What is a Daylily?

What is a Daylily?

Part 1

Thinking about the genus Hemerocallis

The Hemerocallis belong to the order Asparagales (often referred to as 'Asparagoid lilies') and is in the family Xanthorrhoeaceae. The order of Asparagales are considered monocots, which is one of the two large monophyletic groups most plants are divided into based on the number of leaves at time of germination: plants that usually only generate one leaf or one cotyledon, are generally considered monocots. The Hemerocallis are lilioid monocots, characterized by having colored flower tepals on various sizes and expressions of flowers. The hyperlinks in the text for Asparagales and monocots are linked to the Wikipedia pages for those terms. They are good repositories of information and you should read over them if the subject interests you.

All of the Hemerocallis species seem to originate in Asia, specifically from China east of the great deserts to the west and into northern and southeastern to southwestern China, down into the foothills and valleys of the Himalayan Mountains in Tibet and on down into northern India, then on east to Korea, Taiwan and Japan with species of Hemerocallis ranging from the extreme northern islands of Japan to the small southern-most islands of the multi-island country. The islands off the coast of southern China are also populated by Hemerocallis species. While I do not have any references to wild populations of Hemerocallis in Vietnam, Laos, Cambodia, Thailand or Myanmar, or further south into the islands of equatorial Asia - the Philippines, Malaysia and Indonesia, I would be surprised if some of the daylily species aren't grown in many of these countries. I do not as yet know the southern maximum for radiation of Hemerocallis species in the recent historic past - modern times before the beginning of hybrid Hemerocallis breeding for garden flowers. Nor whether that was through natural processes or through human intervention with several daylily species being good candidates for the production of food and easily transported as trade items. This is a subject I hope to gather more data on in the future.

At the other extreme some species are found as far north as Siberia. Species such as H. minor and H. dumortierii are found in the boreal forests of Siberia, in the sub-arctic. The Gobi desert may have cut off populations in the far north from populations further south, creating unique populations of the same basic stock. The species clone Hemerocallis fulva 'Europa' seems to have moved from Asia into the Middle East, the Mediterranean and Europe over many centuries, arriving in Europe in the 18th century or earlier, while the species H. lilioasphodelus has been in Europe even longer, specifically in Greece where it has been grown for about two-thousand years. However, it seems the origin of the genus Hemerocallis lies in Asia. It seems likely that these two forms made their way to the Middle East and Europe through trade, possibly through ports or the silk road.

The monocots are said to go back to the early Cretaceous period (perhaps as far back as 120 m.y.a.) and the Asparagales are thought to branch off from other monocots in the early Cretaceous, making them a very old branch of flowering plants from the monocot group. For this reason there is much diversity in both the monocots in general. There are a host of traits that make monocots different from other plants and there are traits that make the Asparagales different from other monocots.

It is not known exactly how long ago Hemerocallis emerged from the Asparagales lineage. I have found a reference to the genus in 'Early Cretaceous lineages of Monocot Flowering Plants - Bremer, et al., 2000. Their paper records their use of DNA sequence data to determine the ages of splits within the phylogenetic tree of the monocots. The chart in figure 1 shows the Asparagales go back past 100 million years in their arrangement. The Hemerocallis appear to go back to about 35- 40 m.y.a. and find their origin in the middle of the Paleogene period, the Eocene Epoch.

I also found a reference from the abstract from 'The historical evolutionary development of Hemerocallis middendorfii (Hemerocallidaceae) revealed by non-coding regions in chloroplast DNA by J. Noguchi, D.-Y. Hong and W. F. Grant - Plant Systematics and Evolution - Vol. 247, No. 1/2 (July 2004), pp. 1-22, to the split between geographical populations of H middendorfii. The abstract to the paper suggests that the populations of H. middendorfii were split apart when the Sea of Japan formed, separating the two groups - one in China and the other in Japan. The authors estimate this split using molecular testing and geographic history. They estimate that split to have occurred approximately 25 million years ago at the end of the Oligocene epoch, at the end of the Paleogene period.

We may have good DNA and geological evidence that Hemerocallis goes back at least to the end of the Oligocene epoch at twenty-five million years ago. The entire genus must be as old as the species H. middendorfii, if not older, perhaps even going back to the middle of the Paleogene period as suggested by Bremer, et al. 

Paleogene period/Eocene epoch

The Paleogene is characterized as the time after the Cretaceous extinction event where dinosaur populations end and mammalian and avian species arrive on the scene and diversify, the monocots and dicots diversify also. The phase of the Paleogene when the Hemerocallis have been suggested by Bremer to emerge is the Eocene. 

The Eocene began as a warm time in which much of the world was covered in tropical forests. It would end, becoming the Oligocene, in a greenhouse to ice-planet swing over the course of the Eocene. Swings between cold and warm continued into the Oligocene and throughout the Neogene, in which we are in the latest interglacial warm period.

The cooling during the Eocene is thought to begin around 49 m. y. a., which is somewhat earlier than Bremer, et al., suggest the split for Hemerocallis, and could suggest that Hemerocallis emerged during the beginning of the Eocene cold period. We certainly can't say for certain at this time when exactly Hemerocallis first emerged, but Bremer is suggestive,  and while new information could change the dates somewhat, the two pieces of evidence that I sight above seem to indicate a lineage of at least 25 million years ago or (most likely) much more, possibly with a target in the 35-40 m. y. a. range.

