The true lilies — Lilium species

Turk's cap lily, Lilium superbum
Turk’s cap lily, Lilium superbum (West Virginia, July 16th, 2016)

Lilies sure are wonderful! I wish I had words to describe them better. Perhaps I could tell you of my feelings of joy and excitement upon finding some in bloom. Or maybe I could tell you of the sunny summertime days when the lilies flower in their full glory. But it would not be enough.

Canada lily, Lilium canadense
Canada lily, Lilium canadense (Pennsylvania, July 3rd, 2016)

True lilies within the genus Lilium are perennial, though they are relatively short-lived (perhaps no more than 10 years?). Most people are familiar with the garden variety Tiger Lilies (Lilium lancifolium), a native of far east Asia. It’s our native lilies that to me are really fascinating. They can be found all over the continent. In the eastern United States we find Canada lily (Lilium canadense), Michigan lily (Lilium michiganense), Carolina lily (Lilium michauxii), prairie lily (Lilium philadelphicum), pine lily (Lilium catesbaei), Roan lily (Lilium grayi), and Turk’s cap lily (Lilium superbum). In the western United States we find  the western lily (Lilium occidentale), panther lily (Lilium pardalinum), Columbia lily (Lilium columbianum), Washington lily (Lilium washingtonianum), redwood lily (Lilium rubescens), and the coast lily (Lilium maritimum). Doubtless there are more species that I cannot recount. (Additionally there are several related genera with similar structure and ethnobotanical uses such as Calochortus, Fritillaria, and Erythronium; many of these share the same family as Lilium, the Liliaceae family.)

Prairie lily, Lilium philadelphicum
Prairie lily, Lilium philadelphicum (West Virginia, July 19th, 2016)

The lilies may be found in a variety of habitats. The prairie lilies (Lilium philadelphicum) pictured above and below were found in dry upland and prairie meadow habitats such as the plateau of Dolly Sods Wilderness in Monongahela National Forest of West Virginia. They bloom there in July into August and at their peak they dot and paint the meadows with vibrant red color.

Prairie lilies at Dolly Sods
Prairie lilies at Dolly Sods (West Virginia, July 18th, 2016)

The Turk’s cap lily (Lilium superbum) grows primarily in woodland clearings with full or partial sun and the sunny edges along rivers or along roadsides. I have read that they actually require only about 2 hours of full-sun per day, so consider them fairly shade-tolerant. Older Turk’s cap lilies may display dozens of flowers per plant.

Turk's cap lily, Lilium superbum
Turk’s cap lily, Lilium superbum (Maryland, July 13th, 2016)
Turk's cap lilies, Lilium superbum
Turk’s cap lilies, Lilium superbum (West Virginia, July 13th, 2016)
Seedpods of Lilium superbum
Seedpods of Lilium superbum (September 14th, 2016)

The seed pods of Lilium species form after the flower has been pollinated and stops blooming, usually around August or so. The seed pod needs about six to eight weeks to finish ripening around September and the beginning of October. When it has ripened, it turns brown, and dries out (a process called dessication) and splits to reveal numerous teardrop-shaped seeds stacked in vertical columns.

Lilium philadelphicum in flower
Lilium philadelphicum in flower (July 18th, 2016)
Lilium philadelphicum seedpod early formation
Lilium philadelphicum seedpod early formation (August 12th, 2016)
Lilium philadelphicum seedpod ripening
Lilium philadelphicum seedpod ripening (September 13th, 2016)
Turk's cap lily seeds in the pods
Turk’s cap lily seeds in the pod (September 23rd, 2016)

The seeds have a papery coating on them which gives them lift that they may more readily blow about in the wind as a means of dispersal. They are quite easy to store in a cool, dry place out of the light for several months, but they lose viability if stored longer than a year. Germination has some pecularities — when it comes to lilies, folks sometimes talk about “hypogeal” germination. Hypogeal means below ground — in other words, the new plant first focuses on establishing roots before it dare raise a leaf above ground level (but when it does this is called epigeal growth).

