Over a decade ago I decided to try planting the native Frogfruit (Phyla nodiflora) in the maritime grassland exhibit at the South Carolina Botanical Garden. To my amazement, this species that I knew of from the fringes of saltmarsh in the Lowcountry thrived in both wet and dry soils of the upper Piedmont of South Carolina! The plant has proven to have incredible versatility and grows well in sand or clay and can be flooded for weeks and completely dry as well. Unlike many other plants that can accommodate such diverse conditions it isn’t so ugly that only a mother could love it, in fact, it’s charming.
Frogfruit is a low (4” tall) trailing groundcover with 1.25” long leaves that forms a solid mass of foliage but lacks deep root structures and thus does not compete with deeper rooted structural element plants. The flowers are pale pink to lavender and resemble tiny lantanas (a close relative). The flowering season here in the south begins in April and can continue through hard freezes (typically November) but may produce flowers year-round in mild winters.
Phyla nodiflora ‘Ramble On’
Our plants, Phyla nodiflora ‘Ramble On,’ are from a Charleston County, South Carolina collection along the margins of a wet ditch (freshwater), but the species has an amazingly wide range being found from New Jersey west to California and throughout the tropical regions of the world. Another species, Phyla lanceolata, is a more upright plant, with a similar range (but extending north to Ontario) it has longer leaves and is generally less showy as a groundcover.
Phyla nodiflora ‘Ramble On’
This is the ideal living mulch for tough areas of your landscape. It spreads rapidly but is easy to keep contained by trimming the edges of your patch. We placed it in one of our pond overflow pits and were amazed to see it completely transform a time-sink of constant weeding into a mass of lovely little flowers while allowing the Hymenocallis and Hibiscus to continue to rise through the groundcover without obstruction.
The flowers are favored by skipper butterflies, particularly the smaller species and there is an all-day-long collection of hundreds on our patch every day. In addition, small flies, native bees, sweat bees and tiny wasps are fond of their constantly produced flowers. The leaf and stem color ranges from green to deep purple depending on the environmental conditions—generally, the more exposed to sun, intermittent drought or salty soils, the more purple in the plant. If the goal of your garden is to increase the production of life by filling all your spaces with plants that are loved by insects while at the same time reducing the need for mulch and weeding, this plant is definitely worth a try. Look for this in the near future. – Patrick McMillan.
The past couple of weeks the small, freakish flowers of one of the strangest of plants have begun to open in our gardens – pipevines. It’s difficult to believe that nature could summon up anything as strange as the flowers of pipevines. If you remember the “regular” aka actinomorphic and the “irregular” zygomorphic flowers from high school biology then you would have to consider the strange, inflated flowers of pipevines highly irregular.
These plants are related to wild gingers/heartleafs (Asarum) and are members of the basal angiosperms, which are plants with lineages that predate monocots and true dicots. Many of the early angiosperms have flowers that are pollinated by flies or beetles that they lure to the flowers with the pleasant smell of rotting meat or dung. Pipevine is no exception. They trick flies into entering the strangely shaped flowers which often have a funnel-like flare with a narrow opening and trap the fly in a chamber with the pollen and nectar to accomplish pollination.
The general shape of the pipevine flower also approximates the anatomy of a uterus and birth canal. As was all the craze in the early years of medicine, the resemblance to the uterus must mean it is good for treating issues with difficult pregnancies and childbirth. This archaic mode of determining herbal remedies is referred to as the doctrine of signatures – basically if a plant part looked like a part of the human anatomy it would be used to treat ailments associated with that organ. This avenue of medical care led us to the highly unsuccessful use of liverleaf (Hepatica) to treat ailments thought to be caused by the liver, like cowardice and freckles. Medical science has since progressed, maybe?
The pipevines include herbaceous upright perennials, short rambling vines, and enormous lianas (woody vines). The toxic chemical aristolochic acid is contained in all parts of the plant and though it is known to cause kidney failure (which probably didn’t decrease the problems with difficult childbirth) this chemical has also been shown to have antitumor properties. Don’t go munching on the leaves! Pipevine Swallowtails depend exclusively on pipevines for their larval food source and all the species we grow are host to large numbers of caterpillars in the summer.
