Plant Ecology

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Tea Time: Self-heal

Prunella vulgaris can be found all over the place. It has also been used to treat just about everything. What else would you expect from a plant known commonly as self-heal, heal-all, all-heal, and woundwort? The medicinal value of this plant has been appreciated for centuries across its expansive range, and studies evaluating its medicinal use continue today. Being such a ubiquitous species – both as a garden plant and a native plant (and also a common weed) – and because it has so much clout in the world of herbal medicine, it’s an obvious candidate for Tea Time.

Self-heal is a member of the mint family (Lamiaceae), easily distinguished by its square stems, opposite leaves, and bilabiate and bilaterally symmetrical (or zygomorphic) flowers. One surprise is that, unlike the many aromatic members of this family, the foliage of self-heal lacks a strong scent. P. vulgaris occurs naturally across Asia, throughout Europe, and in parts of northern Africa. It is also widely distributed across North America. Apart from that, it has been introduced to many regions in the southern hemisphere and has also been frequently moved around throughout its native range. Eurasian varieties now intermingle with North American varieties, which can make it difficult to determine a native individual from an introduced one.

self-heal (Prunella vulgaris)

Self-heal is an adaptable plant that tends to prefer shady, moist locations, but can also be found in open, dry, sunny sites. Find it along forest edges, roadsides, ditches, and trails, as well as on the banks of streams, lakes, and reservoirs. It occurs in gardens, both intentionally planted and as a weed, and can escape into lawns, vacant lots, and open fields, as well as into nearby natural areas.

P. vulgaris is an evergreen that grows both prostrate and upright, sometimes reaching 1 foot tall or more (but is often much shorter). It has shallow, fibrous roots, and its stems root adventitiously as they sprawl across the ground, frequently forming an extensive mat or groundcover. Its leaves are oval to lance-shaped and measure about one inch long. Lower leaves have petioles, while upper leaves may become stalkless. Leaf margins are entire or can be slightly toothed. As plants age, they can develop a coppery or purple-bronze color.

the leaves of self-heal

The flowers of self-heal are generally a shade of purple, but can also be white, pink, or blue. They bloom irregularly in a spike measuring up to two inches long. Flower spikes are thick, dense, cylindrical, and made up of whorls of sharp-pointed bracts. Flowers bloom irregularly along the spike and occur from late spring/early summer into the fall. Each flower produces four nutlets, which sit within a cup-shaped, purple calyx.

As a medicinal herb, self-heal has been used both internally and externally to treat a long list of ailments. These include sore throats, diarrhea, fevers, intestinal infections, liver problems, migraines, heart issues, dermatitis, goiter, and thyroid disfunction, just to name a few. It has been used topically to treat skin irritations, bites, stings, and minor cuts and scrapes. This is thanks to its antimicrobial properties and its ability to stop bleeding. A report in the journal Pharmaceuticals (2023) calls P. vulgaris an “important medicinal plant” due to its “rich chemical composition” and its “pharmacological action.” Chemical analyses find the plant to be a valuable source of phenolic compounds, flavonoids, rosmarinic acid, and ursolic acid, among numerous other compounds. If you are curious to learn more detailed information regarding this plant’s medicinal value, you can refer to the above report, as well as one found in Frontiers in Pharmacology (2022).

self-heal tea

P. vulgaris is an edible plant, and its young leaves can be eaten raw or cooked. The leaves together with the flowers can also be dried and used to make a tea. This is how I had it. I used about two teaspoons of dried leaves to one cup of water. Feel free to use more if you would like. I thought the tea was pretty mild. It had a slight sweetness to it and a hint of mint flavor. It has been described as bitter, but I didn’t find it to be overly so (although I may have a higher tolerance for bitterness). Sierra tried it and said that it tasted like “water left over from something else.” That might be because it was more diluted than she would have preferred. Overall, I thought it was a pleasant experience and would be happy to drink it again.

More Tea Time Posts on Awkward Botany:

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Garden Plants Gone Wild: The Periwinkles

In a garden setting, a successful groundcover is a plant that is durable and adaptable, spreads readily, and fills in space thouroughly. The point of planting a groundcover is to cover exposed soil and create a sort of living mulch. In fact, groundcovers provide similar benefits to mulch. They prevent erosion, help retain soil moisture, and prevent weeds. It should come as no surprise then, that a plant that fulfills all of these requirements has the potential to become a weed, especially if given the opportunity to escape and establish itself outside of its intended location.

