Buckthorns Gone Wild

Colletia paradoxa photo by James Gaither licensed by CC BY-NC-ND 2.0

When I think of the buckthorn family (Rhamnaceae), my mind conjures up images of battling with Rhamnus invasions around the Great Lakes or the amazing diversity of Ceanothus in western North America. Never have my thoughts drifted to the bizarre and wonderful genus Colletia. Native to temperate regions of South America, this strange group of spiny shrubs is certainly worth a closer look.

Though new to me, the genus Colletia has been known to science and horticulture since at least the late 1700’s. Hailing from temperate climates, at least two of the five known species of Colletia have found there way into temperate gardens elsewhere. Who could blame gardeners for their fascination with these shrubs. Close inspection of Colletia reveals surprisingly complex morphological features.

Colletia paradoxa

For starters, those large, thick, leaf-like thorns are not leaves at all. They are flattened extensions of the stem called cladodes. Instead of relying on leaves for most of their photosynthetic needs, the various Colletia instead produce chlorophyll in their stems. The cladodes function in much the same way as leaves in that their increased surface area maximizes photosynthetic potential. It is likely that cladodes are a means of conserving valuable resources for the plant.

Instead of producing vulnerable leaves that are subject to plenty of damage, these shrubs simply utilize stem tissues. Stems don’t need to be regrown year after year and by adorning the tips of the cladodes with spines, the plant is better able to protect its photosynthetic tissues. That is not to say that Colletia produce no leaves at all. Colletia will produce leaves near the base of each cladode, especially on younger tissues. Leaves, however, are deciduous and don’t stick around long enough to do much photosynthesizing.

Colletia ulicina with its red, tubular flowers. Photo by FarOutFlora licensed by CC BY-NC-ND 2.0

The flowers of Colletia ulicina are pollinated by hummingbirds. photo by James Gaither licensed by CC BY-NC-ND 2.0

Colletia are made all the more noticeable when they come into flower. For most species, clusters of lightly-scented, white flowers are produced at the base of the cladodes. For these species, insects are thought to be the predominant pollinators. Such is not the case for Colletia ulicina. This species produces sprays of bright red, tubular flowers along its stems. In the wild, these are pollinated by the green-backed firecrown hummingbird (Sephanoides sephaniodes).

Another interesting aspect of Colletia ecology is that they are all nitrogen fixers. To be fair, the plants themselves don’t do any of the fixing. Instead, they produce tiny structures on their roots called “nodules,” and those nodules house specialized bacteria collectively referred to as actinomycetes. In exchange for carbohydrates produced via photosynthesis, these bacteria fix nitrogen from the air. This extra boost of nitrogen allows Colletia to survive and excel in the nutrient-poor soils they call home.

Photo Credits: [1] [2] [3] [4]

Further Reading: [1] [2] [3]

Meet the Ocotillo

I love the ocotillo (Fouquieria splendens) for many reasons. It is an impossible plant to miss with its spindly, spine-covered stems. It is a lovely plant that is right at home in the arid parts of southwestern North America. Beyond its unique appearance, the ocotillo is a fascinating and important component of the ecology of this region.

My first impression of ocotillo was interesting. I could not figure out where this plant belonged on the tree of life. As a temperate northeasterner, one can forgive my taxonomic ignorance of this group. The family from which it hails, Fouquieriaceae, is restricted to southwestern North America. It contains one genus (Fouquieria) and about 11 species, all of which are rather spiky in appearance.

Of course, those spines serve as protection. Resources like water are in short supply in desert ecosystems so these plants ensure that it is a real struggle for any animal looking to take a bite to get at the sap inside. Those spines are tough as well. One manged to pierce the underside of my boot during a hike and I was lucky that it just barely grazed the underside of my foot. Needless to say, the ocotillo is a plant worthy of attention and respect.

