Sunday, January 14, 2018

The Daylily as a Permaculture Subject - Part 2

The Daylily as a Permaculture Subject 

Part 2
Selecting Plants for Growing and Base Plants for Breeding

Brian Reeder

*Disclaimer - (I am not recommending in any way, form or fashion that anyone should consume any part of the daylily. While Hemerocallis (daylily) is a common food and medical substance in Asian countries, if you have never eaten daylily or do not have accurate information on the consumption of daylily, it is completely upon your discretion when experimenting with consuming daylily. I strongly recommend that first time consumers be very careful in the amount of any given daylily they eat in order to observe the effects on their individual systems. Some people have experienced diarrhea from consuming daylily in quantity and some daylilies have been known to have sedative effects, which might be referred to as ‘poisoning’ or ‘poisonous effects’ by some sensitive people. The internal chemistry of each daylily, as well as each person, may vary and so no general information about daylilies as food or medicine can be made that will cover all daylilies and all human reactions/interactions with/to daylilies in any given instance. What is perfectly fine for one person may not be for another.)

In the previous installment of this series we looked at the traditional uses for daylily and some of the scientific research that is being done into the nutritional and medicinal uses of the daylily. While that look was in no way comprehensive, it is enough to demonstrate the uses of daylilies beyond the flower as a garden display, and it gives anyone interested some further reading suggestions to begin their own research.

In this segment I want to look at species and cultivars useful for those interested in growing daylilies for permaculture use, or who might want to locate base plants upon which to build a breeding program for any of those uses, whether solely for permaculture uses or even if just as a minor part of a daylily breeding program that encompasses other concerns with the flower as the major focus. I believe that the breeding and selection of traits valuable to nutraceutical concerns and permaculture applications do not have to be in conflict with breeding daylilies for floral gardens. For those interested the two can go hand-in-hand, as is true for any area of selection in a program. In spite of what we are sometimes told, it is possible to select for more than one trait at a time.

For those interested in daylilies in permaculture or food forests, the species and some of the hybrid cultivars may offer some excellent material to work with. However, if the garden has thrips and other insect pests, many daylilies from both groups will be problematic. In short, as the buds and open flowers are the major source of food and some of the medicinal uses, these insect pests damage your harvest, making them less palatable or destroying it partially or completely. The act of predation by these pests leave roughness, bumps, thorns and knots called enations on the bud, and in extreme cases, cause the buds to abort, sometimes even causing the scapes to wither entirely, loosing all the buds.  
Severe thrip damage to scapes on a highly susceptible daylily.

The outer shell of the bud is made up of the three outer petals (called sepals) of the flower. At the very least, there will be roughness on these and they can get very malformed due to this predation on susceptible plants. Buds that are ‘loose’, not tightly closed and perhaps partially open at the tip the day or two before the bud opens can allow these insects to get inside the flower and feed on the inner surface before the flower opens. It is my experience that while there are cultivars showing resistance to these insect predators, there seems to have been very little selection for this trait in the hybrid cultivars, probably because the majority of the wild species also show considerable susceptibility to these pests. However, there is resistance in some small number of hybrids and my own breeding work suggests that this resistance is heritable and can be intensified through crossing resistant individuals and selecting for the trait.

Spraying for pathogens is out of the question in true permaculture practice, and it is also out of the question in a resistance breeding program, so programs that do not spray, for whatever reason, are in the perfect situation to select for resistance to these pests. Many people reading this will be thinking that there are natural ways to control these pests, and there are. There are a number of insects that will predate these insect pests. However, in my garden, the heaviest thrip season falls across May and June, and with the advent of the heat of summer in early July I always see the activity of these pests go into decline. Unfortunately, the two major predators of thrips, aphids, etc, Praying Mantis and Lady Bugs, are not active at that time of year here. In my garden Praying Mantis and Lady Bugs are most active during the heat of summer when these pests have already gone into decline. This then defeats the use of natural predators. For me, the only path that has been very successful has been to select for resistant cultivars and then breed within them to select and increase resistance to these pest in the early and early mid-season flowering types. This is part of why I want to stress this so strongly. It is the only method that is really working for me, proving to be a real solution.