Pipevine caterpillars
The Pipevine Swallowtail accumulates aristolochic acid which deters predators like birds from wanting to consume them (they don’t want kidney failure either). This may explain why Spicebush Swallowtails and Red-spotted Purples look so similar to the Pipevine Swallowtail, they’re taking advantage of blending into the poisonous crowd.
These beautiful butterflies always seem to find your pipevines, no matter how secluded you think they are. Don’t worry, we can’t have enough swallowtails and the vines regrow without issue. These are great plants to include in your garden to support our butterflies. The caterpillars have another interesting behavior, when disturbed they have bright orange “horns” that are technically referred to as osmeteria that it pushes out of the front of its body and emit a very foul odor!
Three native pipevines are found in the Carolinas. Common Pipevine (Isotrema macrophyllum) can become a huge liana, reaching the canopies of old-growth trees and spreads through runners and can be difficult to keep contained. It grows best in cool climates and tends to suffer or not grow in warmer parts of the Southeastern US.
Isotrema macrophylla
Woolly Pipevine (Isotrema tomentosa) grows very well in hot, humid climates and is known from only a few native populations in the Carolina coastal plain. Like its mountain cousin it is very difficult to restrain and may quickly outgrow the space you envision for it.
Isotrema tomentosa
The only other pipevine is very dissimilar to the others, Virginia Snakeroot (Endodeca serpentaria), which is an upright herbaceous perennial that rarely gets a foot tall. If you’re wondering why these former Aristolochia are now in the genera of Isotrema and Endodeca, well, it appears to all be about genetics. They are closely related but currently in different genera.
One of the easiest to grow and showiest, best-behaved species in our garden hails from Brazil, Argentina and Bolivia, Fringed Pipevine or Fringed Dutchman’s-pipe (Aristolochia fimbriata). The plants form scrambling vines that act more like a groundcover than a vine and have beautifully silver-veined small leaves. The flowers are intricately patterned with lines and fringed with long “tentacles” to create one of the most bizarre flowers you’ll ever include in the landscape. Though you wouldn’t pick a plant from Brazil to be rock-hardy here in Raleigh, this one has been fully hardy (zones 7a-9a). For those of you looking to add some swallowtails but unwilling to devote the space to a monstrous mass of vines this one is for you. Fringed Pipevine is also incredibly drought-tolerant and perfect for part-sun to dappled dry shade.
Great new plants for the garden do sometimes just happen. They can occur as a spontaneous sport from an existing planting, as a seed selection that has much better garden traits, but many of our most useful and ecologically important plants in the garden have their start in exploration. I was thinking about this today as I observed the tightly-clumping overwintering rosettes of one of our newest introductions to the JLBG—Oenothera riparia, Riverbank Sundrop.
Oenothera riparia
Sundrops hold a special place in my heart. My grandmother loved them and had large swathes of the old pass along yellow standard Oenothera fruticosa/tetragona in her extensive garden. Every June they would burst into flower for a brief week or so, bringing a brilliant light and foil to her many glorious iris. As I mature, I have come to value plants with connections to our being but also the value of similar plants that can provide the same nostalgia while giving us much more in the landscape.
A plant known as Riverbank sundrops may be a perfect example of a great garden plant that is native to the Southeast and provides more of what gardeners love, while also providing the native insects and other wildlife an abundant resource.
Oenothera riparia was described by Thomas Nuttall in 1818 from plants collected along the Cape Fear River near Wilmington, NC. Nuttall provided a very full description of the plant (for the time) though he presumed it to be a biennial—it is in fact a long-lived perennial. J.K. Small recognized the species in a segregate genus as Knieffia riparia in his 1903 Flora of the Southeastern US. In 1937, Munz reduced the species to varietal status as Oenothera tetragona ssp. glauca var. riparia, and after this it was essentially lost to the minds of taxonomists and plant enthusiasts being considered merely part of the immense variability of Oenothera fruticosa by Radford, Ahles and Bell (1968).
In the mid 1990’s while working in the tidal freshwater swamps and marshes of South Carolina, Richard Porcher and I encountered this species growing at the bases of Bald Cypress trees, on stumps, and on floating logs just above the high-water tidal line on the Waccamaw, Pee Dee, and Edisto Rivers. My frustration with its identification led me to the name Oenothera riparia and its recognition as a completely and consistently distinct taxon—which is very unusual in a genus known for its morphological variability and messy taxonomy.
