Saturday, June 4, 2016

What is a Daylily? 4 - Hybridization 1

What is a Daylily?

Part 4
The Origins of the Garden Daylily
Part 1

To understand the advent of popular flower hybridizing, you needs to consider the time-period of the mid-eighteenth-century from the introduction of the theories of Darwin to the research work of Augustinian friar Gregor Mendel, and his theory's subsequent rediscovery and popularization, at the beginning of the twentieth-century. This time period laid the foundation for the rise of modern plant breeding. From the beginning of the enlightenment in Europe through to the early modern genetic/agricultural revolution at the end of the Victorian Age in England, at the turn of the twentieth-century, ideas that gave us insight into natural relationships between related populations have advanced our understanding of what is actually possible in plant breeding. 

Domestication and most early plant breeding was not documented and probably wasn't really even understood by the people who practiced it, and represents selection more so than 'breeding' ( in the sense of attempts to combine or manipulate traits that are heritable to meet specific aims). Only in the last couple hundred years or so have we begun to gain an understanding of applying statistical analysis to the results of matings to infer heritability, etc. And only since the rediscovery of Mendel in 1901 have we had the tools of genetic science to guide our hand. Now in the age of genetic testing, we are beginning to have a much greater understanding of the actual 'facts on the ground', which should bring about a much more precise view of genetic factors and evolutionary relationships as we move forward. 

Many of the early breeders of daylily hybrids in the west were plantsmen in the old tradition, running huge commercial enterprises of plant hybridization and improvement. Some were trained scientists, while others were laymen influenced by the well-spring of learning and deduction that engulfed their era. They were all influenced heavily by the forces of their time, in the mid to late Victorian period. The advances in the use of science and reason, the development of rudimentary methods to collect data and make inference, and the ability to describe the natural world led to a greater understanding of the nature of plant families, genetic and evolutionary relationships, heredity and environmental responses. This in turn lead to a revolution in actively making hybrids between different species within genus (usually - though wider crosses have been known to be made successfully).

It was from the energy of that time period that the first hybrid daylilies emerged, as did so many of the hybrid flowers and plants that we take for granted now. For a review of the history of plant breeding, see the book 'Hybrid: The History and Science of Plant Breeding' by Noel Kingsbury. There you will find a complete discussion of the times I am referring to above, and their influence on many plant breeders. It is an excellent read. Now on to some specific dates of importance in Hemerocallis hybridization and the development of the modern garden daylily.

I did want to give a short overview of the period though, as it is fundamentally important to any consideration of hybrid plants produced in the last century-and-a-quarter or so. Plant breeding gained a tremendous boost that moved it out of the agricultural and into the scientific, in the late eighteen hundreds and early nineteen hundreds. Thus, by the time of Stout, his position at the New York Botanical Gardens gave him the authority and the insight to treat the Hemerocallis properly. His is still the only monograph we have of the species and gives documentation of his breeding program up to that time. Dr. Stout is not the only early daylily breeder. In fact, he is not the earliest and he built upon the success of those who can before him. However, Stout was able to approach the act of hybridizing from a systematic, information-gathering orientation that allowed him to document his work and leave us a legacy, both of genetic materials (the plants he introduced and all their many, many descendants) and the record of what he did. Due to his work with H. f. rosea, he and those who he gave divisions of the original clones (Amos Perry, Elizabeth Nesmith, etc.), would have a heavy hand in the future of the daylily through the development of bright reds, pinks, lavenders, purples and possibly even whites through the genetic ancestry of the rosea clones.

Some Dates of Significance

From the Beginning...

*Apricot by George Yeld 1893 (flava x H. middendorfii - Questionable) - Stout says in his book, Daylilies, “…”considered to be a hybrid of H. flava crossed with H. fulva or H. middendorfii”, but Mr. Yeld is quoted with the statement that he had lost the details of the crosses. It is stated that Mr. Yeld had been working with Hemerocallis for some time and that he had “a good many seedlings.” Mr. George Yeld states (Rep. Third Int. Conf. Genetics p. 415) that H. middendorfii and H. flava were the parents of Apricot. But Apricot does not closely resemble the hybrids, which the writer (Stout himself) has from this cross.)  - First known Western produced hybrid. 