Carolina lily seeds
Carolina lily seeds (Lisa Tompkins — North Carolina Native Plant Society)
Carolina lily seeds
Lilium superbum, Turk’s cap lily seeds

In the following two pictures are some of the stages of growth of young Lilium plants. In the picture below, we see the single leaves of new emerging plants. The lilies are monocots, therefore, their first-year leaf presents itself in the singular. In the years following, the plant beings its ascent upwards as a rising stem unfolds a whorl of leaves in distinct circles. By the fourth or fifth year the plant might begin to flower.

Lilium canadense 1st year seedling
Lilium canadense 1st year seedling (Delaware, May 1st, 2016)
Lilium canadense 3rd (?) year seedling
Lilium canadense 2nd or 3rd (?) year seedling (Delaware, May 1st, 2016)

Another way to propagate lilies is by corm division. What’s the difference between a corm, a tuber, and a bulb anyhow, you ask? Good question. A corm is defined as something like the underground swollen base to the stem. From it the roots grow, and the corm itself is covered in little scale-like things that are essentially bud or leaf scars. A bulb is a fleshy spherical organ at the base of the plant that is storage for winter dormancy. A bulb is made up of layers, just like an onion. A tuber, by contrast, is an underground storage organ that doesn’t really fit the description for corm or bulb. Its storage organ is generally more versatile, but also more uniform in structure. One learns to notice the difference between the three, but precise definitions are tricky and sometimes overlapping.


Canada lily corm displaying cormlets
Canada lily corm displaying cormlets


In the picture above, we can see how each lily corm is covered in dozens of little cormlets. This is an adaptation to disturbances such as digging. When the corm is disturbed, many of these little cormlets break off the mother, and a chemical signal is sent within each of them that will trigger sprouting. In such a way the plant can clone itself dozens of times! This ability practically encourages digging. Indeed, as M. Kat Anderson writes in Tending the Wild: Native American Knowledge and the Management of California’s Natural Resources —

“When California Indians dug up the parent bulbs or corms, the bulblets or cormlets — many of which were tenuously attached to the parent — were detached and remained in the soil to grow. Those that were not knocked off in the digging process were intentionally removed. The digging sticks churned and broke up the soil in the collecting patch, aerating it and dispersing the detached bulblets and cormlets. Thus the little offsets found themselves in ideal conditions for rapid growth into mature plants. This process, carried on for millennia, ensured each successive year’s harvest.”

She goes on to note the research of botanist Frederica Bowcutt, who states that,

“Redwood lilies [Lilium rubescens] occur mainly in forest openings created by limited logging and in berms of loose soil created by annual grading of a dirt road in and near the park. [..] It is conceivable that the redwood lily was at one time more abundant when large bulbs were periodically harvested and bulblets were left to repopulate the tilled soil. The grading may in part mimic the effects of agroecological methods used by indigenous peoples. Grading aerates the soil, as does the traditional digging of bulbs by native people for food.”

As a food source, the cooked lily corms are fantastic. The texture is crisp like a carrot and starchy like a potato. The flavor is faintly sweet but otherwise fairly nondescript. Lilium corms have an ethnographic history as a staple food, and based on my personal experience with this plant it comes as no surprise. I would love to see a nutritional breakdown, but for now I will have to settle for my intuition which tells me that it is healthy and filling.

The lily is a perfect candidate for reintroduction and rewilding for so many reasons. Beyond it’s beauty and ecological function, it serves as a first food and responds well to digging and disturbance.

It must be noted that digging alone ought to be considered not enough action to ensure the regenerative growth of lily patches. It is important that seed-scattering also occur for the sake of genetic diversity. Clones are sexually incompatible with each other on the individual level, and too many clones grown from only a few distinct genetic individuals will over time result in inbred populations. The indigenous way to kill two birds with one stone is to dig the lily corms after the seedpods have turned brown at the end of the summer or beginning of autumn and are already drying and ready to spill their seed. This also happens to be the period of time when the corm is the largest as a result of being allowed to grow all season, as we saw also in the case of ramps or wild leeks (Allium tricoccum). Therefore, while digging each corm, leave behind in the ground the cormlets and make it a point also to intentionally scatter the seed from out of the pods. This is regenerative harvesting; this is symbiosis!

