller Christy. P. elatior in Britain, Journ. Linn. Soc. XXXIII., p. 172, with map. Bailey. Journ. Bot., XLI., p ' Pax & Knuth. Pflanzenreich ; Priihulacex, pp. 60, 61. Miller Christy. I.e., pp. 197, 198.

2 COWSLIP-OXLIP HYBRID. THE NEW PHYTOLOGIST. VOL. VI. PLATE 5.

3 The Natural Hybrid between the Cowslip & Oxlip. 103 due partly no doubt to the common difference in habitat of the two species the Oxlip living in the vyroods in England, and the Cowslip as a rule in meadows, but more probably to the difference in their times of flowering, the Oxlip usually being in full flower and getting over when the Cowslip is coming into blossom. On the Continent both species are often found freely intermixed' and this is also the case in the woods of the outlying district near Great Livermere, where the plant which forms the subject of this note was found. This small outlier to the north of Bury contains at least two localities where Oxlips may be found. In one, an overgrown swamp at the edge of a lalte, there are only Oxlips, and in the other, a neighbouring wood on a gentle slope, Primroses occupy the higher southern end, and also grow on the outskirts; whilst the northern lower portion is filled with Oxlips and Cowslips and numerous Primrose X Oxlip hybrids. It was in this wood that a single old plant of the Oxlip x Cowslip hybrid was found by the side of a broad path, and it was strikingly different from the Oxlips growing round it. The hybrid was discovered on the 26th of April, 1906, and had about half the flowers in the umbel expanded, whilst the Oxlips in the same locality were fully open. In its somewhat later flowering period the hybrid appears to be intermediate between the two parents. The leaves are flrm and stiff on long petioles and are somewhat stiffly curved, which causes the lamina to lie either on the surface of the ground, or parallel to the surface (PI. V., Fig. 1.) This character reminds one of the habit of the Cowslip,' as does also the obovate-oblong lamina, which is narrower than that of the Oxlip and longer than that of the Cowslip (PI. V., Figs. 2 and 3); the serration of the margin is also intermediate in character between Oxlip and Cowslip. The upper surface of the lamina shews a velvety pile of short hairs like the Cowslip, but the hairs are longer than in that species, and, with regard to the under 8urface, it more nearly approaches the Oxlip. The inflorescence characters are of considerable interest. In the Cowslip the scape bearing the umbel is stout, rather fleshy, and easily flexible (of white-buff colour), and covered with a short velvety pubescence (PI. V., Fig. 3). In the Oxlip the scape is rather thin and wiry, stiff and erect, pale-green in colour, and covered with a tomentum of rather woolly hairs (Fig. 2). The hybrid shews the characteristic Oxlip scape, with a velvety tonientum and green colour (Fig. 1). The flowers of the umbel are pendulous, after the ' Kerner. Oester. Bot. Zeitsch. XV., p. 80. > In the Oxlip the leaves tend to be erect.

164 Arthur W. Hill. manner of the two parents, and are thus in striking contrast to the usual arrangement of the flowers in the Cowslip X Primrose hybrid. The calyx is about 16 mm.

4 164 Arthur W. Hill. manner of the two parents, and are thus in striking contrast to the usual arrangement of the flowers in the Cowslip X Primrose hybrid. The calyx is about 16 mm. in length, slightly inflated, with the teeth about 7 mm. in length, triangular to oblong, acute to acuminate (Fig. 6). The teeth and sharp ribs are pale-green, like those of the Oxlip (Fig. 5), but the membranous portions of the calyx between the keeled mid-ribs are not so sharply differentiated as in this species. In the Cowslip (Fig. 7), the inflated calyx is uniformly coloured of a whitish pale yellow-green hue, varying in length from 9 15 mm., with teeth 4 5 mm., triangular, ovate, acute. In the Oxlip the calyx is from mm. long, with teeth about 6 mm., narrowly triangular, acute to acuminate. The corolla of the hybrid is a clear deep yellow, rather paler thaa the Cowslip, with a deep orange, 5-rayed ring at the throat like that of the Oxlip, with the limb saucer to cup-shaped as in the Oxlip. The lobes are obovate, subrotund, emarginate, being shorter and more rounded than those of the O.xlip. The scent of the hybrid approximates closely to the peculiar peach-like odour characteristic of the Oxlip. The Cowslip X Primrose hybrid or false Oxlip is of interest in comparison with our hybrid Cowslip X Oxlip. The leaves shew the surface-wrinkling characteristic of the Primrose with the velvety tomentum of the Cowslip (Fig. 4). The flower-scape is stout and Cowslip-like^ but with a more dense tomentum (Figs. 4 and 8), and the umbel is composed of more or less erect Uowers, shewing the Primrose "eye," and, like that of the Oxlip X Primrose, has a somewhat untidy scattered appearance, lacking the graceful drooping character of either the Cowsli]? or the true Oxlip. An examination of the flower-scapes of the Oxlip, the Cowslip, and of the various Oxlip hybrids by means of transverse sections has yielded results of some interest. The flower stem or scape of the Oxlip is found to possess a narrow band of thick-walled sclerenchyrtia, which is pericyclic in origin and forms a layer of tissue about flve cells in depth just within the endodermis (Text-flg. 5). In the Cowslip, however, where the stem is usually of a greater diameter than in the Oxlip, tbe thickened band of the pericycle is much broader, and is composed of larger cells with thinner walls, which are only slightly ligniilied.(text-fig. 6). The sclerenchymatous tissue in this species forms a band some seven cells in thickness, and the greater flexibility of the Cowslip stalk is no doubt due to the slight amount of ligniflcatlon of the walls.