One of the most striking aspects of ocotillo life is how quickly these plants respond to water. As spring brings rain to this region of North America, ocotillo respond with wonderful sprays of bright red flowers situated atop their spindly stems. These blooms are usually timed so as to take advantage of migrating hummingbirds and emerging bees. The collective display of a landscape full of blooming ocotillo is jaw-droppingly gorgeous and a sight one doesn't soon forget. It is as if the whole landscape has suddenly caught on fire. Indeed, the word "ocotillo" is Spanish for "little torch."

Flowering isn't the only way this species responds to the sudden availability of water. A soaking rain will also bring about an eruption of leaves, turning its barren, white stems bright green. The leaves themselves are small and rather fragile. They do not have the tough, succulent texture of what one would expect out of a desert specialist. That is because they don't have to ride out the hard times. Instead, ocotillo are what we call a drought deciduous species, producing leaves when times are good and water is in high supply, and dropping them as soon as the soil dries out.

This cycle of growing and dropping leaves can and does happen multiple times per year. It is not uncommon to see ocotillo leaf out up to 4 or 5 times between spring and fall. During the rest of the year, ocotillo relies on chlorophyll in its stems for its photosynthetic needs. Interestingly enough, this poses a bit of a challenge when it comes to getting enough CO2. Whereas leaves are covered in tiny pours called stomata which help to regulate gas exchange, the stems of an ocotillo are a lot less porous, making it a challenge to get gases in and out. This is where the efficient metabolism of this plant comes in handy.

All plants undergo respiration like you and me. The carbohydrates made during photosynthesis are broken down to fuel the plant and in doing so, CO2 is produced. Amazingly, the ocotillo (as well as many other plants that undergo stem photosynthesis) are able to recycle the CO2 generated by cellular respiration back into photosynthesis within the stem. In this way, the ocotillo is fully capable of photosynthesis even without leaves.

Through the good times and the bad, the ocotillo and its relatives are important components of desert ecology. They are as hardy as they are beautiful and getting to see them in person has been a remarkable experience. They ad a flare of surreality to the landscape that must be seen in person to believe.

Further Reading: [1] [2] [3] [4] [5]

Mt. Cuba Center Puts Nativars to the Test

By this point, most gardeners will have undoubtedly heard about the importance of using native plants in our landscapes. Though the idea is not new, Doug Tallamy’s landmark publication “Bringing Nature Home” put native plants on the radar for more gardeners than ever. There is no debate that utilizing native plants in our landscapes offers us a chance to bring back some of the biodiversity that was lost when our homes and work places were built. And, at the end of the day, who doesn’t love the sight of a swallowtail butterfly flitting from flower to flower or a pair of warblers nesting in their Viburnum? The rise of native plants in horticulture and landscaping is truly something worth celebrating.

At the same time, however, capitalism is capitalism, and many nurseries are starting to jump on the bandwagon in alarming ways. The rise of native cultivars or “nativars” is troubling to many. Nativars are unique forms, colors, and shapes of our beloved native plants which have been selected and propagated by nurseries and plant breeders. This has led many to denounce the practice of planting nativars as a slap in the face to the concept of native gardening.

Nativars are frequently seen as unnatural mutant versions of their wild counterparts whose use overlooks the whole point of natives in the first place. Take, for instance, the popularity of double flowered nativars. These plants have been selected for an over-production of sepals and petals that can be so dense that they preclude visitation by pollinators. Another example that will be familiar to most are the bright blue hydrangeas that have become to popular. These shrubs have been selected for producing bright, showy flowers that, depending on your soil chemistry, exhibit a stunning blue coloration. The downside here is that all of those flowers are sterile and produce no nectar or pollen for visiting insects.

It would seem that nativars are a slippery slope to yet another sterile landscape incapable of supporting biodiversity. However, anecdotes don’t equal data and that is where places like Mt. Cuba Center come in. Located in northern Delaware, Mt. Cuba is doing something quite amazing for the sake of environmentally friendly landscaping – they are putting plants to the test.

