Journal of the New York Botanical Garden. 30(354): 129-136 (June 1929)
THE FULVOUS DAYLILIES — I
A. B. STOUT

1. THE HEMEROCALLIS FULVA CLON EUROPA.

The oldest and the best known of the fulvous daylilies is the one commonly cultivated under the name Hemerocallis fulva L. This single-flowered daylily was described by Lobel in 1576 (Historia) as having cinnabar-red coloring in the flowers and as being in this particular very distinct from the yellow-flowered daylily (H. flava), which was also then in cultivation in Europe. In 1601, Clusius (Plantarum Historia) states that this plant was being grown in many gardens throughout Austria and Germany. Nearly two hundred years later, Linnaeus (1753) considered this daylily as a hybrid, but a few years afterward (Species Plantarum, ed. 2, 1762) he gave to it the specific rank and the name Hemerocallis fulva which have since been accorded to it.

But the plants of this particular daylily do not produce seeds to any kind of pollination possible for them alone. They are evidently never able to do so because of a complete self-incompatibility in the processes of fertilization. The propagation of this daylily is by division. All the plants of it are merely branches derived from one original seedling and hence they constitute a clon and not a species or a variety that reproduces true to type by seeds. This distinction was not recognized by Linnaeus, but it is necessary to make the distinction if one is clearly to understand the true status of those groups of plants, both in culture and in the wild, that have arisen from a single individual by repeated vegetative division. The Hemerocallis fulva of Linnaeus, which was in cultivation in Europe for at least one hundred and eighty-six years before his date of naming it as a species (1576 to 1762), and which has continued in cultivation for one hundred and sixty-five years since that date—a total of over three and a half centuries—exists solely as a clon.

The question arises whether this clon, the H. fulva of Linnaeus, is represented among the wild daylilies and is itself typical of a natural species. Botanical and horticultural explorations and studies in Japan and China, beginning about the year 1700 and continuing to the present date, have revealed various fulvous daylilies, but the exact counterpart of the H. fulva of Linnaeus has not been discovered as a wild plant. Nor does there seem to be any conclusive evidence that it has been included among the daylilies recently in cultivation in the Orient. On certain of the fulvous daylilies found, there were bestowed such names as H. fulva var. Kwanso, H. disticha  Donn, H. disticha var. flore-pleno, H. longituba Miq., H. fulva var. longituba Maxim., H. fulva var. angustifolia Baker, and H. fulva var. maculata  Baroni. Thus the rather marked differences between the various fulvous daylilies found in the Orient and the H. fulva  of Linnaeus were recognized. However, conservative botanical treatments, such as that of the Index Kewensis, have included all the various forms mentioned above in the one species, H. fulva. These rather diverse forms are to be recognized as valuable material for use in breeding, especially since the fulvous daylilies seem destined to play an important rôle in the development of new red-flowered daylilies valuable for garden culture.

It is time to review the different fulvous daylilies in respect to their identity and interrelationships. The knowledge now available regarding their natural distribution in the Orient is meagre and inadequate. An attempt may be made to recognize the types of them that are now known both as wild and as cultivated, and to comment on their botanical relationships and their horticultural status. In doing this the botanical status and the horticultural status of each should be clearly considered. Those that are merely clons should be recognized as such. Possibly this may lead to a better understanding of what the wild species are like.

The horticultural name "Europa Daylily" is here suggested for the single-flowered fulvous daylily which has been in continued cultivation in Europe at least since the first description of it by Lobel more than 350 years ago. This is historically the Linnaean type of the species Hemerocallis fulva. But this particular plant is a clon and it is very doubtful if its exact counterpart exists among the wild plants of the Orient. The true botanical status of the clon is simply that of the one original seedling selected for the first propagation. This status is better expressed by using a clonaI name botanically, such as Hemerocallis fulva clon Europa, or merely by using the horticultural name Europa Daylily, than by continuing to use only the species name bestowed by Linnaeus.

FIGURE. 4. At lower left, simple flower of the Daylily Europa; at lower right, a semi-double flower from a plant chiefly bearing such flowers; above a very double flower, All are fulvous daylilies with almost identical coloring.

The Europa Daylily probably arose as a single somewhat aberrant seedling, either wild or in garden culture, which attracted the attention of some Oriental gardener. Possibly the fullness of the flowers and the somewhat bold pattern of their coloring were outstanding qualities. The plant happened to be completely self-incompatible—a type of sterility very common in daylilies. But it also happened to have the habit of spreading vigorously by rhizomes and this made vegetative propagation easy and has maintained the individual character of the original seedling throughout several centuries of garden culture. This clon remains today in vigorous growth and is thus a demonstration that long-continued asexual reproduction does not of itself necessarily reduce vigor and lead to degeneration. We may perhaps assume that the Daylily Europa had its origin in the Orient, together with the Lemon Daylily (H. flava). How and when these two found their way into garden culture in Europe as reported in 1567 are at present apparently matters of conjecture.