Oenothera riparia
This plant is smooth (lacking hairs) throughout with thick textured dark-green leaves and a very bushy habit—it does not produce long stolons or rhizomes, so it forms a dense clump. The stems generally range from a foot to nearly 3 feet in height and the thick stems become semi-woody, providing a stiffly upright growth form. Rather than a single burst of flowers this species produces masses of flowers over the entire summer (June-late August) with sporadic flowering later in the season.
Imagine, here we could have a sundrop that won’t spread like wildfire, doesn’t flop, and flowers for month after month! Sounds like everything you would want in a plant. There is only one problem, it hasn’t been cultivated. This is where the adventure continues and as we embark on this adventure, I would like to take you along for the ride. There is so much that goes into identifying a potentially great garden plant, evaluating it, and bringing it to the trade.
Our initial collections were made this August (2022) when Zac Hill, my wife Waynna, and I were traveling through the SC coastal plain. We made the stop at the Edisto River in Colleton County where I had seen the plant many years before and just like I remembered, there they were, full of seed and with some flowers still present. The only problem is that they were growing far out in the water along the bases of trees! The Edisto is a blackwater river with a large tidal amplitude at this location and it was full on high tide. If you’re a field botanist you forget entirely about the things that concern other more rational folks, like the multitude of large Alligators, and make for the plants. I found some low branches on a neighboring willow tree that kept my feet in only a couple feet of water, and balancing on the branch, in the water, made my way all the way to the edge of the river where a fabulous floating log provided an abundance of seeds and two small plants from a large cluster of individuals. The only casualty was my prescription glasses which promptly fell from my head into the depths of a rapidly moving Edisto River (my wife will not easily let me forget the cost of this single collection). The first step in the process of bringing a new plant into production is done.
Oenothera riparia
Our two small divisions were placed directly into the sun garden at JLBG and the seed was sown. For this plant to be successful in a garden it must be able to not just survive but thrive in a common garden condition far away from its very narrow niche at the very upper edge of the water along the Edisto. You might think this would be unlikely, but other incredible garden plants are entirely found in wetland communities and thrive under very different conditions in the garden. A good example of this is found in another of our introductions, Eryngium ravenelii ‘Charleston Blues’, which comes from high pH wetlands not too far away from the Edisto River.
Will it be hardy here? Will it survive under normal garden conditions? Will it maintain its distinctive and garden-worthy features? Well at least part of this can be answered already. Our tiny divisions are now large overwintering basal rosettes. The plants have not thought about running away from their tight cluster, and they grew very well during their first autumn in soil that was not kept overly moist.
The real test lies ahead. What will the plants nature be under cultivation? Will it be as good as I think it could be? Will it be better? One can only hope, put in the labor, and follow along to find out the end of this story for Oenothera riparia. It could provide all of us with a stately and handsome bit of nostalgia with far greater design and utilitarian use for humans and our native biota. What’s more, there are other seemingly great garden-worthy Oenothera out there—not in far-flung locations but right here in the Carolinas. Have you ever heard of Oenothera tetragona var. fraseri? If not, look for us to tell you more about that in the years to come, there’s a fantastic form with huge flowers in the Blue Ridge escarpment of South Carolina!
The cold and raw weather of late autumn and winter provide the perfect opportunity to sit down with the dissecting scope and put our ferns through the identification mill. Often gardens and nurseries receive a plant into their collections from an exporter or collector who has put their best guess on the identification. After many years in cultivation, we realize what we thought was the right species name for our specimens is incorrect. Today’s nasty weather provided the opportunity to examine, in detail, one of our favorite evergreen fern groups – Dryopterissection Variae.
These firm-leaved evergreens produce thick-textured, durable, medium-sized fronds of varying shape but all display a noticeably longer basioscopic pinnule (that’s fancy talk for the lowest, innermost segments of the divided leaves). All members of the section that we have grown have proven to be very adaptable to our hot, humid summers and unpredictable winters if grown in shade or partial shade in moist woodland garden conditions. The fronds tend to burn if they receive too much light. They are late risers in the spring often not producing a new flush of leaves until late spring or even early summer.