Toadshades and Wakerobins — Trillium species

Painted trillium, Trillium undulatum
Painted trillium, Trillium undulatum (Vermont, June 1st, 2016)

Trillium flowers offer a spectacular spring display. Commonly known as toadshade and wakerobin, they rank among the most beautiful, and the most illusive, of spring ephemeral wildflowers. They come in many different sizes, arrangements, and colors, from the smallest Snow Trilliums (Trillium nivale) to the Sweet Wakerobins (Trillium vaseyi) with flowers as wide as your hand is long! The distribution of Trillium species is all across the North Hemisphere in temperate regions. They are found in forests with rich, loamy soil remaining moist throughout the year.

One of the local native species to the Mid-Atlantic region that I call home has large red flesh-colored flowers. These red wakerobins — Trillium erectum — are colored as well as scented so as to attract the attention of carrion flies for pollination. In the eastern temperate forests of North America, it is quite common to find red, flesh-colored flowers. Everything from trillium,  to pawpaw, sweetshrub, wild ginger, and even skunk cabbage. And every one of these plants are pollinated by carrion flies! Why carrion flies? This is a fascinating ecological aside. As John Muir said,

“When we try to pick out anything by itself, we find it hitched to everything else in the Universe.”

There is more to the flower and fly connection than meets the eye…

When we think of pollination, we think of the honey bee. But the honey bee is not native to North America. Instead, our native bees are ground bees. The ground bees don’t emerge until soil temperatures begin to warm mid-spring, in late April or early May around here in the Mid-Atlantic. (This time also coincides with the opening of nectar-rich nutrient-dense tulip poplar blossoms!). By the time mid-spring comes around, most of our early wildflowers are already in bloom.

Rather than relying on the slumbering bees for pollination, garnering the attention of carrion flies instead seems the better evolutionary strategy. The carrion flies seek out the colors and smells of rotting meat, and they emerge much earlier in the spring — even in the late winter — thanks to the thermogenesis abilities of a plant called skunk cabbage (Symplocarpus foetidus). Skunk cabbage can generate temperatures 15–35 °C (27–63 °F) warmer than the air surrounding the plant! Known as the mother plant by some, its deep red mottled flowers with a foul, rotten smell like flesh prove quite attractive to the carrion flies, who awaken with the warm microclimates created by the skunk cabbages piercing through snow and ice like spears. The flies awaken before the eve of the upcoming spring flower show that will unfold over the coming weeks. These interrelationships demonstrate to us vividly how the ecosystem is more than the sum of its parts.

Trillium erectum growing along the Susquehanna
Trillium erectum growing along Susquehanna (April 15th, 2016)

Trillium species are an occasional food and medicine to humans. The leaves may be eaten as cooked greens, but the root is bitter and inedible, and ought to be considered poisonous. Trillium are within the family Melanthiaceae, which contains some of the most toxic plants such as the death camas (Toxicoscordion/Stenanthium/Zigadenus sp.), fly poisons (Amianthum sp.), and hellebores (Veratrum sp.).

According to folk medicine, a decoction of Trillium root has the medicinal value of controlling blood flow, either to stop internal bleeding or to help in its passage. It was used as an herb to stop bleeding after birth, thus becoming known as “birthwort” or “bethroot.”

Trillium hybrid along Susquehanna
Trillium hybrid along Susquehanna (April 30th, 2016)

The seeds of Trillium develop inside fruit-like capsules in mid-summer. These fruits remain upon the plant for several weeks before they ripen, turn soft, and eventually fall to the ground and split open. Each seed has a large elaiosome attached. An elaiosome is made up of fatty lipids and amino acids, making the seed attractive as a food source to ants who carry them underground into their colonies. Once underground, the ants consume the elaiosome and cast the naked seed into a rubbish pile. Here the seed awaits germination in an optimal environment in terms of light, temperature, and moisture. This process is known as myrmechocory.

