Monday, May 16, 2016

Daylilies in Asia

What is a Daylily?

Part 3
Daylilies in Asia

Once mankind began engaging in agriculture and became settled, our adventure with changing wild species of plants and animals through the process of domestication was set in motion. While hunter-gatherer peoples certainly could impact natural environments and the species of plants and animals within those environments, there is no evidence of any active domestication by hunter-gatherers. This process apparently only fully arose later as humans became sedentary, living in settled communities.

The beginnings of agriculture is generally attributed to the time after the Younger Dryas at around 10,000 b.c.e. or later. The first domesticate is considered to be the dog for animals and wheat for plants. While dog domestication appears to have begun much earlier when humans were still hunter-gatherers through a unique set of circumstances, the domestication of wheat is thought to have occurred in modern-day Turkey after the Younger Dryas. 

Rice was the first major plant domesticate in Asia, generally attributed to 8,000 - 7,000 b.c.e. However, some evidence suggests that rice was domesticated earlier, perhaps are early as 16,000 to 12,000 y.a., though this may represent wild-collection of rice by hunter-gatherers, or perhaps a transitional phase (protoculture) from collection of wild populations of grains to full domestication. What is clear though is that the main body of the work toward domestic strains of rice occurred between 7,000 and 4,000 b.c.e.

It is clear though that rice was not the only domesticated crop raised in early Asia, and plants have been domesticated in Asia up into modern times. Domestics deriving from Asia include Foxtail Millet (which may go as far back as rice in Asia as a domesticate), Bottle Gourd (also as far back as rice), Water Chestnut, Perilla, Burdock, Broomcorn Millet, Garlic, Hemp, Soybean, Peony, Chrysanthemum, Azuki Bean, Chenopodium, Eggplant and Edamame, just to name a few.

Many other plants are utilized in Traditional Chinese Medicine and the historic herbal medicine that proceeded it, but were not domesticated, historically having been gathered directly from the wild. Many of these plants are now grown commercially to meet the demand for them as wild populations dwindle while others are still collected. In between those plants that were fully domesticated and those that were gathered directly from the wild were plants that were grown in captivity, but were not actively selected in the way the true domesticated plants were to change them to meet the needs of the growers. This is generally considered 'vegiculture'. The Hemerocallis seems to fall into this middle group historically in Asia. That is, they were grown in gardens, but they were not significantly changed from the wild types.

Domestication of plants has generally fallen into two main categories - annual crops that are generated through sexual reproductions (seed generated) and those that are perennial crops and are generated through vegetive reproduction such as division, grafting, rooting, etc (vegiculture). With annual crops, there is a complete turn-over of the genome every year with the potential for genetic change that sexual reproduction on such a scale implies, allowing for selection to be applied over time and causing radical changes from the wild predecessors. 

Perennial crops are not always produced through sexual reproduction, and perennials that are easily reproduced through vegiculture may rarely be grown from seeds. This makes the changes one may find in annual crops rarer in perennial crops historically (except for certain examples such as Peony or Chrysanthemum that have been actively bred for their flowers in addition to other uses), and thus little selection may actually be applied to these perennial species. For this reason, such crops may remain very close to the original wild forms for long periods of time even though they have been grown as a domestic crop into antiquity. So it seems to be with the Hemerocallis.

We know from ancient records that Hemerocallis have been grown in gardens and used as food and medicine in China for at least 2,500 - 3,000 years. It seems likely to me that their use goes much further back in time. I would suspect they were a menu-item for the hunter-gatherers in pre-agricultural Asia and as humans became settled, beginning to practice agriculture, they brought wild Hemerocallis species into their settlements both for utility and beauty.

While I cannot say why the Hemerocallis was not treated as Peony or Chrysanthemum in terms of selection for flower type, I might make the following suggestions on why so little modification was made from the wild species in terms of the uses of Hemerocallis as food and medicinal crops.

First, the Hemerocallis seems quite suitable for the needs it was used for. Unlike the shattering seed heads of wild wheat, the wild Hemerocallis species have no traits that make them difficult to gather. As the young shoots, flower buds and spent flowers are the main sources of food from the species, there was simply nothing to modify there. The species were suitable as-is in these regards.

Being easily increased through vegetative reproduction and especially with the running habit of the fulva forms, it seems likely that little sexual reproduction would have been actively pursued, with seeds (and new seedlings) being more of a random occurrence than a regular phenomena. When we add the self-sterility of some species forms and the difficult fertility of the triploids, these factors may have helped to make sexual reproduction of Hemerocallis a rare event in captivity. That doesn't mean it didn't happen on occasion, either by chance or intention, but the lack of forms drastically different than the wild species seems to point toward sexual reproduction being rare over the course of the centuries that this genus has been kept in captivity.

To add to the above, with self-sterility, in situations where entire communities were growing vegetative clones of one form, the chance of sexual reproduction becomes even more slight. While some individual gardeners/farmers may have known to cross forms to get seeds (or were growing multiple forms and simply got seeds through pollinator action), and there may have been individuals actively engaged in this activity, it was clearly not the common event as in Peony or Chrysanthemum, or annual crops such as rice, millet or wheat. 