Here is a closeup of a seedling from my breeding program that shows extremely high thrip resistance. Note in this closeup that that the petal is literally swarming with thrips.

Here is a clump shot of the same seedling showing beautiful, clean flowers with no thrip damage and many buds on tall, well-branched scapes that also show no damage. This seedling also shows extremely high rust resistance. The parents are Ancient Elf x Solaris Symmetry (click the names to see information pages on these two cultivars). Ancient Elf shows extremely high rust resistance but thrip susceptibility, while Solaris Symmetry shows rust susceptibility but high thrip resistance. This seedling shows the desired combination of the resistance traits of both parents and demonstrates that such combinations can be achieved. In addition, this plant also shows the best foliage traits of both parents, being senescent with late emergence and beautiful, dark green coloring.

The same seedling above, here covered in seed pods. This seedling is very fertile, both ways, and so opens the door to a level of breeding that meets many of my personal goals. Note how deeply green and healthy the foliage is well after flowering has finished.

(Left) Ancient Elf and (Right) Solaris Symmetry
Parents of the above seedling.

While I love the many programs within the floral daylily community, I must say that going to modern hybrid daylilies for permaculture subjects is probably a loosing proposition, fraught with pitfalls. One runs the risk of wasting a lot of money on plants that require artificial care regimes, have little ability to compete in a more natural setting and tend to have little resistance to various pests and pathogens. As with pretty much everything else in the modern world, many floral daylily breeders practice ‘better living through chemistry’, and so have strayed for the natural, hardy roots of the species their domestic hybrid garden subjects descended from. While many modern hybrid daylily cultivars may be hardy and vigorous in the environments they are bred in, many are unlikely to be so in a permaculture or food forest settings. 

Mystic Butterfly
One of my favorite flowers I have ever grown. A truly stunning combination of floral traits - colors and forms.

However, when you pull away from the flower closeup, here is the reality of the plant in my garden. The foliage is always late-freeze damaged and the few scapes are always about ten to twelve inches tall with the flowers down in the foliage and seven to ten buds. Now, I am certain this plant flourishes in the garden in Florida where it was bred. I am certain in high-input care and/or warm-winter climates that it is a much better looking plant. I bet it would be stunning in a greenhouse. Yet, this is what it is in my garden. I am breeding with this plant in a salvage project though, because I absolutely love the flower and the plant does have some good traits here such as moderate rust resistance and it does not start growing at the first hint of warm weather. My goal is to bring flowers of this nature onto plants that are similar to the seedling pictures shown above. I want to stress again that this is not meant to be an incrimination of the breeder of this plant. I would not expect evergreen plants bred in Florida to show maximum performance in my cold-winter garden with average soil and low inputs. Such plants can't be selected for my conditions in vastly different conditions. That is simply reality and its not a bad place to reside.

Extreme inbreeding, over-focus on flower traits (and subsequently ignoring plant traits), the use of greenhouses and other artificial, chemically-focused, high-input growing environments and care regimen have caused the modern daylily cultivars to often be pale shadows of their wild ancestors in terms of survivability and ease of management. Never underestimate the ability of selection to do both good and bad things, depending on what is focused upon. While daylily hybridization and domestication has only been practiced in the west for a little over 100 year, that is ample time to bottleneck the gene pool and intensify deleterious traits through ignoring them in deference to the novelty of new visual looks in the flower.

Hemerocallis fulva Korean (Apps accession - 1984)
Growing in my hybridizing garden in two containers, which remain above ground all winter. Note the large, lush, healthy plants with many tall scapes. These two clumps are wildly pot bound, having been in those same pots for five years and I don't do anything to them, no fertilize or extra watering. The only thing I do is cut off the runners that try to come out of the holes in the bottoms of the pots on occasion. Containers are one excellent way to keep running fulva types in check. To read more about this fulva clone, visit my H. fulva Korean information page.