The evolution of the ancestral Asparagales that led to the Hemerocallis occurred in the vast tropical forests of the early, warm Eocene period. With the rise of the Eocene cold period the Hemerocallis-ancestors of those tropical forests would have had to adapt to the colder conditions and the early forms of Hemerocallis would have needed to be able to survive through cold periods, but also survive during the warm periods that have occurred between the extreme cold periods, with this cycle increasing about 2 1/2 million years ago during the rise of the Quaternary Glaciation (we are in the latest interglacial of the Quaternary at this time). In areas where tropical warmth occurs even during cold periods (such as glacial refugia during ice ages in areas toward the equator and below (or above) the boundaries of the glaciers), tropical species could persist even in the coldest ice ages of this period. In areas that have seen long cold periods followed by long warm periods, survival can be achieved by being able to express both deciduous and non-deciduous trait, with the expression depending on environmental triggers that allow the plant to grow or go into dormancy to conserve energy and survive very cold and very hot periods.

We still see this behavior in many daylily species, including but not limited to, many forms of H. fulva and the H. citrina types. During the time in which the Hemerocallis appear to have been around, we have seen extreme planetary climate change, many times, from glacial to interglacial periods of warmth and heat, with the worst cold glacial periods being in the Quaternary. During the cold periods there are generally some warm refugia. There were glacial refugia in both Old World and New World. Warm refugia occurred around the equator in the Quaternary in Asia, Australasia, Polynesia, the Americans with areas in both continents, and the Mediterranean including parts of Southern Europe to the Indian Ocean and the Persian Gulf in Western Asia. Much of Asia was warm. Glaciation at the height of the last glacial maximum (before 12-13 thousand years ago (t. y. a.)) did not cover much of Asia, descending down into Siberia, but not fully down into China or even Mongolia, though much of northern China would be temperate, great swathes of Asia were large, warm ice-age refugia. Glaciers extended much further south into the North American continent than they did in the Asian continent and they extended deep into Eastern Europe/Western Asia above the Current Black Sea. Refugia in North American would have extended from approximately modern day Cincinnati, Ohio south. 

It is also important to remember that in the glacial maximum periods, the water level decreases in the world's oceans, being trapped in massive glaciers, and many parts of the continents are exposed that are under water in our interglacial age. So in addition to the large modern day landmass that was not glaciated in Asia, there would have also been large swathes of lowland such as the Sunda Plain, which is now the site of the Sunda Straits, which is over top the Sunda Shelf, which was exposed during much of the last ice age up until about 12 - 13 t. y. a. Current Hemerocallis species elevations would have been significantly different only 12 - 13 t. y. a. One wonders how many species were lost under the rising seas at the end of the last ice age. 

The Hemerocallis genus may have survived by being generalists that could deal with both warm and cold conditions, high and low elevations and a wide range of soil types, while exploiting those conditions to generate nutrients (or make use of a wide range of available nutrient sources). The flora and fauna most capable of surviving geological and climactic changes tend to be generalists. Specialists are much more vulnerable because they have adapted to very specific sets of conditions to which they may not be able to survive without, while generalists can survive under a broader range of changing conditions. When conditions remain consistent for long periods of time, specialists can survive for long periods of time giving them the veneer of 'survivability', but if conditions change over vast geological scales, the form can decline being unable to adapt to long term changes. Asia, with its large refugia and large tropics would have led to some strains of tropically adapted Hemerocallis species (such as some of the specialized species from the southern-most, tropical islands of Japan - H. sempervirens, etc.). 

Other species, such as H. minor, which is still found as far north as the boreal forests of Mongolia and Siberia, may well have been in those ranges for many, many millions of years. Though they could also be from repopulation at any time during the Quaternary interglacials. They thrive very far north as a handful of other species do as well. The greatest number of species seem to fall in a swath across central and southeastern China, Korea, across to Japan and down to the islands off the coast of China (Hong Kong, Taiwan, etc) then over to parts of the Indian subcontinent and into the Himalaya, especially on the Northern side of the Himalaya which borders on Gansu Provence in the Southwest of China, which is well-known for plant production and its many types of anciently-cultivated plants and its many types of unique wild flora, as well.

Before the end of the last ice age, there was considerably more land above water all around the Chinese coast, up to the Korean peninsula and around Japan. Korea and Japan were much increased and much closer to each other. Many of the current day islands were all connected and connected to the mainland or large islands. The Japanese islands were one landmass at that time. That may indicate that the current populations descend from populations that were at higher elevations in the last ice age. Many current Hemerocallis species can survive in the wild in a diverse range of conditions. Many exploit high altitude ranges but also easily adapt to lower altitudes. Most daylily species seem to be able to tolerate low-water conditions, but the vast majority flourish when they are grown in less-dry conditions. As in the species, our modern hybrids show this same pattern wherein there are very few actual dry-thriving forms, but many forms that will survive a dry period and not reduce or cease to exist, but won't flourish or look very good. Foliage on both species and hybrid forms of Hemerocallis can look terrible, and this is a common survival strategy - for the foliage to die in severe conditions, either extreme heat or cold, as well as dry conditions. There are however amongst both species and hybrids those that suffer more when water rations are reduced below a certain point, while some few will retain an attractive appearance much longer under water rationing or drought. 

When we want to consider the current forms of Hemerocallis species that are demonstrably generalists, we want to look at those that survive and flourish in the widest range and seem to have done so for a very long time. This would include first and foremost the fulva forms, which are very widely distributed, especially forma 'Europa', which is now naturalized through many areas of the world. Many other forms of fulva exist in Asia and this is a huge group of often quite robust plants, especially those that are triploid. Fulva occurs in both diploid and triploid forms. They also qualify as strong generalists in many of their forms, able to survive in wild states, often forming large mats in wild conditions. In addition to the fulva group, the small yellow dayliies, called H. lilioasphodelus, H. Minor, and H. dumortierii are widespread throughout the world as well as in the wild in diverse ranges. 