*Florham by A. Herrington 1899 (H. aurantiaca 'Major' x H. thunbergii) - First known hybrid in America, bred in New Jersey.

*Amos Perry introduced his first daylily 'Amos Perry' in 1906 and through 1929 would go on to introduce 17 more for a total of 18 introductions. He was considered the 'Stout' of England and was tremendously influential early on. He was chosen by Stout to receive some of the divisions of H. fulva forma rosea.

*Luther Burbank introduced Calypso in 1917, one of four daylily hybrids he would introduce from 1914 to 1924. Calypso is a tall yellow.

*Stout had been growing daylilies in the nineteen-teens, and introduced old clones of H. fulva as early as 1917 (Green Kwanso and Flore Pleno) and 1920 (Europa). There was no AHS at the time, no registry board. What he did in those years was introduce these names for these old clones of fulva from Asia and begin to popularize them to the gardening public. Stout then set out to hybridize new forms of daylilies for garden use using species clones and the cultivars in existence at that time.

*Bertrand Farr introduced Ophir (pod parent - H. citrina) and several other daylily cultivars in 1924.

*1924 - Dr. Stout receives the original plants of H. fulva forma rosea, arriving at the New York Botanical garden from Kuling in Kiangsi, China. These were apparently wild-collected plants.

*Franklin Mead introduced Hyperion in 1925 (Sir Michael Foster (1904) x Florham (1899)). Sir Michael Foster was said to be H. aurantiaca 'Major' x H. citrina, while Florham was said to be H. aurantiaca 'Major' x H. thunbergii. Hyperion is thus a three-way hybrid that is the result of crossing F1 hybrids that are each half H. aurantiaca 'Major' crossed to two different yellow types. 

*Stout's first introductions from his own work began in 1929 with Mikado, Vesta and Wau-bun. He also introduced a clone of the species H. multiflora in 1929 through description in the literature of the day. 

*Stout - Mikado - 1929 - One year after the first description, A.B. Stout described it as: " The Mikado Daylily is a seedling recently obtained at The New York Botanical Garden in an effort of selective breeding for the development of fulvous daylilies of special merit as garden plants. It is a complex hybrid having in its parentage the two fulvous daylilies H. aurantiaca and the H. fulva clone Europa, the species H. flava and either H. aurantiaca or "H. aurantiaca major". As may be expected from such a complex ancestry the seedlings thus obtained show much diversity, and the colors of their flowers range from pale lemon yellow through many shades of yellow, of orange, and of fulvous red, both in single color effects and in various combinations of eyed patterns. In the flower of the seedling which was named Mikado the zone of color is more intense and in greater contrast than in any other daylily now known. There is also good size, form, and fullness of flower, and an excellent semi-robust habit of growth. All who have seen this daylily in bloom readily agree that it is a plant of unusual charm and of distinctive merit as a plant for flower gardens. The descriptions of new daylilies published in "House and Garden" for January, 1929, contain the first printed reference to this clone. "

"The foliage of the Mikado Daylily is medium coarse, ascending curving and reaching a general level of about 20 to 24 inches. The flower stems rise about ten inches higher and are ascending rather than erect. The dome of leaves remains freshly green and in tidy attractive appearance until the heavy freezes of autumn arrive. The flowers are about five inches in spread; the segments are fairly broad, somewhat stiffly spreading-recurving, and of good texture. In the middle of each petal there is a large blotch of dark and almost purplish red of the shade called mahogany red, bisected by a strip of the same color as that of the blade, and in the open flower these combine to form an undulating zone of bold coloring which is in sharp contrast to the rich orange of the rest of the flower. The season of bloom is mid-summer, or chiefly during July at The New York Botanical Garden. The clone Mikado does not set seed to self-pollination. The capsules, obtained by cross-pollination, are nearly ovate in outline, somewhat grooved, and in general character not closely resembling any one of the species involved in the parentage. "
(cited from: Addisonia, 1930, vol. 15, p. 13 (plate 487) ) - Parent of Rajah and others."

