Read the previous paragraph again - you might have missed something. I have tentatively identified this seedling as a hexaploid, 66 chromosomes in the plant, rather than the usual 22 (diploid), 33 (triploid) or 44 (tetraploid). To my knowledge, it is the only hexaploid in existence; certainly, the only "natural" (no doubling of chromosomes by conversion) hexaploid.

I waited four years for this seedling to bloom, from selfings of ED MURRAY that I made in 1994 (and in subsequent years, too). Of course, anybody who has tried to set seed (with any kind of daylily pollen) on ED MURRAY has observed that the plant vigorously sets pods - appearing to be very fertile - but all (with rare exception) of the pods will fall off, classic triploid behavior.... one of the reasons I think ED MURRAY is a triploid. From all my efforts to self ED MURRAY I got one pod in 1994, containing one seed, and this seedling is the result.

For an update on the ploidy of ED MURRAY, please see Letter from Peter Mueller to Nick Chase, July 20, 1999 in "The Hexaploid Project".

The first bloom I got in the summer of 1998 was so ugly and misshapen, I wondered why I'd bothered to nurse the plant for so long. I was sad that I would have to trash it after such a long wait. But then the plant settled down, and the rest of the blooms looked like the picture. I assumed the plant was a tetraploid, because that was what I had been trying for - a fertile, usable-for-hybridizing, tet version of ED MURRAY - and because my use of ED MURRAY pollen on the triploid species H. fulva 'Europa' had produced Implausibility, a fertile tet. (In 1994, I did not think that ED MURRAY was a triploid; I only knew that its pollen would set tet seed on tetraploid plants.)

So, I put tet pollen from my very fertile seedling TR95-08 on each of the blooms, and Hexxed set pods very nicely. Yeah, it's a tet, I told myself. I tried putting Hexxed's pollen on various tets, but without success, as it is very difficult to use (see below).

As the fourth bud opened and bloomed, the first pod I had set began to shrivel up. Only then did I realize that I might have something other than a tet.

But what? It could conceivably be a triploid recreation of the mother, or even a diploid. But neither of these was likely because of the characteristics (described below) of the plant which indicate the chromosome count has been substantially increased. It could be a pentaploid, but the odds of creating either a triploid or pentaploid, on top of the odds thet such will survive "triploid block" or "pentaploid block", are miniscule. In the world of daylilies, nothing is impossible, I have discovered.... only that the odds are very low. That left hexaploidy, or the possibility that Hexxed is an aneuploid or polyploid or of ploidy greater than hexaploid, as the only probable alternatives.

For a hexaploid to have been created, there would have to be evidence that ED MURRAY produces triploid (33-chromosome) gametes. Is there any such evidence? The answer is, yes.

Nona Ford has created two tetraploid plants from diploid mothers using pollen of ED MURRAY ancestry. The first is THORNBIRD (1986), a cross of diploid LITTLE GRAPETTE X ED MURRAY, and the second is NORMAN LEE HENNEL (1991), a cross of diploid RECENT VINTAGE X THORNBIRD. (Both plants are registered as diploids, but are in fact tetraploid; I use them in my tet hybridizing with great success. For example, my tetraploid ROSOCOFSKY is a cross of NORMAN LEE HENNEL X BEST KEPT SECRET.) Now, the normal assumption would be that both LITTLE GRAPETTE and RECENT VINTAGE provide unreduced gametes of 22 chromosomes (with ED MURRAY providing the other 22). But they don't. I know, I've tried. I have put tet pollen on these diploids, and gotten nothing. Nada. Zero. Zip. By contrast, in the summer of 1998 I put THORNBIRD and NORMAN LEE HENNEL pollen (which carry the large pollen grains characteristic of ED MURRAY pollen) on lots of diploids, and got quite a few tet-size seeds, which were planted in early June 1999. By mid-July 1999, I had a few lovely green sprouts poking up, though the germination rate was low. (I found myself in the patch exclaiming YES! YES! each day as they grew.) Sadly, one was an albino; three were lost to insects or varmints of some kind. What's left, as of late-July 1999, are seedlings from JOLLY LAD (dip, Bill Spalding, 1980) X THORNBIRD; and my diploid lavender-pink seedling DP96-04 (a sibling of "Esther Gelade") X THORNBIRD. I also have a sprout from GARNET ROBE (triploid, Carl Milliken, 1948) X THORNBIRD, while most of my GARNET ROBE X tet crosses have yielded seed that doesn't germinate.