Mt. Cuba has been running trial garden research and experiments on native plants and their nativars for over a decade. The goal of this research is to generate and analyze data in order to help the public make better, more sustainable choices for their yards. Mt. Cuba aims to better understand and quantify the horticultural and ecological value of native plants and related nativars in order to better understand the various ecosystem services these plants provide. In collaboration with academic institutions in the region, popular nativars are established and grown under similar conditions to those experienced in the yards of your average gardener. They are monitored for years to assess their overall health, performance, and ability to support wildlife. Thanks to the help of countless volunteers, these trial gardens paint a holistic picture of each plant and related nativars that is sorely lacking from the gardening lexicon.

This is very exciting research to say the least. The data coming out of the Mt. Cuba trial gardens may both surprise and excite gardeners throughout the mid-Atlantic region of North America. For instance, their latest report looked at some of the most common Phlox varieties on the market. At the top of this list is Garden Phlox (Phlox paniculata). This lovely species is native throughout much of the eastern United States and has become quite a rockstar in the nursery trade. Over 580 cultivars and hybrids have been named to date and no doubt many more will be introduced in the future. Amazingly, many of these Phlox nativars are being developed in the Netherlands. As such, Phlox arriving in regions of the US with vastly different climates often fall victim to novel diseases they never encountered in Europe. What’s more, people often plant these nativars in hopes of attracting butterflies to their garden. Despite their popularity for attracting various lepidopterans, no one has ever tested whether or not the nativars perform as well as their native progenitor.

Phlox paniculata 'Delta Snow'

Starting in 2015, Mt. Cuba began trials on 66 selections and hybrids of Garden Phlox along with 28 other sun-loving types of Phlox. The plants were observed on a regular basis to see which of the nativars experienced the least amount of disease and attracted the most insects. The clear winner of these trails is a nativar known as Phlox paniculata ‘Jeana’. This particular selection was discovered growing along the Harpeth River in Tennessee and is known for having the smallest flowers of any of the Garden Phlox varieties. It also has the reputation for being rather resistant to powdery mildew. Alongside other selections such as Delta Sno’ and David, Jeana really held up to this reputation.

As far as butterflies are concerned, Jeana blew its competition out of the water. Throughout the observation period, Jeana plants received over 530 visits from butterflies whereas the second place selection, Lavelle, received 117. A graduate student at the University of Delaware is studying why exactly the various nativars of Phlox paniculata differ so much in insect visitation. Though they haven’t zeroed in on a single cause at this point, they suggest that the popularity of Jeana might actually have something to do with its small flower size. Perhaps the density of smaller flowers allows butterflies to access more nectar for less effort.

Phlox paniculata ‘Jeana’

Monarda is another genus of North American native plants that has seen an explosion in nativars and hybrids over the last few decades. The popularity of these mints is no surprise to anyone who has spent time around them. Their inflorescence seems to be doing their best impression of a fireworks display, an attribute that isn’t lost on pollinators. These plants are popular with a wide variety of wildlife from solitary bees to voracious hummingbirds. Even after flowering, their seeds provide food for seed-eating birds and many other animals.

As with Garden Phlox, a majority of the commercial selection and hybridization of Monarda occurs in Europe. As a result, resistance to North American plant diseases is not top priority. Many of us have experienced this first hand as our beloved bee balm patch succumbs to aggressive strains of powdery mildew. Though there are many species of Monarda native to North America, most of the plants we encounter are nativars and hybrids of two species – Monarda didyma and Monarda fistulosa.

Monarda fistulosa 'Claire Grace'

Again, Mt. Cuba’s trial gardens put these plants to the test. A total of 40 different Monarda selections were grown, observed, and ranked based on their overall growth and vigor, pollinator attractiveness, and disease resistance. The clear winner of these trials was a naturally-occurring form of M. fistulosa affectionately named ‘Claire Grace.’ Its floral display lasts a total of 3 weeks without waning and managed to attract over 130 visits by butterflies and moths. Though plenty of other insects such as short-tongued bees visited the flowers during the trial period, they are too small to properly access the nectar inside the flower tubes and are therefore not considered effective pollinators.