*Stout - Wau-Bun - 1929 - It has for its immediate ancestry the two species Hemerocallis flava and H. aurantiaca and the Luteola Daylily which is in itself a hybrid (a hybrid between H. Thunbergii and either H. aurantiaca or H. aurantiaca major). Full siblings include Soudan (1932) and Vesta (1929).

*In 1930 Stout introduced Cinnabar - (Luteola x H. aurantiaca).

*In 1931 Elizabeth Nesmith made her first introduction, Fulvax. The first major woman in daylily hybridizing, she would go on to make a huge mark on the plant. She began gathering every form of daylily she could locate into her garden for growing and hybridizing as early as 1910. She was a friend of Dr. Stout and used his introductions extensively. Her reds descend from Theron and Vulcan. Her pinks, for which she was well known, all descend from the original H. f. rosea, which was given to her by Dr. Stout. Her first pink introduction - Sweetbriar - is a direct descendant of rosea. She is one of the pioneers of the pink daylily.

*Stout introduced Bijou - 1932 - it is a complex hybrid having in its immediate ancestry the species H. aurantiaca, H. flava, and H. fulva clone Europa. The plant of H. multiflora used as the pollen parent of the Bijou Daylily has tall, erect, much-branched scapes and it is somewhat earlier in blooming than the other plants of the species. The Bijou Daylily is a hybrid with four distinct species involved in its parentage. As to flower color it is to be classed as fulvous; in flower size it is very similar to the H. multiflora but the flowers are of a different shape.

*Stout introduced Theron in 1934. A complex blend of H. flava, H. fulva ‘Europa’, H. aurantiaca and H. thunbergii. “A climax in selective breeding for dark red color. ‘Theron’, the first truly red daylily, was produced by Dr. Stout and described in 1934. Four different species and eight selection seedlings were involved in the ancestry. It took twenty-five years of painstaking work. “ “The Theron daylily represented a new combination of genetic factors with intensified pigmentation is from Hemerocallis aurantiaca and H. fulva ‘Europa’. Quite possibly, the genes for color intensification are from H. flava and/or H. thunbergii.” While the chart showing the ancestry of Theron is complex and weaves somewhat between the generations, Theron is basically the fifth generation from the four species. The three F1 seedlings are 6#1 (H. f. ‘Europa and H. flava cross), 12#1 (H. aurantiaca and H. flava cross) and Luteola’ (H. aurantiaca crossed with H. thunbergii). There are two seedlings in the second generation - 70#1 and 70#2 (both are H. f. ‘Europa’ x seedling 12#1 (from the F1 set). In the third generation, there is one seedling. This seedling is not designated with a seedling number, but the artwork reveals a richly colored, fulva like eyed orange-red flower. This seedling 70#1 crossed to Luteola. The fourth generation has two seedlings, also not labeled. The first is the third generation seedling (70#1 and Luteola crossed) crossed to seedling 6#1. The other fourth generation seedling is seedling 6#1 crossed with seedling 70#2. Amongst the resulting seedlings of crossing the two fourth generation seedlings was Theron in the fifth generation.

*Stout introduced Midas in 1935 - (Luteola x H. aurantiaca)

*In 1936 Stout introduced LindaThe ancestry of the Linda Daylily includes the species Hemerocallis Thunbergii, H. citrina, and two different seedlings of H. flava which came from the wild in central China. It is then very interesting that four yellow species created a second generation, four-way hybrid that seems to show the eye and anthocyanic overlay to the petals that we expect to only come from fulvous types...Regardless, this an exceptional plant that is still exceptional to this day. It is my favorite of all the Stout introductions I have grown and anyone interested in good traits and spidery types should have this plant on hand and make every effort to always keep some of it.