Myrmechocory, ants of Trillium seed
“Myrmechocory” — Ants on Trillium seed (not my image)

See the beauty of ecological relationships… Symbiosis is found everywhere. The skunk cabbage fosters carrion flies to pollinate Trillium flowers which form a seed that gets dispersed by ants. What a wonderful, interconnected world!

Trillium erectum gone to seed
Trillium erectum gone to seed (West Virginia, July 14th, 2016)

As the fruit of Trillium begins to soften after a few weeks upon the plant — generally in July and August — you can begin to harvest it. Pick whole fruits and keep them somewhere moist where they can ripen further if necessary, becoming even softer until they split open.

Trillium fruits
Trillium fruits in July (Pennsylvania, July 6th, 2016)
Trillium seed spilled upon the ground
Trillium seed spilled upon the ground (Pennsylvania, July 3rd, 2016)
Seeds inside the fruit of Trillium erectum
Seeds inside the fruit of Trillium erectum (July 3rd, 2016)

Trillium seeds, like ramps (Allium tricoccum), are double-dormant, meaning they need to wait not one, but two winters for germination. However, by harvesting the seeds a little bit early when they are slightly premature, the seeds react by germinating faster, after only a single winter. Some horticulturalists will do this in order to speed-up production times. I don’t have any experience doing this. For my purposes, reintroducing this plant species into new areas, I have allied myself with time and do not have such a sense of urgency. For the sake of genetics and plant health, I prefer my seeds to be as mature as possible and that they take their time.

To store Trillium seed, keep them in a cool, dark, and moist environment. If they dry out, they change to a darker color, shrivel up, and completely lose viability. To prevent this, I place them in a bucket filled with damp forest soil and I leave them there until I am ready to plant them into the ground.

Growing Trillium from seed is another one of those long-term plant investments. After sowing (year 1), there’s the double dormancy, and then the first monocotyledon leaf grows out of the ground and stretches out (year 2). The Trillium may continue with just one leaf another year. But by year 4, all three leaves ought to be showing. It may not be until another 3 years has passed before the Trillium flowers and can set seed of its own. They take overall approximately 7 years from seed to flower. Colonies will form, but can take a very long time to establish. However, with consistent sowing for six years straight, by the seventh year a colony ought to be self-sustaining.

Wild Leeks aka Ramps — Allium tricoccum

Ramp patch
Ramp patch (Pennsylvania, late April 2014)

Everyone who has heard of ramps, loves ramps! Also known as wild leeks, they are one of the most exciting and abundant foods to greet us in the springtime. The flavor is like an exquisite onion — strong, pungent, and healthy as garlic but surprisingly sweet. Every bit of the plant is a premier wild edible, from the tip of the leaves to the base of the bulb.

Fun fact, the name for the city of Chicago comes down to us from the Algonquian word chicagoua, which means the wild leek, Allium tricoccum. “Chicago” — the stinking place of onions!

It is important to understand the anatomy and life-cycle of Allium tricoccum in order to harvest appropriately. As with many onions, these plants have three basic parts:

1) The roots. These grow from out of a nodule underneath the bulb. This is area contains the genetic information of the plant and guides its growth each year, expanding vertically like sedimentary layers of rock. It is possible to guess the age of a plant by counting the layers in the nodule, much like we would with tree rings. The root nodule, if separated from the bulb that grows above it, will grow a new bulb the following year if kept alive or in the ground.

2) The bulb is the storage-organ of the Allium tricoccum plant; it grows seasonally, waxing and waning in size like the moon. Smallest in winter, largest at the end of spring. If undisturbed, the bulb will increase in overall size through the years until it is mature enough to split and divide, thereby cloning itself into two or more plants.

3) The leaves. They emerge from out of the bulb. Younger plants have one leaf; more mature plants have two leaves. Sunlight is absorbed by the green chlorophyll in the leaves, which feeds and powers the rest of the plant’s structures underground. The leaves may be harvested in a sustainable way. However, keep in mind that while removing the leaves will not kill the plant, it will diminish the plant’s ability to gather and store energy.