At the beginning of the day, we started with 8 accessions of Dryopteris varia, 2 accessions of Dryopteris bissetiana, 6 accessions of Dryopteris formosana, one accession of Dryopteris saxifraga and a couple of unknowns. From these numbers you would expect that the one plant we would know best would be Dryopteris varia.
Dryopteris bissetiana comparison
Well…it turns out all the plants we had received or had identified as D. varia were actually representative of other taxa. If you’ve never tried keying ferns using The Flora of China or The Ferns and Fern Allies of Taiwan, you would have no idea just how difficult a process this is. The floras of these areas are notoriously difficult to use and often contradictory or difficult to assess using illustrations or pictures (yes even plant taxonomists google names to find images). Very quickly we became intimately familiar with the nature of the stipe and rachis scales, frond outlines, and disposition of the vestiture (yeah you think that sounds easy, right?).
We found most of our collection was actually composed of Dryopteris bissetiana, which are mostly from collectors who sent us tentatively identified wild-collected material. The majority of these were from Sichuan in China, however one very beautiful, deep green and glossy selection that is only half the size or less of the others was Tony’s collection in Korea, and has tentatively been identified as Dryopteris saxifraga. All of these are remarkable garden plants, but we are very excited to some day offer the choice dwarf from Korea which we have named ‘Cheju Dwarf.’
It was a pleasant surprise to find that our collections of Dryopteris formosana were correctly labelled, but we weren’t prepared for there to be two distinctly different looking plants represented in our garden that are the same species.
One of these is the plant that has been shared among fern enthusiasts for some time that is the typical sexually reproducing diploid. The other is an apogamous triploid that looks like a completely different species. In a diploid (like you and I) the pairs of chromosomes uncouple and one copy of each goes into making the male and female gametes.
Dryopteris formosana forms
Thus, each gamete has only one set of each chromosome (haploid) and when combined with those from the complementing sperm or egg results in another diploid. Plants sometimes have a mistake in their cells that lead to the production of gametes with twice as many chromosomes as they would normally have and when such tetraploid plants breed with a diploid the result is a gamete with 2 copies of each chromosome combining with a gamete with only one—thus triploid. In your average plant this triploid is a dead end for reproduction by seed or spore because they have an uneven base number 3—which can’t be divided into an equal number of chromosomes, so it is sterile.
Dryopteris formosana comparison diploid left triploid right
This triploid avoids the curse of having an uneven number of chromosome pairs by avoiding sexual reproduction and producing spores that will result in new plants without the traditional interplay of sperm and egg on a germinated gametophyte (yes apogamy in ferns is still legal in all states and countries). We were puzzled when two very different looking ferns keyed to the same species. Everything that was in the key matched. The bullate hairs, the shape, the color, the basioscopic pinnae and the overall shape.
Our taxonomist, Zac Hill, very quickly uncovered a recent paper by Kiyotaka Hori, et al (2017) which explained and beautifully illustrated our conundrum. The triploid produces a wider, far more pentagonal frond with a less erect nature in the way the blade is held, and a deeper green, highly pleasing color—now that’s pretty darn cool! This new discovery we have named ‘Yushan 2 X 4.’ A new plant for us all to grow in the years to come and now you know why we chose the name diploid (2) X tetraploid (4).
Dryopteris formosana triploid
Now that we realize we grow seven different forms of Dryopteris bisettiana, each collection will be given a cultivar name, which will refer back to their specific origin and uniqueness.
Every day brings discovery when you manage a collection of 30,000 taxa but one thing we know for sure, these are amazing, well-behaved, slow growing woodland plants that are the essence of what makes Juniper Level Botanic Garden so amazing.
Patrick McMillan, director of horticulture and gardens
Literature cited:
Hori, Kiyotaka, L. Kuo, W. Chiou, A. Ebihara and N. Murakami. 2017. Geographical distribution of sexual and apogamous types of Dryopteris formosana and Dryopteris varia (Dryopteridaceae) in Taiwan. Acta Phytotax. Geobot. 68 (1): 23-32.