The modern species are interesting, but are difficult to make absolute statements about. In China and most of Asia, there has traditionally been two major divisions for Hemerocallis types - orange and yellow, with a further division in the yellow group for small yellow and tall yellow. I would agree that this division - orange and yellow - seems to be a real point of division representing the evolutionary history of the Hemerocallis genus. Gene testing has shown there to be two major clades of Hemerocallis. Interestingly, these major clades fall along the 'orange and yellow' categories. One major split from the ancestral form of Hemerocallis lost anthocyanin pigments becoming self gold to yellow in various shades, with no eye an little to no midrib. When viewed in the ultraviolet spectrum, most of these "self" colored yellows and golds show intense eyes that glitter with color. But we cannot see them in our limited visual range. We perceive a self colored flower missing anthocyanin. We can't see it, but that doesn't mean it isn't there, in some way, even if suppressed in certain pigment ranges. This line split from the fulvous, anthocyanic line very early on and the two lines have evolved forward since. 

I have found three phylogenetic trees done in different research projects on Hemerocallis species using genetic comparison to group clades of relationship. These studies have been consistently similar in showing a deep and early split between the fulva family and the yellow family. The relationships within the yellow group shows two major forks within it. However, height is not consistent between any of these two groups of yellows.

The papers are as follows:
A google search of any of these titles should bring up the papers, references or places where the articles may be obtained.

DNA Fingerprinting in Daylilies - Genetic Variations and Relationships among Species - Part 2 
Daylily Journal Vol. 56, No. 3 Fall 2001 - Jeffery P. Tomkins - Clemson University Genomics

Phylogenetic Relationships of the Genus Hemerocallis in Korea using rps16-trnK Sequences in Chloroplast DNA
Man Kyu Huh *, Oh Sung Kwon , and Byeong Ryong Lee
Department of Molecular Biology, Dong-eui University, Busan 614-714, Korea
Department of Biology Education, Seowon University, Chungbuk 361-742, Korea
Received January 24, 2013/Revised May 20, 2013/Accepted June 28, 2013

Genetic and Phylogenetic Relationships of Genus Hemerocallis in Korea Using ISSR
Joo Soo Choi, Hong Wook Huh, Seol-A Lee and Man Kyu Huh

Department of Molecular Biology, Dong-eui University, Busan 614-715, Korea
Department of Biology Education, Pusan National University, Busan 609-735, Korea

We will look at this in greater detail soon when I write about the species from historical times.

So what is a daylily?

It is an Asparagales monocot of the genus Hemerocallis. The Asparagales contain many interesting plant families. In addition to Hemerocallis, the group also contains orchids (Orchidaceae), irises (Iridaceae), Hosta (Agavoideae), Amaryllis (Amaryllidaceae), Agapanthus (Agapanthoides), Allium (Allioideae), and of course Asparagus (Asparagaceae) among many others. The Amaryllis clade and the Asparagus clade are connected from a common branch that is very deep in the ancestry of the Asparagales and represent a deep split with all the other Asparagales families. The Hemerocallis comes from that other line. 

The Hemerocallis, along with all the Asparagales, are very old lineages of perennial flowering lily-like grass-relatives (monocots).  They are often generalists that can grow in many biotypes. There are two main groups within the Hemerocallis - those which show anthocyanic, water soluble pigment in the upper layers of the tepals, usually in orange in the wild species, characterized as the fulva clade. The other main clade doesn't show visible anthocyanin in the upper layer of the tepals, though eyes and mid-ribs are visible in many in the ultraviolet spectrum that many birds and insects (pollinators) see. It remains to be seen in the case of the pattern visible in the ultraviolet spectrum if the pattern is the result of a pigmentation or a refraction effect. To our eyes though, this second line of Hemerocallis appear to be single colored yellows and gold.

The Hemerocallis have been cultivated in Asia for millennia. The main use is for food in the form of dried flowers for soups and other dishes, and there are other food applications also. There are reputed medicinal properties and some research suggests this may be true. Many different forms of many species are grown in cultivation in many parts of Asia. There are likely to still be wild populations of species in many parts of Asia. Some of these could be escaped cultivations. It is very hard to discuss specifics of the species because they have been cultivated by man for so long. Many of the forms that we attempt to call "species" or even "clones" may represent garden forms, bred or (more likely) accidentally produced within captivity, rising to prominence for one or many reasons. The age of the genus, its wide-spread distribution, would have allowed for many interesting variations over the millennia, both in the wild and in captivity. We can see from the rapid advances in hybrid flower breeding within the genus has turned up tons of interesting genetic variations, all arising from the same species materials that we still have available to us. Those species then must contain a complex mix of genotype that is not being expressed in their phenotypes. The species are repositories of genes that are either blocked within their species or are heterozygous features that need to be brought to homozygosity in later breeding and selection that only we can apply in domestic garden settings.

The domestic lines that have risen from the species within the last 100 years or so are an amazing collection of art and selection, showing the vast range of genetics found within the genus. Soon we will look at the species themselves.

Hemerocallis fulva 'Europa' growing under bamboo of the Phyllostachys genus. This stand of 'Europa' has been growing here for over thirty-five years.

Looking Back - Statement of Intent

LOOKING BACK-

The Statement of Intent

I was reading over the Statement of Intent page here at my blog today. I was pleasantly surprised to find that it was all still applicable. As I am beginning the next cycle of my breeding program, I thought a good point to start out with, to close out the fifth year and start my sixth year and next cycle, would be to post this page as a blogpost - a reminder of where my focus has been and remains.

STATEMENT OF INTENT/DISCLAIMER

This blog, all of its posts and its permanent pages are to be seen only as my own personal musings and thoughts on daylilies. I love to grow daylilies, but I have very specific standards for what I expect from a daylily. No one should ever feel I am suggesting that they should share my feelings or opinions. I think there needs to be a lot of diverse people doing very different things with daylilies, as this can assure diversity in the genus and lead to truly unique cultivars and lineages. In this way, breeders can explore the outer limits of the daylily genome. I can only say to anyone reading these pages or posts, take what works for you and chuck the rest.