Another clear winner in terms of pollinators was possibly one of the most stunning Monarda selections in existence – Monarda didyma ‘Jacob Cline’. This tall, red-flowering nativar was a major hit with hummingbirds. During the observation period, Jacob Cline received over 270 visits from these brightly colored birds. Researchers are still trying to figure out why exactly this particular selection was such a hit but they speculate that the large flower size presents ample feeding opportunities for tenacious hummingbirds.

Monarda didyma 'Jacob Cline'

Claire Grace and Jacob Cline also outperformed most of the other selections in terms of disease resistance. Even in the crowded conditions experienced by plants in the trail garden, both selections faired quite well against the dreaded powdery mildew. Though they aren’t completely resistant to it, these and others did not succumb like some selections tend to do. Interestingly enough, most of the other pure species tested in the trial faired quite well against powdery mildew as well. It would appear that Mother Nature better equips these plants than European breeders.

These reports are but two of the many trials that Mt. Cuba has undertaken and there are many, many more on the way. Thanks to the hard work of staff and volunteers, Mt. Cuba is finally putting numbers behind some of our most commonly held assumptions about gardening with native plants and their cultivars. It is impressive to see a place so dedicated to making our landscapes more sustainable and environmentally friendly.

If you would like to find out more about Mt. Cuba’s trial garden as well as download your own copies of the trial garden reports, please make sure to check out https://mtcubacenter.org/research/trial-garden/

Pollination Plasticity

© Danny Keßler

Pollinators are great -- that is, unless they also feed upon the plant they are pollinating. In the arid regions of western North America, Nicotiana attenuata, sometimes referred to as coyote tobacco, has this very problem. 

Blooming at night, its white flowers are heavily scented, which attracts its pollinator, a species of hawkmoth known to science as Manduca quinquemaculata. Female hawkmoths do a little bit more than just grab a sip of nectar. Their larvae feed on members of the tobacco family and, as anyone with tomatoes can tell you, they have a voracious apatite. Visiting female moths use the meal break as a chance to lay their eggs. However, this does not have to be a death sentence for the plant. Researchers noticed a strange thing about N. attenuata plants that had feeding damage from hawkmoth caterpillars. Their flowers seemed to change.

Photo by Stan Shebs licensed under CC BY-SA 3.0

And change they did. Coyote tobacco plants with caterpillars will start to produce flowers that open during the day, instead of at night. The plants also stopped producing a scent. What's more, the flowers didn't open very far either. What is the reason for these drastic changes? Are the plants stressed out from the caterpillar attack?

Not exactly. In fact, the answer is quite remarkable. As it it turns out, plants with caterpillars munching on them were intentionally shifting their entire reproductive strategy to avoid the larvae of their intended pollinators. Flowers that open during the day no longer attracted the attention of moths, which reduced the number of new eggs being laid. Instead, the flowers started attracting the attention of hummingbirds. Hummingbirds are pretty effective as pollinators and their offspring don't eat the plants that their parents feed on. 

Manduca quinquemaculata adult male. Photo by Didier Descouens licensed under CC BY-SA 4.0

So, how does the plant know when its being fed upon? Caterpillar spit. Chemicals in the saliva of the caterpillar trigger a chemical response within the plant that tells it to start ramping up defenses (of which nicotine is one). This signaling cascade also tells the plant to start producing day opening flowers instead of night opening flowers. It just goes to show you how a little attention to detail can uncover some amazing aspects of the world around us. 

Photo Credit: Danny Kessler, MPI chemische Ökologie, Wikimedia Commons

Further Reading: [1] [2]