Linda - Stout - 1936

*In 1938 Stout introduced Rosalind. In the introduction year, A.B. Stout described it as:
"The name Hemerocallis fulva var. rosea refers to a group of fulvous daylilies which have rosy pink tints of coloring in their flowers. The first of these plants came to the New York Botanical Garden in a collection of wild plants obtained in the interior of China. There were several of these plants and they differed somewhat in stature, in the width of the segments of the flowers, and in the precise shades of coloring especially in respect to the mid-zone. Some were almost without this zone of more intense color.

"The individual selected for illustration in color (Addisonia 15: plate 484. 1930) at the time this variety was first described, has flowers with a pronounced mid-zone of darker coloring. Historically, this plant is to be considered as the botanical type of the variety. Divisions of this individual plant, itself a wild seedling in origin, have been distributed to some extent. But these divisions do not constitute the variety rosea. Collectively they comprise a single clone which has the same status as any one of the many individual seedlings of the variety. But it is probably correct to say that no other seedling or clone will duplicate this one. It possesses individual differences in character that are of some significance in horticulture.

"In order to give these propagations a desired horticultural identity, I will here give to the clone the name Rosalind daylily. The divisions now offered for sale by several nurseries as Hemerocallis fulva rosea are, I believe, of this clone, provided the stock came from the New York Botanical Garden under this name. Plants of this clone have thus far been incompatible to self and intra-clonal pollinations. Hence, seeds set by the plants of this clone are certain to be from cross-pollination with other plants, and the seedlings obtained from such seeds will not closely resemble the Rosalind daylily. This particular plant can be multiplied only by clonal propagation. Under the name Rosalind one should obtain only divisions of the one plant. " cited from: Horticulture, 1938, vol. 16, p. 226 )"

The Rosalind daylily then appears to actually be a fulva seedling from the wilds of central China that has been vegetatively propagated and is one of the original clones brought from Kuling, Kiangsi in 1924. I have no idea if the plant I grow as 'Rosalind' is the same as what Stout would recognize as 'Rosalind', but it looks similar. Stout introduced many species clones, including many fulva clones. I believe, however, that the 'Rosalind' daylily got officially introduced because someone else had "introduced it to commerce" for Dr. Stout previously and without his knowledge. As you can see above, he explains that the introduction of Rosalind is to give provenance to those daylilies in commerce at that time that derived from his original importations.

The H. fulva rosea forms are, I believe, very important to the story of the modern daylily, as they likely opened the door for colors other than yellow, gold, orange, red-orange and red-brown to burgundy tones. 

*A search of the years 1930 to 1960 with the name Stout at the advanced search on the AHS Daylily Database reveals ten pages of introductions for a total of 95 cultivars.

*A search of the dates 1890 to 1960 reveals 8,622 cultivars registered in that time period. Thirty-nine (39) of those registered by 1960 are registered as tetraploids. 

*A search of 1890 to 1930 reveals 133 registered daylilies, including species clones.

*A review of the results of the search 1890 to 1930 (AHS Database advanced search using the dates 1890 as the 'not before' date and 1930 as the 'not after' date with no other search criteria selected) is very helpful in understanding what the next thirty years would be building on, as the plants registered and made available to commerce in this time period are the foundation that most other breeders bred from or were using in their own crosses with species clones to make their own lines during the period from 1930 to 1960. I would point out also that plants being called Hemerocallis aurantiaca and H. aurantiaca 'Major' show up with great frequency in many of the early pedigrees.

*The period from 1930 to 1940 shows 456 registrations, while 1940 to 1950 shows 2,844 registrations, while 1950 to 1960 shows 5,713 registrations. This shows a remarkable increase in introductions per decade.

*There were many daylily breeders from the start and the numbers clearly swelled greatly in the post-war years and there are many noteworthy breeders from those decades, but A. B. Stout had an especially important impact, as his introductions, of both species clones, hybrid seedlings and later-generation hybrid cultivars, represent the advent of a great many of the traits we are still working with today and his impact on the emerging hybrid garden daylily shouldn't be underestimated.

*LinksThere are several excellent sources of information on the work of Dr. Stout.

