Hemerocallis Fulva in our Tetraploid Breeding Program

Note:   'F1' refers to the first generation of a cross of Fulva with something else.  'F2' is an 'F1' crossed again to another daylily, and so on.  

 Why in the world would anyone breed with Ditch Lily?

When I gave my 'Fulva Presentation' to the Michigan Daylily Society, Ginny Pearce asked why I started this program and  I was not able to vocalize a coherent response, though we are ten years into this, having started in 2013.  I guess my best thought is 'why not?' or just to see what I could get from it. Yes, Fulva is orange, but that is a 'plus' in my book. It is also hardy and healthy. It's the source of color for all modern daylilies that are orange, red, pink, and purple, as well as brown.  So, why would I not give it a try?

'Fulva' is not a single uniform species

Most people assume that the 'ditch lily' is the one and only representative of Hemerocallis Fulva. This is not true.  The species is actually a fertile diploid, and the ditch lily is a several thousand-year-old garden variety.  It is a nearly sterile triploid properly called Hemerocallis Fulva 'Europa'. The double forms,  Green Kwanso (Flore-pleno?) and Kwanso variegata are also garden selections of the species, both are also sterile triploids.  'Korean' is a triploid garden form that was collected by Darrel Apps and Barry Yinger in South Korea in 1984.  It is not a registered variety.  Some other Fulvas are Hankow, Cypriana, Rosea, Rosalyn, and 'Switzerland' and 'Yunnan', both collected by Gil Stelter, (a Canadian hybridizer who is at the forefront of breeding with Fulva). Several of these varieies are fertile diploids, but most of them are naturally triploid garden forms. So, there are a number of distinct forms of Fulva, all selections of the species. The triploids were likely often selected as garden forms because the extra genes give them a taller, sturdier scape. 

Fertility of types of Fulva

The variety commonly called ditch lily, Fulva Europa, is triploid and is nearly sterile.  I have never found any record of seeds being set on it naturally, despite the fact that the plant exists in the hundreds of thousands on roadsides.  However, Arlow Stout, the father of modern daylilies, was able to get a small number of diploid seeds from it after making thousands of crosses.  These formed the basis for modern hybrids of all colors.  Originally all daylilies, including Stout's hybrids, were diploids until the late 1940's when colchicine was used to create modern tetraploids. The diploid species such as Rosea are all fertile with other diploid daylilies. Interestingly, several of the triploid forms are fertile with both diploids and tetraploids. This is true of Europa, which has been used to breed modern diploids and tetraploids, both as a pollen and seed parent.  I have never been successful with it, though Stout used it for diploids and a number of hybridizers have used it since in both ploidies. However, Cypriana, Hankow, Korean, Switzerland, and Yunnan are all much easier to use as seed parents if modern tets are crossed with them as the seed parent.  Most yield from one to eight seeds per pod, which is not that far below average for many modern tet crosses. Note that some seeds in almost every pod are sterile and hollow, leaving maybe an average of three good seeds per pod. As far as I have been able to determine, Hankow is a diploid, but will cross with tet pollen and give good seeds.  The others are triploids. As Jurassic Park's Ian Malcolm said, "Life will find a way."  Plants can survive being triploid (animals cannot!) and can produce gametes that are compatible with tets from diploids from unreduced gametes and from triploids from a hit and miss division of an uneven number of chromosomes. 

Hybridization Results

Overall, I found that within two generations, Fulva crosses  produced clear colors to rival any modern daylily, and eyes and edges, too. I have worked chiefly with round forms and I did find that the bloom size stayed on the small side for the first several generations, but I did start seeing larger blooms in the third generation away from Fulva. Gil Stelter (mentioned above) worked mainly with large, open-faced blooms and he created many that are of the same large size as modern tets within one or several generations. 

Aggressive growth?

One commonly expressed concern with growing any hybrids of Fulva is the fear that they would be as aggressive growers as the species.  This has not turned out to be true.  None of the F1, F2, or even F3 hybrids have shown any rhizomatous growth. This seems to be totally lost in the offspring. Some of the vigorous growing varieties do produce several new fans per season, but so far these seem to be produced near the parent plant and show no more of a tendency to run than any fast-growing modern hybrid.

Seed Size and Enhanced Fertility

Any grower who has examined both diploid and tetraploid seeds knows that they usually vary in size, with diploid seeds being almost consistently smaller.  In breeding tets to our fulvas, it is interesting that the pods contain a mixture of seed sizes, though all are tetraploid or possibly triploid, most of which won't germinate. Some seeds are small and others appear from their size to be tet seeds, that is, the same size as modern tet seeds.  No actual follow-up has been done to see if the smaller 'tet' seeds germinated and grew, however.   