A plant of the Europa Daylily is among the most robust of daylilies. In coarseness of foliage and vigor of growth it is surpassed only by the double-flowered form. The dome of leaves reaches a height of about three feet. The leaves are light green, strongly distichous in arrangement, and rather coarse. The plant extends itself vigorously by widely spreading rhizomes. The roots are numerous, and some are enlarged or fleshy. The strong scapes are stiffly erect to a height of about 50 inches, are branched at the summit, and bear as many as 15 to 20 flowers.

The flowers have a fulvous overcast of color in the outer zone of the open flower with reticulated veins of darker shades. An arching mid-zone of darker shade in the petals is a conspicuous feature. The throat of the flower is orange only. The petals are rather thin, slightly wavy along the margin, and of delicate texture, yet they retain form and color well during the day. The segments are rather broad and overlapping, giving a full flower.

The capsule, produced only rarely and to the compatible fertilizations of certain cross-pollinations, is about one inch in length, broadly ovate, with the apex truncate and indented.

The persistent self-unfruitfulness of the entire clon of the Europa Daylily makes it impossible to obtain selfed seedlings of the clon. Its sterilities in cross-relations are also so decided that it is only within very recent years that the clon Europa has been used successfully in hybridization with other daylilies.

2. THE D0UBLE-FLOWERED FULVOUS DAYLtLIES

The varietal names "Kwanso" and "flore-pieno" have been given to double-flowered daylilies that are very closely related to the Daylily Europa. The first record of such a daylily is by Kaempfer, a physician and botanist who was among the first of the Europeans to live in Japan. In his volume on Japanese plants (Amoenitatum Exoticarum ... Descriptiones, 872. 1712) he mentions an "Iris" and briefly describes it as having large double flowers of the color of fire. But he gave the Japanese names for it (Ken, Quanso, and Wasrigusa), through which the plant was identified later by Thunberg (Flora Japonica, 1784) as a Hemerocallis.

A Japanese work on plants (Somoku-Dzusetsu, 2nd edition, 6: 13. 1874) gives an uncolored illustration from a drawing of a double-flowered daylily named Yabukwanzo. In a later edition this plant was identified as H. fulva L. var. Kwanso. Various publications on Japanese plants mention this double-flowered fulvous daylily as rather widely cultivated in Japan.

The first introduction into Europe of a double-flowered fulvous daylily was by a Rev. W. Ellis, who brought living plants from Mauritius. The firm Veitch & Son exhibited this plant under the name H. disticha flore-pleno before the Floral Committee of the Royal Horticultural Society in London during 1860, receiving for it a First Class Certificate (Gard. Chron. 1860: 482). A year later a colored plate was published (The Floral Magazine 1:  pl. 13) which is in very poor coloring but shows a flower that is much doubled. The excellent volume which surveys the botanical and horticultural contributions of the Veitch firm mentions (Hortus Veitchii, 446. 1906) the daylily introduced from Mauritius as H. fulva var. flore-pleno and says of it "A semi-double form of the common Day Lily, with spikes of orange flowers similar in color to the type, possessing the important quality of remaining longer on the plant."

Soon after the introduction of the fulvous Daylily Flore-Pleno by Veitch & Son, plants known as H. fulva Kwanso were brought into Europe directly from Japan by von Siebold. The first mention of this plant appears to be a mere note in the Gardeners' Chronicle (1864: 654) that a plant of Hemerocallis Kwanso foliis variegatis"  had been exhibited. Two years later a plate in good color, showing the double flower and the white-striped or variegated foliage of this plant was published (Gartenflora 15: pl. 500. 1866).

The Gardeners' Chronicle in 1867 (32: 292) speaks of this plant introduced by von Siebold under the name Hemerocallis Kwanso flore-pleno and states that it "is merely a variety of H. fulva, with the leaves more or less distinctly marked with white stripes, and the tawny-red flowers filled out with a tuft of smaller petaloid segments in the center."

Thus it appears that the H. fulva Kwanso first introduced was variegated as well as double-flowered. The variegation is evidently of the chimeral type and plants with this type frequently produce all-green branches. Hence it may well be that some of the double-flowered plants with all-green foliage now in cultivation arose from the Kwanso Daylily.

The propagation of these two daylilies is solely by vegetative means. It can not be otherwise, for the pistils of the flowers are entirely impotent. The two types are merely clons.