For the reasons cited in the last paragraph, and in combination with my own personal experience, I think there is actually little modern hybrid material that is suitable for use as a breeding base in developing daylily cultivars for permaculture settings. With that said though, I will also say that I firmly believe that any visual phenotype known in daylilies can be integrated into a program of breeding for permaculture use, IF the proper base plants are used to layer those visual traits upon AND proper attention is paid to a wide range of traits and proper selection pressure is applied to the seedlings. However, we do not have the wide range of data to tell us what effects, in terms of nutrients or chemical components, may be found in any given phenotype that has been heavily intensified through selective flower breeding. We don’t know, for instance, what effects the huge green throats that are so popular now many have. We don’t know what effect the selection toward blue-purple has intensified, or selection for bright, baby pink coloring may do to the nutraceutical traits. Personal and professional investigation is warranted to make further assessment. However, we know the species that are eaten and used medicinally in Asia, and so that is probably a good place to start.

Hemerocallis sempervirens
The latest blooming daylily in my garden, starting in Late August/September and going until frost in late October/November.

In my personal experience, the best base plants for breeding for permaculture uses are the clones of H. fulva (including the closely allied H. sempervirens) and H. citrina (including the closely allied H. vespertina).  Of the H. fulva clones, I find H. fulva ‘Korean’ and H. fulva ‘Hankow’ to be the most useful. The former shows less thrip resistance, but can produce resistant seedlings, especially when bred with other plants showing thrip resistance. The later is late blooming and so tends to miss the thrip season in my garden. It can throw both late blooming seedlings, and when bred with earlier blooming cultivars with thrip resistance, can also produce earlier-flowering resistant seedlings. Other fulva clones are probably useful, and I am currently testing out several other forms, but ‘Korean’ and ‘Hankow’ seem to be exceptional for breeding in my program. 

Of all the fulva clones, ‘Europa’, ‘Kwanzo’ and ‘Flore-pleno’ seem to show the least utility in any type of breeding, as they are near-sterile triploids. My experience is that I can’t produce enough seedlings from any of them to actually apply the type of selection I wish to make. Others may find them more useful. While ‘Korean’ and ‘Hankow’ are almost certainly triploids as well, they are quite fertile with both diploids and tetraploids in my own personal experience (and that of others) and so can be used in programs at either diploid or tetraploid levels. H. sempervirens may be its own species or it may be some form of fulva, but whichever it actually is, it is a good breeder and is very late flowering, even later than ‘Hankow’. Whether it has thrip resistance or not, I don’t know, but I have never seen problems with insect predation on it. This may be because of the late flower. It is probably a diploid, and I have produced numerous seedlings with it when crossed with pollen of diploid cultivars. I haven’t yet used it with tetraploid pollen, so can’t yet speak to that. Amongst its seedlings, some are earlier flowering and they also have not shown much insect predation, but that may be because they are still flowering too late to have heavy exposure to the problem. I do think H. sempervirens is useful, especially for breeding late flowering plants, and it is heavily branched with many, many buds and a long flower season that is later than any of the other species I know of.
A clump shot of H. vespertina showing the extremely tall and branched scapes and the large plant. A late bloomer, typically starting in August and going for about six weeks.

A closeup of the flower of H. vespertina on an immature plant showing less branches, buds and lesser height of scape than is to be expected on a mature clump. The flower is typical of the mature plant.

Amongst the yellow types, I have found all the clones of H. citrina I have grown to have high thrip susceptibility. H. vespertina, which may be a form of citrina or may be its own species, is far superior to all the citrina clones I have grow, perhaps because it blooms quite late in the season. It is very tall and heavily branched with many buds. Seedlings I have produced from it have also tended to show resistance to insect predation, however, this again many be due to their flower in July and August. The yellow species are thought to be diploid. While one might get an occasional set from tetraploid pollen, this would be a very rare event. I haven’t yet tried H. vespertina with tetraploid pollen, and I spent an entire summer putting tetraploid pollen on H. citrina clumps to get two pods with seventeen seeds. While one might persevere and create plants with chromosome counts above the diploid level using the yellow species, generally they are easiest to use at the diploid level.