5 The Natural Hybrid between the Cowslip S Oxiip. 165 VIS' Text-fig. 5. Transverse section of theflower-scapeof the Oxlip. 'c, cortex : p,, pericydc; t., endodermis ; ph., pith ; vh, position of vascular bundle. Text-flg. 6. Transverse section of the flower-scape of the Cowslip. Lettering as above. The two scapes can thus be very easily recognized in transverse section. With regard to the structure of the scape in the various hybrids, it is found that the Cowslip has handed on its characteristic scape-structures to its hybrid with the Primrose, perhaps* owing to the fact that the Primro6e is practically acaulescent. For a similar reason no doubt the internal structure of the scape of the hybrid Oxlip Primrose shews all the characteristics of the true Oxlip. But in the Cowslip X Oxlip hybrid, where both parents have scapose umbels it is of interest to notice that the Oxlip stem-structure is dominant over that of the Cowslip, for the narrow band of thickwalled sclerenchyma is as well developed as in the Oxlip itself (cf. Pigs. 1-4).

6 i66 Arthur W. Mill. It is, of course, impossible to say whether the hybrid just described is a first cross between an Oxlip and a Cowslip or not, though the evidence seems to be in favour of this view. Its characters are more or less intermediate between those of the two parents, though the infiuence of the Oxlip appears to to be somewhat the stronger. In general hairinesd, shape, and markings of the fiower, and scent, as well as in the internal structure of its fiower-scape, the Oxlip characteristics are predominant, whilst in the shape, position, and appearance of the leaves, and in the orange colour of the flowers, the influence of the Cowslip is clearly seen." The hybrids between the three species of Primula in Britain are of such interest that it is to be hoped that they will soon be subjected to careful scientific experiment. There can be no doubt tbat inter-crossing experiments will yield results not only of considerable interest as to the various forms found in a wild state, but will also throw important light on questions of hybridisation in general. EXPLANATION OF PLATE V. ILLUSTRATING MR. A. W. HILL'S PAPER ON THE NATURAL HYBRID BETWEEN THE COWSLIP AND OXLIP. Fig. 1. The Cowslip x Oxiip hybrid, P. offitiiialis x elatior; shewing the Cowslip-like leaves and the flowers and flower-scape more like those of the Oxlip, cf Fig. 6. The corolla is orange, and the calyx shews Cowslip characters. Fig. 2. An umbel and leaf of the true Oxlip, P. etatior L, cf. Fig. 5. The corolla is of a pale cream colour with an orange ring. The calyx is contracted at the throat. Fig. 3. An umbel and leaf of the Cowslip, P. nfficinalis, L, cf. Fig. 7. The corolla is deep orange with characteristic markings and the loose calyx has short blunt teeth. Fig. 4. An umbel and leaf of the hybrid Cowslip x Primrose, P. offkinalis x acautis, cf Fig. 8. Note the wrinkled leaf and hairy flower-sc^pe. The corolla is yellow, with the flve spots of the Cowslip shewing oit.the starlike eye of the Primrose. The calyx is inflated and hairy. Figs. 5 8 for a comparison of the floral characteristics of these four plants. Fig. 5. Flowers of the Oxlip. Fig. 6. The Cowslip x Oxlip. Fig. 7. The Cowslip. Fig. 8. The Cowslip x Primrose. The photographs were taken by Mr. Altard Foreman of the Botanic Garden, Cambridge. * Owing to the innumerable gradations between the Oxlip and the Primrose, which are- found as natural hybrids, it is not possible at present to discuss the influence of each of the parents in the flrst cross.

Plant Crib VERONICA. 1. Veronica serpyllifolia

Plant Crib VERONICA. 1. Veronica serpyllifolia VERONICA 1. Veronica serpyllifolia Illustrations reproduced, with permission, from M. McC. Webster (1978). Flora of Moray, Nairn & East Inverness. Aberdeen. Subsp. humifusa (Dicks.) Syme Subsp. serpyllifolia