It is generally considered that the clon of the Daylily Flore-Pleno introduced by Veitch & Son is less double than the Daylily Kwanso. It is to be noted, however, that these conditions are exactly reversed in the first illustrations of the two. The plate of van Siebold's plant published in 1866 (Gartenflora, pl. 500) shows a semi-double flower, while that of Veitch's plant (The Floral Magazine 1: pl. 13) shows a flower much more double. The writer has obtained entirely green-leaved plants under both names Kwanso and Flore-Pleno from various nurserymen and from various botanical gardens. These have been grown side by side for comparison and study. There seems to be no difference in the color of the flowers and the various aspects of the habit of growth. Some plants, however, have all flowers very double, while other plants have semi-double flowers, with also many inter-grades; various grades of doubleness may be found on the same plant.

In 1924 Mr. Henry H. White, of the Department of Agriculture, American Presbyterian Mission, in Anhwei, China, sent to The New York Botanical Garden living plants of daylilies which he had collected from an old abandoned cemetery. These plants have green leaves and bear flowers of various grades of doubleness with many flowers only semi-double.

When plants of the Europa Daylily are grown beside plants having double flowers the former are earlier to bloom, have foliage somewhat less coarse, and have taller scapes. The color of the flowers is the same. The zone or eye spot is, however, less developed in the smaller and inner accessory segments of the double flowers than in their more primary segments and in the petals of the single-flowered Daylily Europa.

Of the origin of the double-flowered type there is no record. It was in existence in Japan in 1712 and it is known to be now rather widely distributed in Japan and probably also in China, both in culture and as an apparent escape, evidently, quite as the Daylily Europa now exists in such old-settled areas as Long Island about abandoned homesites, along roadsides, and wherever the activities of man have given the plant a chance to spread vegetatively. All the distribution of the double-flowered fulvous daylilies in the Orient as well as in Europe and in America has been by vegetative propagation. The very close relationship between the double-flowered types Kwanso and Flore-Pleno and the single-flowered Daylily Europa is obvious. The double-flowered type probably arose either directly from the Daylily Europa as a bud-sport, or it may have been a seedling member of the same race. Some evidence bearing on the origin of these rather anomalous types of fulvous daylilies will undoubtedly be obtained from the character of the seedlings derived from using them in cross-breeding.

The double-flowered daylilies have not become popular garden flowers. The color of the flowers is perhaps too dull. The flowers also seem monstrous, coarse, and lacking in pleasing symmetry. As in the case of the double-flowered true lilies (Lilium tigrinum flore-pleno, for example), they are not very attractive. Besides, the flowers stand rather high above the leaves. Possibly double-flowered daylilies may yet be developed with sprightly colored and more symmetrical flowers, and with a more pleasing habit of growth.

3. FULVOUS DAYLILIES WITH VARIEGATED FOLIAGE

Thunberg in his volume on Japanese plants (Flora Japonica, page 142), published in 1784, makes mention of a white variegated variety of what he called Hemerocallis fulva. He also speaks of varieties with simple and with double flowers but does not say which had the variegated foliage. A daylily with variegated leaves is mentioned in 1829 (Roemer & Schultes, Systema Vegetabilium) as a variety of H. disticha and since a variety with "pleno" flowers is also mentioned one might infer that the variegated plants were single-flowered. In the various descriptions of the double-flowered fulvous daylilies usually there is no mention of variegated leaves. In 1864, The Gardeners' Chronicle (page 654) mentions the display of a "Hemerocallis Kwanso foliis variegatis" and states that it was a "well-marked variegated plant, very near the variegated variety of H. fulva." Evidently the H. fulva Kwanso introduced from Japan by von Siebold was both double-flowered and variegated, according to the plate of it published in 1966 (Gartenflora 15: pl. 500) and the early description of it.

From time to time after 1866 two sorts of variegated daylilies have been mentioned. One with single flowers is usually called H. fulva variegata; one with double-flowers is generally known as H. fulva Kwanso variegata. The latter has been spoken of as "almost as striking in appearance as the well-known Pandanus Veitchii, which at first sight it somewhat resembles (Gard. Chr. 51: 681, 1882).

The variegated daylilies are seldom offered in the trade either in. single or in double flowers. It is possible that plants of one or of both of the variegated forms exist in certain gardens in America, but if so the writer has not heard of them. Through the special efforts of Director Fred J. Chittenden, of the Royal Horticultural Society Gardens at Wisley, England, The New York Botanical Garden has at last received plants of variegated daylilies, which, however, are yet to come into bloom.

There appears to be no accurate description by which one may determine whether the flowers of the single-flowered variegated daylily are like those of the Daylily Europa or are different. For a more precise description and a correct naming of the clon or the clons of the variegated daylilies, the flowers are necessary.

Part II