It has been of interest that subsequent generations to the F1, crossed to standard tetraploids, seem to have enhanced fertility over regular tets, producing large, round, black seeds of the highest quality. Brian Reeder (Sun Dragon Daylilies) has also noticed this and has written quite a bit in his blog about it. 

Seedlings and Results

Fulva, more than many species, seems quick to shed it's 'species look' and 'go modern'.  Maybe this is not surprising, since Fulva is the basis of all modern daylilies and the species has an eye, slight bicolor look, has sported to a double (Kwanso), and shows a tendency to have a ruffled edge.  None of these characteristics have been observed in other species, and I attribute the existence of toothy edges and eyes to Fulva.  So, what have we seen in our seedlings? 

As expected, crosses to fulva in the first generation did produce a lot of oranges in various shades.  However, it was interesting that some desirable characteristics from the non-Fulva parent started showing up in this first generation:

Fulva Cypriana x Better by Design

The thought process of making this cross was to get a Fulva hybrid with a watermark. Though it does not have a watermark, it does show a ruffled edge and great color in an F1 cross. Crosses with Cypriana tended in general to show rich orange to scarlet colors, not much of an eye, and pronounced tenderness often passes from the tender tet parent.  Apparently Cypriana is not sufficiently hardy to overcome the evergreen tendencies of the other parent,  Brian Reeder also has observed an extreme susceptibility to rust, also in Cypriana and its offspring.  So, on to bigger and better things...

Fulva Yunnan x Samsational (F1)

This one has fabulous color, though not a fancy face.  Definitely a keeper!  This was its first year to bloom, so I'm not sure how tall or floriferous it will be, but love that color!  Being the offspring of two hardy plants, I am sure this will be more hardy than the one above.

Fulva Korean x Mitch Lynd (F1)

I have always like bicolors and was surprised to find one in my first couple of crosses when the other parent wasn't a bicolor.  It has since been used to cross with other bicolors.

                                              Fulva Yunnan x Empire of Dawn and Dusk (F1)

This F1 seedling was exciting!  It is 50% Fulva, and has clear, golden yellow color and an edge and an eye!   Used quite a bit to carry on the program. This cross shows how compatible Fulva is with those bred from it originally, that is modern tetraploids.  An F2 seedling from it follows...

(Fulva Yunnan x Empire of Dawn and Dusk) x (Jules Michael Vinkman x Empire of Dawn and Dusk)

Here is an F2 from the above seedling, trying to lighten the background to white and adding more color to the edge.  Success!

(Fulva Yunnan x Empire of Dawn and Dusk) x Chico Tim

This one is probably our best yellow/red effort to date among the Fulvas.  It is a cross of the F1 above with a fancy one from Tim Bell that has proven hardy for us. 

Fulva Korean x Mitch Lynd (F1)

Not all F1 hybrids ended up as a shade of orange.  Here's a dusky (muddy?) one in a dark shade. More work to be done on this one!

Fulva Korean x English Lavender

While showing none of the lavender I envisioned, this F1 seedling does show how amenable Fulva can be toward giving a larger, more modern form when it meets the right partner.  This one was an outlier in size for an F1.  As is usual in hybridizing, I got some features I wanted (form and size), but not the color!  

As far as breeding for lavender is concerned, remember that all purple pigment in daylilies was originally from Fulva, hence a bit of muddiness in the species (hence 'fulvous') since the orange is overlaid with some purple. Unbelievably, also, orange and purple 'play well' together...when crossed, you don't end up with muddy color, which I would have expected, but the pigments tend to sort themselves out in the offspring.  I attribute that to them both pigments originating with Fulva.  Check out the following seedling:

Fulva Korean x Queen's Circle ('Korean Queen' from Brian Reeder) x 

(Gus Townsend x Desire of Nations #3)

This one well illustrates the paradox of orange + purple =  orange or purple... and not mud!

It's the offspring of Korean Queen (F1), which is orange (from a cross of orange and purple),  crossed with a rich, fancy deep purple seedling of ours. Interesting result!