This isn’t a hypothetical. This exact scenario has played out numerous times. A good example of this are the periwinkles: Vinca major and Vinca minor. Both have been popular garden plants for centuries. Their introduction to U.S. gardens came as early as the 1700’s. Today, both species (including cultivars of each) can be found for sale in nurseries throughout the country, even while escaped periwinkles proceed to spread across natural areas and uncultivated spaces. Even in gardens where periwinkles have been intentionally planted, they can be deemed no longer welcome due to their aggressive nature. Eliminating them, however, is a formidable task.

greater periwinkle (Vinca major)

Periwinkles are relatively easy to identify, yet telling the two apart can sometimes pose a challenge. Knowing what to look for can make this a fairly simple task. Vinca major (greater periwinkle) is the larger of the two. Its stems are tough and sprawl up to 5 feet long, rooting adventitiously when in contact with the ground. Vegetative spread also occurs via stolons and rhizomes, modified stems that spread horizontally both above and below ground and whose main purpose is to produce new roots along their length. This extensive rooting creates dense mats of stems and foliage, precisely what you’d want from an effective groundcover. Leaves are arranged oppositely and are semi-evergreen with fine hairs along their margins. They are thick, glossy, simple, dark green, and ovate to ovate-lanceolate in shape. They have a relatively long petiole, and some leaves can even appear heart-shaped.

The flowers of V. major are blue to purple and fused at the base to form a tube, separating into five distinct lobes and creating a shape similar to a pinwheel. They are borne on a long stalk in the axils of leaves and measure about two inches wide. Their sepals are long, slender, pointed, and lined with bristly hairs. Fruits rarely form, but when they do, they are narrow follicles.

comparing the petals and sepals of Vinca major (left) and Vinca minor (right)

Vinca minor (lesser periwinkle) is a slightly smaller plant with a similar habit, spreading vegetatively in all the same ways as V. major. Stems are slender and smooth and leaves are evergreen. Compared to V. major, the leaves of V. minor are smaller, narrower, and have hairless margins and short petioles, otherwise they are very similar. Same goes for the flowers, which look identical on both species, except that those of V. minor are slightly smaller (about one inch wide) and borne on shorter stalks. Sepals are shorter, broader, more rounded at the tips, and lack the bristly hairs of V. major.

Because periwinkles only rarely produce seed, their main method of getting around is vegetatively. Fragments of roots or rhizomes hide in soil and are moved from one location to another inadvertently. Periwinkles are often used in hanging baskets and containers, and when these things are cast aside at the end of a season, the perennial roots of periwinkles may continue to grow, spreading out beyond the potting mix and into the soil.

Dump soil, yard waste, and improperly disposed of containers are the main ways that periwinkles find their way into natural areas. Both species can be found in the understories and edge habitats of woodlands, as well as along roadsides and pathways, and in vacant lots and old homesites. They can also be found in riparian areas, where waterways can carry fragments of plants to new locations. The Invasive Plant Atlas compiles reports of both V. major and V. minor growing outside of cultivation and tracks them on a map. They also track which states include them in noxious weeds lists or laws.

lesser periwinkle (Vinca minor)

The best way to keep periwinkles from continuing to spread outside of cultivation is to refrain from growing them. If you choose to have them in your yard, dispose of plant parts properly. If you keep them in containers, send those containers to the landfill when you are done with them. If your property is adjacent to natural areas, the risk may be too great and you may want to consider a different groundcover. Depending on where you live, alternatives vary. In the Intermountain West, potential substitutes include wild strawberry (Fragaria virginiana), woodland strawberry (F. vesca), kinnikinnik (Arctostaphylos uva-ursi), and wild ginger (Asarum caudatum). Each of these are low growing, evergreen to semi-evergreen, spreading plants that do well in shade and can handle some degree of drought.

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Horticulture’s Weedy Introductions in a Changing Climate

In case I need a reminder that the horticulture industry has a history of introducing weedy plants to natural areas, I get one each time I bike to work. Riding along the Boise River Greenbelt, a trail that for much of its length is flanked by cultivated landscapes on one side and a highly modified but largely naturalized river bank on the other, I see a mixture of both native and introduced plants. Of the introduced plants, many are horticultural species that have escaped cultivation and established themselves on the bank of the river. There are catalpa and black locust trees brought in from the other side of the country, St. John’s wort and chicory from Eurasia, honeysuckles primarily from Asia, and a few different cherry species and hybrids with varied provenances. And this is just a small sample of what can be found along my three and a half mile bike ride.