Notify Ground Crew early morning 8 AM

Notify Ground Crew afternoon 5 PM

Notify Ground Crew at sunset 9 PM

Notify Ground Crew at night 11 PM

For yellow near-species/species-like hybrid cultivars my favorite cultivar as a base plant is Notify Ground Crew (click name to see information page about this cultivar). It is a large plant with tall (6’) scapes, well branched and with many buds. It shows much greater thrip resistance than the citrina clones I have grown, and it descends from a tetraploid conversion of H. citrina made by Orville Fay (via pod parent Tetrina’s Daughter) on one side and tet. Purity on the other as pollen parent, which is also close to species. Notify Ground Crew is a great breeder and I have produced many seedlings from it that also show strong thrip resistance. Another useful hybrid cultivar, descending from two converted species-like yellow parents, is Ancient Elf, though it has fairly high thrip susceptibility in my garden. Crossed with plants showing high thrip resistance, it can produce much more resistant offspring and it has many other good plant traits including very high rust resistance.
In addition to thrip and other insect resistance, most of the species-clones and cultivars I discussed above tend to have moderately high to very high rust resistance. This may be useful for some programs where that is also a concern, such as in the deep south or anywhere that daylily rust can survive and there is no winter freeze to kill the rust fungus. 

A daylily plant showing heavy daylily rust infection. Such a plant is susceptible to the rust fungus. Some plants show resistance to this fungus. For more information on cultivars resistant to daylily rust and breeding for rust resistance, click here.

Another point about all those mentioned above is that they can be grown throughout a wide range of climates. Daylily cultivars are registered as ‘dormant’, ‘semi-evergreen’ and ‘evergreen’. These categories are very loose and are not botanical descriptions, but are hobby shorthand. While some think these are hard-and-fast categories there are no true, binding botanical rules as to what gets registered how, and so they are not reliable guides. However, plants having leaves that die in fall and then form a resting bud may require some cold and many of these fail in southern gardens. Some plants retain some amount of foliage, green, above ground throughout the winter regardless of climate and only vanishing to or near the ground when they are frozen off. Many of these are hardy in cold climates, but others are tender and are harmed or even killed by freezing conditions. 

This picture shows a clump of Ancient Elf in the winter. Note that you can see the name tag, but you don't see any plant. This demonstrates the full senescence of the foliage. The resting bud is below the dead leaves and just at or under the ground level and will begin to emerge when spring commences, showing moderately late emergence allowing it to skip many freeze/thaw cycles. Daylilies of this nature may fail in areas without sufficient winter cold.

The fulva clones and citrina clones are adaptable. In cold climates they tend to go into dormancy, with only a tiny amount of plant above ground or only a resting bud, allowing them to thrive in very cold climates. In warm climates they tend to be ‘semi-evergreen’ or ‘evergreen’, and this also allows them to thrive in even very warm climates. For this reason, these species are recommendations I can make for almost any climate. Notify Ground Crew behaves much as the citrina and fulva types, though it is registered as ‘dormant’. Of all those I mentioned above, H. sempervirens is the most evergreen (sempervirens literally means ‘evergreen’) and while it survives in the far north, it doesn’t thrive, and Ancient Elf is the most dormant, with foliage that dies back in fall forming a resting bud for a period of time. It may not thrive in the deep south. While the species clones and hybrid cultivars I have mentioned here have strong applications for permaculture settings, they also have excellent applications in floral breeding programs, opening the gene pool of inbred and bottlenecked lines and possibly bringing genes not found in the modern hybrids into those lines.

Here is a picture of an evergreen type in my garden in winter. Some evergreen daylilies are hardy in my garden, while others are not and are referred to as 'tender'. These daylilies tend to attempt growth at each warm period throughout the winter and early spring and can sustain repeated damage that can reduce their performance. Such daylilies prosper in warm-winter climates such as the deep south of the US, and while some are perfectly hardy in the cold-winter climates, some are tender and not able to thrive or survive in repeated freezes. Ironically, some of these evergreen types do better in the far north than they do in my zone 6/7 garden, as some northern climates provide an insulating blanket of snow throughout the winter, protecting these plants from beginning growth repeatedly before spring has fully arrived. My area never has the long-lasting, insulating blanket of snow.

In part three we will look at going beyond the base plants and developing programs targeted at specific phenotypes.