Root nodule of Allium tricoccum
Root nodule and bulb of Allium tricoccum
Early spring ramp growth
Ramp leaves growing from thin, hungry bulbs in early spring (late March 2016)

The most sustainable way to harvest ramps is to wait until they have finished growing for the season. This will be the period of time leading up to just before the foliage begins turning yellow on the tips, marking the beginning of the dormancy stage which will last until the next spring. (The ramp leaves will continue turning yellow and brown down the stem, wilting away until all of their above ground portions have disappeared.)

At this point in the end of their growing season, the bulbs have attained their largest size for the year, and the roots are strongly bound within the earth. Because there is a weak point between the foliage and the roots at the nodule just underneath the bulb, by gently pulling and tugging at the plant by its leaves, a crack will be heard. This is the sound of the root nodule separating from the bulb! As the forager continues pulling the plant out of the ground, they are delighted to find that the roots and nodule have remained in the ground to grow for another year, but the bulb and leaves are harvested and in hand, ready for eating. This is a win-win situation for both human and plant, and the alternative to destructive digging. Harvesting in such a way can be considered regenerative, or symbiotic.

It is also worth mentioning that the ramps are easier to clean when pulled in such a way; with the root nodule left in the ground, the outer skin of the bulb slides off easily, and with it, most of the dirt or mud. See the picture below…

Ramp bulbs
Ramp bulbs with the root nodule snapped off (Vermont, June 2016)

In 2016, I gathered some snapped-off root nodules into a pot with good soil, as a test to see how (or if!) they would regenerate. In spring of 2017, I uncovered the roots and was pleasantly surprised to find that around 80% of them had definitively regenerated, and were rooting and growing new bulbs! This is scientific proof that ramps can indeed be harvested regeneratively via the method explained above, carefully leaving the nodules in the ground but harvesting the bulb and leaves.

Propagation of Allium tricoccum from root nodules
Propagation of Allium tricoccum from root nodules

When you are gathering ramps for food in late spring, do not collect any that look like the picture below: it is the beginning of a flower stalk. If left undisturbed, the raceme will continue to grow up 4-10 inches high above the litter of leaves, and at its tip will bloom forth a white, spherical umbel of flowers.  These flowers, after pollination, will each go on to develop seeds. These seeds hold within them the future life of the population.

Raceme of Allium tricoccum
Raceme of Allium tricoccum (Pennsylvania, May/June 2016)
Ramp seed head developing
Ramp seed head before the flowers unfold (Pennsylvania, late June 2016)

Pictured above is an umbel of Allium tricoccum before the flowers have opened on a late June day in Pennsylvania.

Ramp flower
Ramp flower (July/August 2016)

Above is a picture of a ramp flower in bloom. Note how within each flower in the umbel is the ovary, and it is made up of three ovules. This means every individual flower has the potential to develop up to three seeds.

Ramp seed heads
Ramp seed heads (mid-September 2016)
Ramp seed head
Ramp seed head (mid-September 2016)

The seeds of Allium tricoccum ripen from mid-September through mid-October.

Ramp seeds
Ramp seeds

If the seeds are kept dry and relatively cool in a space out of the light, they will store well for a few years.

Ramps, much like Trilliums, are double-dormant. This means they will not germinate after a single winter cold period, but instead require two winters. If sown in the fall, expect to see the first grass-like seedlings not the next spring, but the following spring after. From germination to flower takes approximately 5 years. Therefore, a ramp plant from seed to flower is a 7 year investment. Consider this before digging up any portion of a patch!

My philosophy is to scatter ramp seed in the same spot for six years. By the time the seventh year arrives, the first seeds sown will flower and soon set their own seeds, and there will already be six generations more lined up on deck. At this point the patch can be considered effectively self-sustaining.

Ramps grown from scattered seed
4-6 year ramps from 2010-2012 seed (Green Light Plants)

Ramps are an excellent candidate for ecological restoration. They are quite hardy, and will compete well against “invasive” plants such as multiflora rose, japanese honeysuckle vine, and others.

Go create more patches!

Bluebells — Mertensia virginica

Virginia bluebells are among our most beautiful spring ephemeral woodland wildflowers. I love the lush green-and-blue carpets that blanket the forest floor in mid-April in southeastern Pennsylvania.