Elliott’s aster (Symphyotrichum elliottii) is the absolute last of our asters to flower at JLBG. It doesn’t begin to flower until the first of November and withstands the mild frosts of October like they didn’t even happen. It is naturally found in tidal freshwater marshes and other moist open sites from the Virginia and Carolina coastal plain south to Florida and west to Louisiana. Though it hails from moist environments it thrives under general garden conditions if the soil isn’t allowed to become too droughty.
Symphyotrichum elliottii
The plant has a lot to recommend it besides the time of flower. It forms stiff stems rising 5-6’ tall crowned with a dense pyramidal arrangement of inflorescences of pale pink with a hint bluish-purple ray flowers and bright yellow disk flowers. The lack of lanky branches allows this tall aster to display its flowers without flopping all over the rest of your garden in the manner typical of asters. It spreads via rhizomes, so you need to be sure to give it space to roam a bit. It provides a dramatic impact when planted at the back of borders. Though it spreads, it doesn’t roam far from the parent plant and can be easily kept in place by yearly thinning of the outer edges of the clumps.
Symphyotrichum elliottii
The most outstanding feature of this beautiful aster to me is the number and diversity of pollinators it supports. I don’t think I’ve ever seen a plant that attracted more. In addition to swarms of honeybees, bumblebees, carpenter, and solitary bees the flowers draw in numerous pollinating flies, halictids, moths and skippers. I love plants that extend the color season and though we all think about early spring, we really should also plant to extend our love affair with color into the leafless season and Elliott’s aster does this is a big way.
History is replete with examples of new plant species that are first encountered by intrepid plant explorers, yet described later by taxonomists. Salvia darcyi was discovered and introduced into cultivation by Carl Schoenfeld and John Fairey of Yucca Do Nursery. Three years later, they guided researchers to the site who subsequently described the species without acknowledging the original collector. It is unfortunate that the act of discovery by those in horticultural circles are so seldomly recognized (not to mention the indigenous peoples who have known many of them for eons).
Upon my first visit to Heronswood in the autumn of 2019 I was shown a splendid robust Begoniawith heavily lobed leaves, short upright stems, and amazing tight-clumping habit with yellow (yes yellow!) flowers. I immediately confirmed that this was a heretofore undescribed species. The plants had been grown from the seed collection made by Dan Hinkley along with fellow collectors and nurseryman, Shayne Chandler and Leonard Foltz, from Arunachal Pradesh India. These plants were shared by Mr. Hinkley with Monrovia who immediately released it under the name TectonicTM Eruption Begonia (Begonia sp. DJH18072).
The unknown Begonia has now been given a formal scientific name Begonia lorentzonii by Swedish taxonomist Eric Wahsteen and the Indian researcher Dipankar Borah, based on two specimens collected by Borah in November of 2018 (incidentally, after Dan Hinkley’s collection). No mention of the plant in cultivation or the contribution of Dan is found in the publication despite the fact that quite a few of the Begonia aficionado crowd around the globe had by then become familiar with the plant. Regardless of the name, this species is among the most spectacular hardy garden plants for cool but not cold climates.
Begonia lorentzoniiBegonia lorentzonii
Begonia lorentzonii has proven hardy at Heronswood (zone 8a) where it was left outside with only a covering of leaves and straw in temperatures ranging into the low teens and at least a week long stretch of consistently below freezing temperatures which resulted in ground freeze. It forms 2-2.5’ tall dense clumps with one of the best forms I’ve seen in a cold hardy species.
In late summer through late autumn it is adorned with yellow flowers beset with hairlike projections on the outer surface of the tepals produced on stems that equal or are slightly shorter than the leaves. Begonia lovers should visit the Renaissance Garden at Heronswood to see mature plants in their full glory and a pilgrimage to Heronswood is a must for all hardy Begonia lovers as the collection of rare and unusual cold hardy species is probably the best among our public gardens. While this startling plant appears to be perfectly adapted to life in the mild Pacific Northwest it remains to be seen what its tolerance for heat will be. It was an honor and pleasure to grow and nurture these plants during my time at Heronswood and I must admit my heart and mind will forever be drawn to that sacred space of ground whenever I glimpse a Begonia of any species.