When we look at the changes made to the genus Hemerocallis in a mere century in the West, we can see there is considerable genetic diversity within the genus. The lack of that phenotypic expression in pre-western garden forms in Asia, I feel, is another datum pointing to little active breeding in Asia throughout the centuries. Clearly, much change was made to Peony (especially the tree peony) and Chrysanthemum from their wild progenitors over the centuries in Asia, so we know the process of breeding and selection was being utilized historically in eastern Asia. I can only surmise that there were reasons the same was not applied to the Hemerocallis.

It certainly isn't  that they were unpopular. They were a common garden flower and food/medicinal crop for many centuries, with much lore and myth attached to them, as well as there being a good knowledge of the uses of the plant. So I can only suspect that a combination of factors kept the Hemerocallis very close to the species-types throughout centuries in Asia. As I have suggested above, I suspect the growth habit and fertility issues are at the basis of this lack of change within the genus in captive situations.

However, none of this is to say that no changes occurred. 

Areas where I think some change is implied occurs within the two main branches of the genus - the oranges and the yellows. 

In the fulva group, the "oranges", there are many forms known in Asia. While any of these could be wild-occurring regional variations, chance mutations or interspecies intergrades, some of them may well represent forms derived from garden-produced seedlings. Certainly, the diploid forms of fulva are fertile, and even when self-sterile, they can cross with other forms of fulva and other species of Hemerocallis, as we have seen from the work of Stout and others within the last hundred years. This would open the door for, at the very least, chance seedlings if not actual intentional production of new forms. 

Areas where domestic selection may have occurred is the doubled tepals of 'Kwanso' and 'Flore pleno', the variegated foliage of some 'Kwanso', the more pink or red flowers of forms such as 'rosea' formas and 'Chengtu', etc. While any of these traits could have arisen in wild populations and simply been collected and brought into captivity, these are also the types of ornamental traits one might expect to find favored and valued in captive-produced seedlings. Further, even where these traits may have come from wild-collected specimens, any garden where wild-collected specimens were brought together to grow would have been fertile ground for seedlings to emerge carry variations of these traits, and thus allow for captive selection to be applied. 

None of these traits would have inhibited the use of such specimens in their traditional food/medicinal roles, and in fact, all the above forms mentioned are grown in Asia for these traditional purposes, as are the other fulva clones. My experimentation with the forms of fulva I grow show that each form has a slightly unique flavor - an important point in a food plant and a culture with a highly developed cuisine tradition. Some clones are grown exclusively in some areas, while in others multiple clones are utilized. 

Another point to consider is bloom-time. By growing clones that are early (Europa), mid-season (rosea), late (Hankow) and very late (sempervirens), a much greater supply of food would be available throughout a much greater portion of the year. The same effect could also be achieved by growing both yellow and orange types that flower at different times throughout the season.

The triploid forms of fulva imply that the diploid versions on rare occasions produce unreduced gametes. Thus the triploid forms could have arisen in the wild through chance or in captivity through chance. However, the presence of the triploids, with their low-fertility, robust size and rambunctious rhizomatous growth habit may have made the chances of new seedlings that much slimmer, as the triploids are cultivated extensively in Asia and would have been a natural choice for their superior production of larger shoots, buds and flowers.

In the yellow category, many species are attributed, but how many of these are really species or wild-occurring species hybrids and how many are captive hybrids and forms? That is a difficult question to answer. First, it is very difficult to determine what the actual species are. Second, while some species forms show self-sterility, not all do, and some of those that do show self-sterility are sterile with their own clone, but not with other individual clones of the same species.

When we look at the yellow types grown in captivity in Asia, there are forms (just as we see in the fulva complex) that have not been shown to occur in the wild. For instance, while wild H. citrina is known, there are many clones of citrina in captivity. Likely some of the yellows in culture are clones, intergrades or hybrids of the true, wild-type yellow species.

Within the yellows, there are certain traits that may have been enhanced in captivity. Two that spring to mind are branching/bud count (important traits for the production of flowers as food) and the length of tepals in certain types such as some H. citrina forms (i.e., 'Baroni', etc), which would also give more food per bud. Such selection would fit with the uses of the plant and it is important to remember that in China there are only considered to be two basic types of yellow daylilies - tall yellow and short yellow.

H. citrina clone showing very long buds/tepals

A final point to consider is that there may have been many garden forms of Hemerocallis that have been lost throughout the centuries. Eastern Asia, especially on the continent, has seen many wars and much upheaval and destruction during its long history right up into modern times, and there is simply no way to know what wondrous forms of the many domestic plants and animals may have been lost over the centuries.

While it is impossible to make any definitive statements on the specific origins of any particular type, we can say that the many forms of Hemerocallis that were found in captivity and in the wild during the nineteenth and twentieth centuries in Asia provided the materials that Western horticulturalists would use to produce the amazing array of daylily cultivars that we know in the West today.

Next we will look at the early hybridization of the Hemerocallis that set the ball rolling in the West toward the amazing array of garden daylilies that we now know.