St. John’s wort (Hypericum perforatum) on the banks of the Boise River

This is certainly not a new concern. We have been aware of the role that horticulture plays in introducing invasive species for quite some time now. Several years back, while doing a deep dive into the topic of invasive species, I wrote about this issue right here on this very blog. According to a study published in Frontiers in Ecology and the Environment (2021), out of 1285 plant species identified as invasive, 61% are currently sold in nurseries. If that’s not concern enough, an additional factor to consider is climate change. Plants that were less likely to escape cultivation and head for the wild, may take the opportunity to do so in a changing climate. Plus, horticultural plants that are already problems in certain areas could expand their range as climates become more favorable in new locations, especially if these plants continue to be sold in nearby nurseries.

These concerns and more are the topic of a paper published in BioScience (2023). Evelyn M. Beaury, et al. looked at nurseries across the United States and the plants they sell in order to determine where invasive plants are still being sold in regions where they are invasive. Additionally, they looked at plants known to be invasive but that are not currently invasive in the regions they are being sold. Using climate models, they predicted whether or not these plants could become invasive under changing climates.

Plants are being moved around with a lot more ease than they once were, and the sales of problematic plants are increasingly difficult to regulate. For one thing, plants prohibited for sale in one state can be purchased at nurseries in neighboring states and brought back to be planted in regions where those plants are invasive. And while mail order has existed for a long time, online ordering makes the process even simpler; and many online plant vendors are not liscensed nurseries, making them much more difficult to regulate. But even regulation is typically a response to something that has already become a problem, rather than a proactive measure to prevent plants from escaping into natural areas.

Beaury, et al. identified 672 nurseries across the United States, both online and traditional retailers. Each of these nurseries were selling one or more of the 89 plant species that became the focus of their research. These are plant species that are either on federal or state noxious weed lists or that have been identified as invasive by Invasive Plant Atlas. The reach of each nursery was determined by using customer reviews to compute distances that plants might travel after being purchased at nurseries or from online stores. Obviously, not every customer that purchases a plant leaves a review, but this is a good way to get a general idea how far away customers are from nurseries without having access to more detailed records. These geotagged reviews can also be cross-referenced with known distributions of invasive plants. Using climate models and environmental predictor variables, the researchers determined areas of current and potential invasion for each of the 89 plants.

tansy (Tanacetum vulgare) – one of the 89 plant species looked at in the study

The first question was about proximity to current records of plant invasions. Results showed that “49 of the 89 ornamental invasives were sold within 21 kilometers (13 miles) of an observed record of invasion.” When invasive plants are sold and planted near locations where they are already known to be invasive, it gives them the opportunity to add new plants to existing or developing invasions. In ecology, this is known as propagule pressure. When it comes to current and future climate, most species in the study are being sold by nurseries where the climate is either currently favorable for range expansion or may eventually become favorable. Specifically for future climate, 40 of the 89 plants are being sold in regions that are currently suitable for invasion and will continue to be suitable as the climate changes, and 25 of the 89 plants are being sold in regions where the climate is currently unsuitable but will become suitable as temperatures warm.

Particularly for plants being sold in areas that are not yet suitable for invasion, there is time to educate both the nursery industry and the general public and to look for alternatives to these plants. However, as the researchers point out, their analysis “only examined about 10% of the larger pool of U.S. ornamental plants known to be invasive,” and they “sampled only a subset of the nurseries that could be selling invasive species in the United States.” It is highly likely that the results of this study are an underestimation of the problem. Clearly the work of education and finding alternatives to problematic plants is monumental. The hope is that studies like this can help with education and can assist with working out ways to regulate sales of invasive plants.

coltsfoot (Tussilago farfara) – another one of the 89 plant species looked at in the study

Regulating the sale of plants is beyond most of our control, and how much regulation we should be enforcing on nurseries in the first place is a debate we should be having. Outside of those questions, there is a responsibility that we should take as gardeners and as residents of the planet. If we choose to grow plants, it is crucial that we get to know them. We should be taking the time to observe the degree to which they spread and how they are being dispersed. When they do move around our yards, where are they going, and are they able to grow outside of our care? Are they leaving our properties and coming up elsewhere? If we choose to plant non-native species, we should be mindful of how they might affect nearby, wild landscapes if they were to escape our yards and establish themselves in these locations. We should also be aware of where we live in the city. If our gardens are in the middle of a dense urban landscape, perhaps there is less concern that our plants will move beyond the borders of our gardens. But if we garden near natural areas, we should be significantly more selective about the things we plant, and we ought to be more observant as to what those plants are up to.