  1. A great deal of information about Dr. Stout can be found at Charlotte’s Daylily Diary
  2. Another excellent source of information on Dr. Stout's many daylily introductions is the Manatawny Creek Farms Stout Daylily Introductions pages
  3. New York Botanical Garden Libraries (Go here and search the term ‘stout’ to find the A.B. Stout archives - this is where A.B. Stout did much of his work with Hemerocallis), and for pictures of daylilies grown at the NYBG.
  4. Another great link is Pictures of Stout Introductions by Artur Jasinski.
  5.  One of the most important sources concerning the early years of species clone importation, hybridization and introduction is Dr. Stout's own book, published originally in 1934 and re-issued in 1986, on his work and the first few decades of Hemerocallis hybridization - Daylilies: The Wild Species and Garden Clones, Both Old and New, of the Genus Hemerocallis.
  6. Another excellent source of daylily history is A Passion for Daylilies: The Flowers and the People by Sydney Eddison

Commentary to come next time...

But I want to say a few words in closing. I note a heavy presence of plants being called Hemerocallis aurantiaca and H. aurantiaca 'Major' in a good many of the early hybrids. The 'Major' type is said to be a larger flowered clone that is less hardy than the original version and with no fulvous eye. We can't know exactly what all of those so-called 'aurantica' were. Stout mentions in his description in 'Daylilies' (A. B. Stout 1934, reprint 1986) that, "Several clones somewhat different from this one and of uncertain origin are sometimes called H. aurantiaca." The original clone of aurantiaca came from about 1890 at the Royal Botanical Garden (Kew), thought to be derived from Japan. This plant is fulvous, but considerably more gracile than the fulva group. H. aurantiaca is an evergreen that can show winter damage and freeze damage even in zone 6. Stout says, "...suffer somewhat from winter injury, but...plant soon recovers and so can be classed as hardy about New York City" (zone 6).

Of H. aurantiaca 'Major', Stout has this to say, "The so-called "H. aurantiaca major" is very similar to the H. aurantiaca in habit...but the flowers of (Major) are much larger and have no trace of fulvous coloring and the plant is less hardy."

Later in 'Daylilies', on pages 99- 101, Stout discusses 'Winter Injury'. It is an extremely interesting discussion of the hardiness, tenderness and growth issues in the daylily species and popular types in 1934. Stout goes on to point out which daylilies grow best in various climates and encourages (predicts?) the growth of evergreen daylilies in Florida. Again, he reports about H. aurantiaca 'Major', "...clone Major suffers so severely from winter injury that it usually does not survive the winters throughout most of the northern United States. Many seedlings obtained by hybridizing H. aurantiaca and H. aurantiaca clone Major with more hardy daylilies are not fully hardy."

I wanted to point these things out, but I don't want to make any rush to judgement. I think there were reasons the early breeders used aurantiaca a lot. First, they had it. It had been in the west since at least 1890. Next, it was tamer, more gracile, than the big, rambunctious fulvas. Then, its difficulty gave it a veneer of 'rarity' or 'exoticness'. As well, aurantiaca and Major bloom at the same time (generally) as Europa and some of the early yellow species, allowing for on-the-spot hybridizing or even accidental bee-crosses. Finally, several breeders at the time noted the large flowers of Major were just too good to resist. 

When you look over the dates above, you will notice that H. aurantiaca and clone Major are prominent, especially in many of Dr. Stout's breeding lines. I have highlighted all traces of aurantiaca in breeding records in the date section above in red. There is nothing wrong with those plants being used, but each choice creates consequences and can tip the balance one way or the other.  In addition to aurantiaca, we see thunbergii, flava and fulva with good frequency, and of course we also see H. f. rosea as a significant player from the late 1930's. Citrina was used enough to be considered foundational and middendorfii and dumortierii as well H. minor all play a role. All in all, there are said to be 13 or 14 species involved in the modern daylily. I suppose that depends on whether you are a lumper or a splitter. Next time we will look at some of the possible consequences of the early selections that went into the advent of daylily hybridizing.

Linda (Stout 1936) background 
modern seedling (Brian Mahieu Sdlg x Lily Munster) in foreground