You might think of this blog as a diary that I am using to help me define and refine my breeding aims. These pages are not instructional. Think of them more as conversational. I am asking questions here and discussing my own experiences, not providing answers. My intention for this blog is to write about my own process of developing breeding lines of daylilies reflecting the traits I value. Those traits may vary widely from yours, or they may be identical.

I feel that in the process of developing base lines, I need to simply focus on what performs wonderfully for me, in my own conditions.  Much of my writing here will reflect my process of choosing the cultivars (and later seedlings) from which to build strains that reflect my ideas and ideal about what makes a good daylily. I will also share my experiences in breeding, strain development and in breeding and selecting for disease resistance.

All daylilies aren't created equal, and there are different types of daylilies. I generally work with three main categories of daylilies in my own gardening design, though there are many sub-categories and cross-overs, so I don't mean for these three broad categories to be seen as mutually exclusive.

Regardless of ploidy, the three major categories I utilize in my own garden for daylilies are:

1. Landscape/Borders/Background and Mid-range Fill - These tend to be medium to tall.

2. Specimen/Focal Point/Shock Value - The freaks - all the extremes of flower phenotype.

3. Border Fill or Specimen/Potted Plant - These tend to be very short/shorter to medium height.

Each of these categories generally utilize quite different daylily cultivars. While there is some cross-over, there may be instances where I find a given cultivar really only works in one of these categories. As an example, for me it is most common that those that belong to category 2 are the least likely to fall into category 1 or 3, but there are a few that do. Different family lines can be developed to fill each niche category, and thus different selection criteria could apply to each category.

Most of my writing will speak in terms of these three categories, though I will touch on selection for things such as disease resistance, vigor, fertility and/or rebloom in a more general way.

I have grown a lot of cultivars that for one reason or another, are not suited to one or two of my three categories. A few cultivars have a flaw or two that makes them unsuitable for any of these categories, in my own garden. Those cultivars (and later seedlings) are removed from my growing environment. I fervently believe that breeding from plants that show the traits you want, in terms of overall plant performance, is essential to producing lineages that reflect those desired traits. Others may choose to follow other criteria and a cultivar that is not suitable in my garden or my breeding program may work wonderfully for someone else in another environment. Never think I am suggesting otherwise.

I want to focus my discussion of cultivars to those that do well in my environment. Those that do not do well in my environment may do well in another environment or care regime. I do not want to seem as though I mean that a plant which has shown an elimination flaw in my garden is necessarily a "bad cultivar". Rather, they are simply not suited to my program. It is up to each individual to determine their own criteria of selection and their own tolerance-limits with any given flaw or problem. As well, each person must determine what cultivars will work for them, in their own site, for their own aims.

Daylily Domestication

Daylily Domestication:

A Process That is Still in Process

I have noticed this year, as I have observed the daylilies, that my mind has been drawn over and over to thoughts about the domestication process. My hobby is actually the study of the process of domestication, more than it is any particular species or 'genetics' in an abstract sense...specifically the study of the genotypic/phenotypic variations, breeding and selection of captive populations of plants and animals that are genetically changed from their wild type ancestors in various ways, is what most interests me.

When I observe the daylilies, I think about how far we have come in such a short time period. The advent of daylily phenotype breeding in the west began in the 1890's and from the start the early breeders utilized the fledgling science of selection and breeding - heredity - to guide their hand. The work of A.B. Stout is phenomenal and important. I am both surprised and disappointed how few daylily people have actually read Stout's book, or his many articles. His records are an unparalleled look into the earliest phases of a domestication process, detailing the hybridization of species into clonal lines and then away from their species roots to become very unnatural and very beautiful variations, family-lines that each reflect the genetic legacy of those original species in all their backgrounds, as well as mutations and new visual effects due to unique genetic combinations in the individual hybrid populations.

I want to say that I am awed and amazed by where I find the daylily today. In a mere century, daylily breeders have created a bewildering array of styles and variations, and they have done so on two ploidy levels. I am very proud of what we now have and I think all the extremes and oddities are a testament to the artistry and skill of the breeders who made them. I do not expect that pre-historic hybrid populations would have become so extreme as quickly as our daylilies have. I suspect that chickens took centuries to become something other than odd interminglings of two, three and four-way jungle fowl hybrids. It may have been hundreds or even thousands of years before various traits were put together in exclusive combinations in such a way that they would later become landrace breeds leading to the breeds we now know today. In the development of the modern garden daylily, we see a very accelerated development that is rather astounding, and I would also have to add: special.

But just because we see that fast progress, that doesn't mean we won't be susceptible to long-term trends. Provided that humans persist, I expect some will grow daylilies. Some will likely breed them as well, if anyone does grow them. So should daylily culture persist for millennia, what can they become? What amazing desirable traits can we combine and enhance and what might our oversights of bad traits have accrued into in the future? I always like thinking about the edges of the known in daylilies, as I suspect there are many directions as yet untapped. Those directions may never be tapped, but they are still there waiting. Any variation seems amenable to selection, if you start applying selection to it. But even beyond that fringe of thought, wild traits as yet unrealized, what does any of this mean for the day to day breeding of daylilies? I would suggest it means we need to pay attention to a wide range of traits and be very intolerant of undesirable traits.

I recently posted an article on corn domestication at my Facebook page. (The link is here. I highly recommend reading this article!) Corn and wheat domestication are very fascinating to me. Both are ancient events on opposite sides of the planet that came to the same basic outcome. There are other examples - each fascinating - but these two have always captured my interest, especially. So when I had first read the article a some time back, the next few days in the garden had my mind abuzz with thoughts of examples from corn breeding applied to daylilies. The early corn breeders weren't selecting for flowers, but they still made aesthetic and practical selection decisions that ripple down through the ages to us today. 