[An aside here: It struck me that the easiest way to create "natural" tetraploids would be to cross THORNBIRD and NORMAN LEE HENNEL with each other, which I did in 1998, and I planted the seeds in early June 1999. Four weeks later, all of my "regular" 1999 sprouts were coming along fine, but nothing from these two crosses. Six weeks later, and I am now seeing sprouts from the crosses both ways of these two plants.... about a 20% germination rate. Also sprouting after an interminable wait are my 1998 crosses of ED MURRAY X (either THORNBIRD or NORMAN LEE HENNEL, whichever pollen was available on a given day). Though "back-crossing" the child and grandchild onto the ED MURRAY parent yielded only four seeds, the germination rate is 100%, and this is a darn sight better than my previous years' efforts from putting tetraploid pollen on ED MURRAY, which yielded either no seeds at all, or seeds that didn't sprout.]

So, my conclusion is that, for THORNBIRD, LITTLE GRAPETTE provided the usual 11 chromosomes, and ED MURRAY 33. For NORMAN LEE HENNEL, RECENT VINTAGE provided 11 chromosomes, and THORNBIRD provided 33 (from one of the scattered large pollen grains in its otherwise-normal-looking tet pollen).

If ED MURRAY's pollen can provide triploid pollen gametes, are the odds good that the plant will also be able to sometimes produce triploid ova? I would think so.

Let's look at the physical evidence that Hexxed is a hexaploid. First, the pollen. Shown below, on the left, is normal tet pollen from WEATHERLY. On the right is Hexxed's pollen. These two pictures were shot through a small hand magnifier, so they're not very clear - I'll try for better pictures in 1999.

 

WEATHERLY's normal tet pollen is uniform, and the pollen grains are of customary tet size. ED MURRAY's pollen (sorry, not shown, the plant had finished blooming when I was taking pollen pictures) is sparse and crystalline-appearing, except for a few scattered, huge pollen grains. In Hexxed, as you can see, the pollen is made up almost entirely and exclusively of enormous grains, many more than appear in ED MURRAY's pollen. This pollen is extremely difficult to use. I got nothing (no pods, no seed) putting it on tets; once I realized that Hexxed is probably a hexaploid, I put it on several diploids, and got one pod (on HAMLET) but no viable seed. As I said, the pollen is extremely difficult to use; I suspect the pollen grains are simply too large to pass down the pistil of most blooms (especially diploids) and impregnate them.

Secondly, let's look at the evidence in plant habit that supports hexaploidy. Shown below, in the left-hand or upper picture, depending on how your browser displays them, is the perianth of a bloom on ED MURRAY that's about to open - the one in the center. (The perianth is the base of the bloom, the part that connects the petals and sepals to the scape.) Note the slenderness of the perianth - what one would typically expect of a diploid or triploid. The right-hand or lower picture, depending on how your browser displays them, shows the perianth of a Hexxed bloom that's about to open. Notice the width of the perianth - about two-thirds that of the opening bloom itself. A sure sign that the chromosomes have been doubled (or have come reasonably close to doubling).

Further support from the plant's appearance: The petals and sepals of the blooms are like shoe leather - really tough (but surprisingly, the blooms have no difficulty opening). Hexxed is totally and completely sunfast - the flowers look the same at 7 PM as they do at 9 AM. There is no slicking, even in the hottest sunny weather. The blooms also withstand rain well. The foliage is not notably larger than ED MURRAY's, but the backsides of the fronds have ribs which are larger and protrude more, another sign of doubled chromosomes.

For comparison, below is a picture of ED MURRAY. The color in this photo is not quite true - the flower is more black-red, like Hexxed - but even so, you can see how the colors have been intensified in Hexxed, especially the green throat. (The petals of Hexxed are also a bit wider, but I attribute this to my constantly watering the seedling bed, while ED MURRAY survives only on rainfall - and 1998, when these pictures were taken, was a dry summer.)

Of course, one obvious test remains, which I will try in the summer of 1999: Put diploid pollen on Hexxed, and see if I get tetraploid seed.