These bluebells, Mertensia virginica in Latin, are members of the borage or forget-me-not family (Boraginaceae). Their electric-blue coloring as well as the elongated, pendulus, trumpet shape of their flowers look alike to kindred cousins such as borage, viper’s bugloss, comfrey, waterleaf, and others.

Woodland carpet of Virginica bluebells
Woodland carpet of Virginica bluebells (April 15th, 2016)
Bumblebee on Virginia bluebell flowers
Virginia bluebells (April 15th, 2016)

Mertensia virginica flowers are excellent insectaries. When a colony first blooms, it may be one of the year’s first bonanzas for pollinating insects, such this bumblebee pictured above drinking the sweet, sweet nectar of the blue flowers.

While the root of Mertensia virginica is not appetizing to humans — I have tasted it and can attest that it has a strong bitterness to it — the leaves, stems, and blossoms are quite palatable. The blossoms are deliciously sweet raw, much like sweet violets, and make for a delicate and beautiful garnish on a salad. The foliage has a stronger flavor, and is most often cooked steamed. Some report that the flavor is like that of oysters, though this trait is more often associated with Mertensia maritima, a sea-side coastal species within the same genus.

Virginia bluebell blossoms
Virginia bluebell blossoms

After the blossoms erupt in early and mid- April and get pollinated, the petals begin to drop off the calyxes one-by-one, leaving behind the prominent central pistil and the ovary to nurture the young seeds.

Mertensia virginica beginning to form seed
Mertensia virginica beginning to form seed (April 30th, 2016)
Mertensia virginica seed capsules
Mertensia virginica seed capsules (April 30th, 2016)
Unripe seeds of Mertensia virginica
Unripe seeds of Mertensia virginica (April 30th, 2016)

In this picture above we see the unripened seeds of Mertensia virginica. This picture was taken at the end of April, and it was perhaps another week before I would begin to avidly collect the stems for seed.

Mertensia virginica calyx
Mertensia virginica calyx showing 4 seeds

Here’s a closeup of an ovary showing the four seeds which are typical of each capsule.

Ripening seed of Mertensia virginica
Ripening seed of Mertensia virginica (May 15th, 2016)

By mid-May, some of the seeds have ripened in their capsules, and they almost immediately fall to the ground. If one wishes to gather good amounts of the seed, some strategy is required. The technique that was shown me is to gather whole stems of plants just before the seeds begin to drop off, but not much earlier as the seeds may not mature properly. This requires some familiarity with the plan’s lifecycle and helps develop one’s powers of observation.

I lay out the stems I’ve collected, and the juices and nutrients remaining in the foliage finish the job of ripening off the seeds, which drop off one-by-one onto a surface for catchment, such as a table, sheets of cardboard, cloth, or whatever other setups one might imagine.

Letting seeds ripen and drop
Letting seeds ripen and drop (May 26th, 2016)
Mertensia virginica seeds
Mertensia virginica seeds

When the seeds are fully ripe, they turn a rich, dark-brown color. The green ones pictured will also turn brown with time. Generally, the darker the seed, the more time it has had to mature and the better its viability.

Mertensia virginica seed
Mertensia virginica seed closeup
Mertensia virginica seedling
Mertensia virginica seedling

I have heard some conflicting information about the viability of bluebell seed after drying. Some say drying destroys a lot of their viability. Others say they are quite tolerant of drying. In any case, I air on the side of caution and scatter collected seed as soon as I can, mimicking their life-cycle in the wild. I collected most of my Mertensia seed around the end of May; by the end of June I had none of it remaining. All of it was scattered in the wild in optimal habitats where spring ephemeral growth was limited or non-existent.

Finding suitable locations for more bluebells is a fun adventure and wonderful excuse for aimless woodland wanderings (a favorite past-time of mine). Fortunately, the bluebells are quite resilient and will grow in a number of places, including under multiflora rose! It is thus an excellent candidate for ecological restoration.

The seedlings pictured above are one- to two-years-old. It will be another 3 or 4 years before these little guys are popping flowers and setting seed of their own. What a joy to watch them grow!