One of the major selection decisions they made was to move away from the wild-type seed-scattering phenotype and toward the desirable non-scattering seed head that allowed all the grains to be harvested at one time. In essence - a selection decision toward a specific seed phenotype. All the hundreds of domestic versions of corn and wheat tend to show these non-scattering genes to this day, so it is likely this selection decision dates from the early days of their domestication. It is in essence the basis of the domestic forms of these plants. This makes me think that if we apply selection to any traits where there are variations in our daylilies, we can select toward, and even intensify, desirable types.

But it is important to realize that selection is not just for the flower traits, even though that should always be a major point of selection, constantly pushing the boundaries and bringing out new traits. However it needn't be the only area of selection. There are many plant traits, seed traits, disease-response traits and environmental-adaptability traits that can (and should) be selected for. With that said though, I am not saying each breeder needs to select for every trait, just that they should be aware these things can be selected for and then choose those traits to select for that are of the most importance to themselves.

I fully understand how hypnotic the flowers can be. I have to actively remind myself to look at the plants when the daylilies are in flower. If daylilies didn't flower, I can guarantee you that I wouldn't grow them, but there is more to the daylily than the flower. There are so many traits in the daylilies that can be selected for. Try looking closely at your plants. Look at the traits other than the flowers and when you see a trait that displays variations across different cultivars (or seedlings), that is a trait to which you can apply selection, choosing those which display the variation of the trait(s) that you prefer. The selection possibilities are at least vast, if not unlimited...

Why Daylily Foliage Matters To Me

Why Daylily Foliage Matters To Me:

Using Daylilies in the Landscape

The foliage of daylilies is an oft-neglected area and one that is ignored in some circles, but daylilies can have beautiful foliage and can serve wonderful effects in the garden; in background groupings, in mass plantings, as a substitute for ornamental grasses (with the added bonus of flowers) - but not just any daylily will work for these purposes. Some daylilies are bred for fancy flowers alone and require more intensive care than the average homeowner or gardener cares to spend on their gardens. 

My first  fifteen to twenty years with daylilies were spent mainly working with daylilies as landscape plantings rather than specimen plantings grown strictly for flowers. Though I grew a wide range of daylily types, I quickly found that only certain daylilies were actually what I would consider 'good plants', by which I mean landscape quality plants. To me, a landscape quality plant is a fast multiplier that recovers quickly from division and increases quickly but does not decrease its scape count as the plant matures and has beautiful foliage. Thus such plants can be frequently divided to create groupings and mass plantings that can be very beautiful in the landscape and can be accomplished fairly quickly, and without a massive monetary investment.

However, such plants are often overlooked in the daylily hobby and are not reliably-enough identified in the general home garden market. We often hear that daylilies make excellent ground covers, slope covers for erosion control, and xeriscaping plants, but not all daylilies successfully can fulfill those roles, which can make buying daylilies confusing and sometimes disappointing.

Over the years I have gravitated toward using simple flowered, species-form and more primitive cultivars in order to find just the right balance of vigor, flower display and beautiful foliage. However, in recent years, I have begun testing newer cultivars, both dips and tets, and have begun to locate a few that may have the extremely fine foliage and garden habit that can make a long-loved garden classic.

I wanted to show you some of the uses I have made of daylilies in my own landscape. As you can see, these daylily plantings are not just about the flower. They are important parts of my landscape year-round. The flower is an accent to the foliage, which is the primary landscape use for these plants. We frequently see short daylilies like Stella De Oro used in landscapes as borders or edging, but I like daylilies more as a backdrop and as a substitute for ornamental grasses. For that use, daylilies that are more robust in foliage and scapes are most appealing to me.

Here in the entry to my house, you can see a mass planting of daylilies to the left.

Here is a more complete view of the entry with daylilies in mass plantings to both the left and right of the stairs that lead down to the deck and house. You can see they have a lovely, grass-like, textural look that is very pleasing and is a wonderful contrast to the red foliage of the Japanese maple.

A study in contrast.

The contrast of the daylily foliage and the stark concrete retaining wall is very attractive and softens the hard edges of the concrete wall. The daylily growing behind the wall and to the left in front of the wall are H. fulva 'Korean', which tends to show lovely lime green foliage that is very golden when it first emerges in the spring.

From this view you can see that behind, the Japanese maple is trimmed flat and creates a hidden testing area where modern cultivars are being tested for their garden value. The soil in this area is backfill from when the house was built and is basically red sand and red clay. In other words - horrid! So this is a marvelous testing area for vigor and toughness. You can also see the large clump of H. Fulva 'Korean' that is against the retaining wall and you can see how golden it is. I believe that daylily foliage shows the same range of possibilities for color that hostas exhibit. Daylilies just haven't been selected for it like hostas have. The daylilies against the wall to the left of the fulva 'Korean' are the old classic Hyperion. You can see the lovely dark green foliage that it has from its ancestor H. citrina. I love the contrast of the two foliage colors together.

A closeup on Hyperion, and in the background you can see a mass planting that includes several species and cultivars. Notice that the planting is split in half with lime-foliaged plants to the right and darker green foliage plants to the left.

Here is a closeup on the mass planting I just mentioned above. I did plant the daylilies to have the two foliage colors as contrasts, but the ivy and sedum did their own thing and also intensify the contrast pf the daylily foliage. The daylilies to the left are the dreaded 'ditch lily', H. fulva 'Europa'. They have been there for years and haven't eaten creation :-) There is a clump of Stella or two in front of them also, and the daylilies to the left are a mixture of "Nashville Star" (an unregistered G. H. Wild cultivar that is slightly root-spreading), Frans Hals, H. sempervirens and H. fulva 'Kwanso'. When daylilies are mass planted like this, they tend to shade out most weeds. 

Here is the view from the bottom of the stairs of the daylilies on each side of the retaining wall. Notice how the golden spring foliage of fulva 'Korean' contrasts so beautifully with the Japanese maple and the faded red paint on the railing.