      

         Seed Parent, Korean Queen                                  Pollen Parent, Gus Townsend x Desire of Nations 

                                                                                                                      (seedling)

(Fulva Korean x Empire of Dawn and Dusk)  x Sonoran Desert Rose

The color could be cleaner, but this one is here to show that doubles show up spontaneously at about the same rate as they seem to in our other programs, working with modern daylilies. As with most of the second-generation Fulvas, this one is fertile, so I'll work on the color, I promise!

(Fulva Korean x Transylvanian Tango) x Crimson Stories (F2)

The one above is a second-generation Fulva with a quite modern face... a favorite of 2022!

(Fulva Yunnan x Empire of Dusk and Dawn) x Crimson Stories (F2)

Several things to point out with this bloom.  First, it's a fused bloom, which also occurs about like in any group of daylilies.  It is not a polymerous bloom because it has two stigmas and 12 stamens!  A poly bloom will have four or five petals and double that number of stamens, but only one stigma. Fused blooms, like polys, only happen occasionally and most blooms on this plant are normal.  As far as breeding, it has wonderful, clean color and shows how selecting from your own seedlings can move your program ahead.  Empire of Dusk and Dawn gives clean color to its offspring.  It is bred from Richard Norris' Clarification and tends to 'clean up' a background and often gives a colored  edge, too, though not in this instance. Crimson Stories is tender here in Indiana (a beauty from Ludlow Lambertson in central Florida), but has been a great parent for giving us some beautiful faces. Note that this bloom does have a ruffled edge. 

Fulva Hankow x Firebird Suite

(Morning sunlight, not photoshopped!)

This bloom is of interest because it's a reverse bitone (sepals are the same color, but darker, than the petals).  We have seen several of these from orange crosses in the F1 generation.  Reverse bitones and bicolors are a special color group that is fairly scarce, especially among the tetraploids.  The late Bob Faulkner felt that with diploids you could not breed for a reverse bicolor, that they just 'showed up.'  We are working with them in our diploid lines, so we'll see what we conclude. As far as tets, there are very few reverse bicolors around to work with. Lamb's Breath from Dan Hansen is one: 

Hopefully we will be able to try our hand at bybridizing reverse bitone and reverse bicolored tets starting next season.  It should prove interesting!

(Fulva Korean x Ted's Tribute to Linda) x Colorful Chaos Nelli

 

Of all the Fulva crosses of the past 13 years, this one probably has the most toothy edge.  The color is saturated, though it could be more clean.  A pretty nice edge for a plant that is about 25% fulva!

Fulva Korean x Crimson Stories

Keeping with the theme of showing the possibilities of breeding with Fulva, here's a crested seedling that is an F1, from that influential parent, Crimson Stories. Not so beautiful, but an interesting result, nonetheless.  

(Fulva Yunnan x seedling) x Ashwood Summer Sunshine

Like most modern daylilies, you can get the unexpected.  I was hoping for a big watermark with this one, but it just decided to give up it's eye and become a self-colored cream.  At least it has a bit of an edge. Ashwood Summer Sunshine is good for giving larger blooms in its offspring, and that is true of this one at almost 6" from a smaller seed parent.

(Fulva Yunnan x Empire of Dawn and Dusk) x Hip Hop Music

 

Yes, this really is an F2 Fulva.  Not all F2's are this fancy, but this one has great color and a ruffled double edge.  Time will tell if it gets taller since it is still in the seedling bed.  It is only about 28" tall, so it needs more height and a purer red edge.  We have bred it to our future introduction yellow and red seedling to see what we get.

(Fulva Korean x Empire of Dawn and Dusk) x (Bodacious Bite x Gaudeamus)

This lovely pastel pink is an F2, so about 25% Fulva.  An amazingly clean pink from a yellow Fulva crossed with a pastel seedling. 

(Fulva Yunnan x Empire of Dawn and Dusk #1) x Crimson Stories

Another lovely pastel pink from an F2 Fulva.  I attribute the lovely color and double edge to Lambertson's Crimson Stories.  This one is tall and average sized so is a likely candidate for introduction in the near future.  Like all our tetraploid Fulvas, it is very fertile and has already been crossed with other fancy moderns. 

The above are a sampling of what we've come up with from hundreds of crosses with Fulva.  So far, the rhizomatous nature of the species has disappeared,  but we are still monitoring.  A few of the more vigorous ones show a tendency to spread out, but no more than any of the moderns that are great at growing new fans.  Hopefully the influx of Fulva genes will add hardiness and resilience to the strain. Time will tell if the Fulva hybrids can rival the moderns in appearance but give stronger growth, but that is the goal.