This view shows the area of the garden on the west side of the house. This pathway leads up to chicken houses (that no longer have any chickens...) and the west gardens and then up to the top of the hill where the hybridizing garden and grow-out beds are located. This pathway used to be full sun before the bamboo ate it. I grew many daylilies along that path until the bamboo took over. I now focus on Hosta and Hellebores in this area. The Japanese maples have done well in both full sun and heavy bamboo shade. Many of the daylilies that I am showing in this post once grew along this pathway and were rescued from under the bamboo. The bamboo killed several cultivars in this area, so those that survived tend to be extremely hardy and flourish. Now the last two winters have taken a serious toll on the bamboo. I hope in time it will recover, though we will probably need a few mild winters for that to happen, but that's gardening - everything you plant doesn't work and things evolve.

Here is the western side retaining wall and the bed behind it. Both sides of the house have mirror image beds - both have the same horrid soil and I use both as testing areas. Anything that flourishes in these beds is very hardy and thus very useful for many purposes. This garden on the side of the house is steeper, and this area just behind/against the retaining wall had problems with erosion. I did two things that have stopped that. First, I added a McCourt rectangular tub and filled that with Louisiana iris and used a small McCourt three-tier waterfall behind the tub and in it I am growing Iris versicolor. These stop the dirt from being washed down over the bank and into the bamboo. The second thing I did was to cover the steep bank with daylilies. These have filled in well and have worked to retain the soil and stop erosion. It is also really beautiful, lush and requires very little weeding.

Here is the bank when the daylilies were first added in early spring of 2011. You can see the terrible soil, These daylilies are rescues from under the bamboo. They are "Nashville Star" and a fulva like cultivar I bought at Lowes in 2000 that was simply called "ground cover daylily".  It is slightly root-spreading, which is very good on this slope. 

Here is "Nashville Star" blooming in the summer of 2011, only a few months after they were saved from under the bamboo. You can see how much those little single fans I had planted that spring had grown in only a few months.

Here is the same planting in the spring of 2012. You can see the great growth of these plants. I did not fertilize them at all, as the bamboo will run into any area and get very vigorous if there is much fertilizer added. So these plants have to be very vigorous to flourish in this awful soil. This is essential for daylilies that are going into landscape situations and that won't be pampered!

Here is the planting from above in the summer of 2012. You can see the fulvous types at the top of the planting and the Nashville Star plantings below. The hot pink flowers to the right are the old cultivar Carmine Monarch, and it too survived under the bamboo for many years before it was rescued.

Here is the planting this spring (2015). I actually moved some of the Nashville Star that was directly behind the tubs, as they weren't necessary for stopping erosion (the plants up higher seem to do that enough on their own) and they were crowding the irises. They are now in the entry planting.

Here is the planting in the winter. I included this picture to show that these are all dormants and why dormancy is important to me in this type of landscape setting - no 'boiled-lettuce', ragged leaves struggling to grow all winter... The dead daylily leaves and tree leaves work to stop erosion through the winter. The look of this bed is very clean and open all winter - a stark contrast to the lush look of the growing season. I really enjoy that contrast.

Here is the planting this spring from the back. I included this to show how the clumps along the edge of the slope have become huge and serve to stop most of the erosion on their own, by channeling the runoff away from the slope.

Here is a wider shot of the slope showing more of the garden that is behind it, mostly daylilies, but also a few peonies. Peonies take years to establish in this horrible soil, so there aren't many in this garden. The tree peony has been there for over twenty years. I extensively cull poor performing daylilies in this area, and you can see the results in beautiful plants, as well I am gathering data on what daylilies make excellent landscaping subjects.

And here is the full view of the western testing bed from the northern view. In addition to being a great place to test daylilies for vigor and resilience, it is also very pretty year-round. That is because I vigorously cull those cultivars with bad foliage and poor performance and I focus on those that perform beautifully and there are almost no semi-evergreens or evergreens here so the bed looks clean and neat all winter. The lushness of these beds on the sides of the house is beautiful and relaxing - a lovely place to relax in the late evening after a hard day of working in the rest of the garden, and it is not just beautiful when the daylilies bloom.

This is a view of my hybridizing garden and pigeon houses (formerly chicken houses...). This garden was established in 2010. Even though it is mostly daylilies here, I have still approached this planting as a garden and with an eye toward pleasing aesthetics. I am very influenced by Asian gardens, especially Japanese gardens, so an ocean of moving, variable greens to me is just as pleasing as when the garden is in full flower, and that is the garden that I see for much longer each year than the garden in flower. Since most of my property is on a south-facing slope, I like the effect of the daylily foliage as it appears to almost cascade down the hillsides. The lush, green effect is in part due to culling out anything that showed ugly, unattractive foliage. Look at the beautiful lushness of this scene and you can see why daylily foliage matters to me as much as the flowers.

Here is the hybridizing garden in 2011, just to show you the progression of this garden.

Here is a detailed shot of lovely foliage in the hybridizing garden. The daylily cultivar in the front of the picture is Substantial Evidence, which has beautiful golden to lime green foliage, while the taller clump behind it is Notify Ground Crew. Both cultivars have wonderful foliage, beautiful flowers and tons of presence in the garden.

Here is a picture from summer of 2012. You can see there are still a few cultivars with less-than-beautiful foliage hanging around at that time. 

Here is a view of the hybridizing garden this spring. I took the picture while standing on the porch of the pigeon house. You can see that the layout of the garden, even though it is just a garden for hybridizing, is still designed to be attractive, have easy access and be beautiful throughout the year. I am in this garden every day of the year, all year, as I have to feed the pigeons, so it maters to me that the garden is attractive and the plants that I grow here are beautiful, even when not in flower. The area to the front of the picture that is surrounded in landscaping timbers is a seedling bed. There is one plant in the front of the row against the timber to the right that has very unattractive foliage. I have seedlings from it, many with equally bad foliage, but a few with much better foliage. That plant is unlikely to be here after this season. The other seedlings around it have much better foliage and much better prospects.

Another shot of the hybridizing garden. The shot also shows some of the seedlings in their testing tubs. Notice even they have nice foliage. That's because I removed and composted those that didn't have beautiful foliage. I love the sea of beautiful green foliage in the background.

Here is Challenger looking gorgeous and shading the base of a beautiful white clematis. The plant in front of Challenger is one of the least attractive foliage plants left in the garden, but it has very strong rust resistance, so it has been given a reprieve for the time being, but as soon as I get some better seedlings from it...

The white clematis in bloom today. There is much more to the hybridizing gardens than just daylily flowers :-)

And to close our tour of foliage and mass plantings of daylilies, here we are back at the west retaining wall and the daylilies that stop the erosion. I have to admit, this is one of my very favorite areas in the garden - simple, elegant, and beautiful year round. This is why daylily foliage matters to me, and these are some of my methods for using daylilies in the landscape.

Advice For The Beginning Breeder

A Little Advice For The Beginning Breeder

**I want to start this post with a disclaimer. What I have to say in this post are my opinions based upon my own experiences. If these thoughts don't apply to you or you have other opinions then please ignore what I have to say. However, if you feel there is something in what I have to say that you can apply, please feel free to do so.**

In my many years of breeding plants and animals, I have gained some insight into the process of beginning a breeding program. I want to share some of these observations with you, the reader, so that you may avoid some of the pitfalls and old-wives-tales that are so common in many hobby realms.

One of the first observations I would like to make is about obtaining breeding stock. The standard story is invariably that you must buy the most expensive and advanced cultivars (plants)/strains/lines (animals) you can afford. In plants the implication is that you must get the latest and greatest introductions and in animals you should buy individuals of the highest placing show lines, if you even want to bother with breeding. However, I have two objections with this. The first is that many beginners may not have the experience to sink a large amount of money into the 'finest' stock and may not really be capable of giving these (often touchy or delicate) examples the care they may need. Conversely, the second point is that beginners may gain much more valuable experience by not working with "the best" (as if the best show lines or the 'latest and greatest' introductions actually even are "the best" either in terms of hardiness and survivability or genetic potential). I think the whole notion of 'buying the best' is sometimes more about the sellers purse than the buyers needs, in some instances.

A beginner to breeding any plant or animal is in need of experience. Once some experience is gained, then he or she will have a much better idea about what they want to do and actually can do. So my advice to the beginner is to buy older and affordable cultivars and gain some experience, or in animals buy the second tier stock usually called "breeding quality" and gain some experience. In poultry, an excellent source of stock to gain experience with are those obtained from the commercial hatcheries. Unfortunately, breeders of other animals don't really have this luxury. Plant breeders, though, are in the absolute best situation when it comes to purchasing stock. Any older cultivar that is still in commerce can generally be obtained fairly easily and often fairly inexpensively, and there are thousands to choose from in all forms, styles and both ploidy levels.

Don't let anyone fool you into thinking that an older cultivar is useless. No matter how many offspring were or were not produced from a given cultivar in the past, those breeders didn't have the same gene pool to choose from to combine with those cultivars that we have today. Gain some experience with older, less expensive cultivars that have traits you like and then in time you can bring in more expensive, newer cultivars as you gain experience and combine over the best of those older cultivars to learn about breeding, the making of bridge plants and the retrieving of desired traits in later generations.

Don't assume that because a cultivar is new or expensive that it is superior. Some may truly be superior plants, but others are merely fancier flowers. Like everyone else, I use the occasional fancy flower on a lesser plant in a breeding project, but I only combine them onto the hardiest and most vigorous plants, generally older things that I have a lot of experience with and can trust to reproduce those characteristics, even when crossed to weaker cultivars or seedlings. However, when you are new to breeding any type of plant or animal, you might not be able to recognize the truly hardy and vigorous nor the weak but fancy. The inexperienced eye can be deceived by flashy and extreme phenotype traits. So can more experienced breeders, sometimes, as well.

It is important to remember that delicate and hard to maintain does not equal quality. This is an old-wives-tale that seems to be far too common in all breeding groups. Quality starts with performance and may be compounded by phenotype extremes. However, an extreme or beautiful phenotype in and of itself does not imply quality. These things simply imply an accumulation of genes, usually major and minor, for the trait in question. That is nice, but does not imply a well rounded cultivar or strain. When you are starting out, it can be hard to recognize the more subtle performance traits through the glitz of the accumulated phenotype extremes.

In daylilies, it is important to remember that not all cultivars flourish in all environments. Some southern cultivars do not do well in cold climates. Some hard dormants don't survive in the south. This is just reality. However, most daylilies will do fairly well in most conditions. We haven't managed to turn them all into hothouse delicacies just yet. With that said, the beginner is best advised to buy from a climate similar to their own and even then, buy those that are flourishing in a similar climate. Just because something can survive in a given climate does not mean that it will also thrive. What you want to gain experience with at first are plants that thrive, because you want to reproduce plants that also thrive. 

There will be plenty of time to add "breeder plants" that may not thrive but are still useful in crossing over plants more suited to your environment in order to bring in advanced genes of one kind or another. Once you have some experience, you will also have a better chance of using such plants well. Starting with such plants can be discouraging, but even worse, you might think that such plants are the 'norm' and never gain discernment of what a great plant really is.

Beginners with any type of breeding project are rarely going to jump in and produce an enduring classic in their first batch of seeds, so don't feel pressured to buy very expensive or delicate stock from the get-go. Beginners are often poorly equipped to care for such stock and don't have a point of reference so may not recognize the problem traits in such lines and may actually compound those problems without realizing it. With experience you can learn to recognize the problems and work to eliminate them from such lines while working to combine the desired phenotype traits with desirable performance qualities.

It is so important to not get in a rush. There is no finish line in breeding. If you get started breeding anything and stick with it, you will always be breeding, selecting, working to improve both phenotype traits and performance traits. That's what breeding is. It doesn't end. There is no point at which anything is 'perfect'. There will always be room for improvement. Take your time in getting started and learn what good traits, on both phenotype and performance, really are. Don't let people convince you that poor performance traits are ok because a line or cultivar is 'so advanced' or 'so whatever'. You might use a weak individual, seedling or cultivar at times, but you should be working toward improving the line by eliminating the bad traits while retaining the desirable traits. The bad traits are not ok and should not be overlooked just because of x,y or z phenotype traits. Don't ever believe they are acceptable, even when you are using such an individual or paid a lot of money for it.

By starting with tried and true stock, stock that has proven to have many good traits over many years and in many locations over the country, you will learn what good traits are and then can recognize them. Once you know what real performance traits look like in your garden, it will be very hard for someone else to pull the wool over your eyes and convince you that something pretty but weak is 'superior'. The point of gaining experience is to have a point of reference for future endeavors. 

Remember that everyone will have opinions. Almost any system can bring success if the breeder follows through in what they are doing, but 'success' can mean different things. For some success means they made a lot of money. For others, success means that they have taken a phenotype and made it more extreme, more concentrated. For some success is the production of well-rounded strains or cultivars that are consistent in good traits and have both performance and phenotype extremes combined into one package.

Beware of the notion of not 'reinventing the wheel'. This is something I have heard a lot in the poultry world for many years. The animal world is somewhat different from the plant world in that the animals tend to come in set 'breeds', so the admonition to not reinvent the wheel for animal breeds usually has to do with not trying to make new breeds or new varieties of existing breeds, but in plant breeding, homozygous strains that reproduce themselves in clone-like fashion are the exception, not the rule. So I was a bit surprised to occasionally find this same admonition in the plant world. Doesn't every breeder reinvent the wheel in some way or another? Isn't that the whole point? If we aren't reinventing the wheel, what on earth are we doing? Just churning out more of the same, maybe with a little bit more of this or that trait? There does seem to be a lot of that in almost all breeding circles. 

Sometimes, though, the wheel needs to be reinvented. When there are problems in a line, sometimes it has to be outcrossed and remade from the ground up. So many people seem to fear doing this and so in too many domesticated organisms, we see a steady decline over generations through inbreeding and narrowing of the gene pool. In many animals lines, the fear of outcrossing produces inbreeding problems and creates lines that are not viable, often with disease susceptibility problems set in the line, as well as infertility. This is less of a problem in plants, but we do see some of it. One area where the wheel might desperately need to be reinvented in the daylily world is in regards to the many types that show high rust susceptibility.

Another worrisome trend I have noticed in the daylily world is the tendency to blame failure with a given cultivar on the person growing the plant. This is especially prevalent when dealing with evergreen and semi-evergreen cultivars that fail in the north. Instead of just saying that maybe that cultivar is not a good choice for all cold climate areas, some will blame the failure on 'cultural conditions'. We accept that many hard dormant cultivars won't survive in the south, so why then is it sometimes not also accepted that some evergreen cultivars won't survive in the north?

In regards to the notion that certain cultivars won't survive without specific cultural practices, my question is, why then would I want that cultivar when the six hundred cultivars I have that surrounded it in the garden all survived without those 'cultural practices'? A beginner may not think to ask that question, but having had many years of experience with growing and raising plants and animals, I have learned that delicacy doesn't equate rarity, value or specialness. 

Now, I might use a delicate or demanding specimen (for as long as it survives or for one or two seasons) in a breeding program, but I am not going to think that delicacy is ok. I recognize that such delicacy is a serious flaw and I will work to eliminate that problem in future generations while seeking to keep the desirable trait(s) from such a cultivar within the line. However, you have to be able to recognize the flaws in order to not perpetuate them. To do that, you have to be able to see those flaws and even understand that they are flaws to begin with. Only by gaining experience with excellent performing individuals or strains will you have that point of reference to recognize the flaws when you see them.

While it may not be a popular thing to say, there are plenty of five and ten dollar daylilies that are wonderful, beautiful garden plants and are also excellent breeders, especially when crossed to excellent modern cultivars. Age is not a determiner of breeding value. If you need any further evidence, just look at some of the work of Brian Mahieu or Gil Stelter, both of who have gone back to the species in their programs with wonderful results. Talk about reinventing the wheel. Can you reinvent it any more thoroughly than going back to the beginning?

Another fine example of someone who 'reinvented the wheel' is Richard Norris, who created his line of flat flowered cultivars by using the old, flat flowered cultivar Lights of Detroit. Lights of Detroit had been around for years when Richard started working with it to produce a flat flowered line, yet almost nothing of any interest had been done with it. Richard had both the vision and the persistence to just do it. He had an idea and a vision and raised large numbers over several generations to produce what he was looking for. There must be so many currently obscure traits in the daylily gene pool that could be, but currently aren't being, exploited to produce amazing and breathtaking new phenotypes. We will never know if someone doesn't try.

It is important for beginners to study what has gone before them. Look at what other did. Don't just play follow the leader or jump on every latest trend. Try new things, but also, don't be distressed if your ideas fail. Sometimes our failures are our greatest teachers. I am deeply grateful for my failures, sometimes more grateful for the failures than the successes. Learn to value your failures and your successes, as both are your teachers. Listen to the experiences of others, but follow your own ideas, dreams, intuitions and learn what is possible and what is not possible while mapping your own path. If you do, you might just give the world something extraordinary. Above all, be patient, take your time to learn and don't spend more than you can afford. There is always time for that when you know what you are doing and are sure of what you want to do.