Nurseries generally sell the plants that gardeners want to buy, which means we can choose not to buy problematic plants and instead demand alternatives to these plants. Seeking out nurseries that sell the types of plants that are better suited for our regions and do not exhibit invasive behaviors can send a message to other growers that they should phase out certain plants and start growing the plants that gardeners are asking for. This may be a simplistic take, and as with most things, it’s complicated. While one of the goals of this research is to help influence regulators, another goal is simply to “[share] information about high-risk ornamental invaders across states and regions, and [work] with horticulture and community members to reduce the escape of ornamental species into natural areas.” This is precisely the area where gardeners can make a difference.

On that note, I will be starting a new series of posts to discuss some of the ornamental species that have gone weedy. By getting to know the plants that find themselves in this predicament, we can be better situated to make informed decisions about what to do about them.

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Winter Trees and Shrubs: Box Elder

Box elder is a maple that doesn’t often get credit for being a maple. Moreso, it is a tree that is not thought highly of, and it may not even be welcome in certain discussions around maples. You could even say that box elder is a “rogue maple,” as Arthur Plotnick deems it in The Urban Tree Book. It should come as no surprise, but if people are going to talk about a plant this way, it’s only going to make me like it more.

First off, the leaves of Acer negundo are palmately compound, compared to the simple, palmately lobed leaves of other maples. A box elder leaf is generally composed of three leaflets, sometimes five or seven. The center leaflet is on a longer stalk than the side leaflets, which are oppositely arranged. Sometimes the leaves are confused with ash tree leaves (Fraxinus spp.), which is the reason for another common name, ash-leaved maple. It’s also not uncommon for box elder leaves to be mistaken for poison ivy, which is maybe not a plant you’d want to be confused with. In the fall, unlike the beautiful reds, oranges, purples, and yellows of so many other maples, the leaves of box elder turn a drab yellow before dropping, a trait that Plotnick calls “an embarrassment.”

leaf of box elder (Acer negundo)

It’s not really its leaf, however, that bothers people. Box elder is a fast-growing and prolific tree. Basically, it’s weedy, and because it grows so quickly, its wood is generally weak and its structure unpredictable. It falls apart easily, and it doesn’t always grow in typical tree fashion. Sometimes its more of a shrub, and sometimes it leans so far over that it practically lies on the ground. It sprouts and suckers in abundance, and as John Eastman puts it in The Book of Forest and Thicket, it “generally ignore(s) human standards for an attractive tree.”

Maybe before people knew better, or during a time when people just needed a fast growing tree to plant in their developing cities, box elders were widely planted across North America and beyond. That and their ability to spread readily helps to obscure their native distribution. Box elder is naturalized across North America, but its original distribution (whatever that means) appears to be more midwestern and eastern. Box elder grows in a variety of conditions, and is actually quite drought tolerant, which has allowed it to feel right at home in the American west, whether it is truly native here or not.

Identifying box elder in the winter can be pretty easy, especially if you come across a seed-bearing individual. Box elders are dioecious, meaning male and female flowers occur on separate individuals (another thing that sets them apart from other maples). A samara-bearing tree is a female tree, and the samaras often hang from the branches in great numbers. Like other maples, the fruits are paired together, each seed with a prominent wing. The samaras of boxelder point downward and hang from long stalks, persisting on the tree throughout the winter.

persistent samaras of box elder

The twigs of box elder are glaucous (covered with a thin, powdery bloom) and can be either green, bluish-white, or reddish-purple. They have tiny, white lenticles, and their buds are oppositely arranged. Leaf scars are a broad U-shape, and scars opposite of each other meet at the tops of the U’s, coming to a raised point. At least three bundle traces (sometimes more) are visible in the scar. Leaf buds are protected by two scales that are covered in dense, white hairs. The scales are similar in color to the twigs, but appear more white due to the fuzz. Lateral buds are 1/8 inch long or longer. Slicing open the twigs lengthwise reveals solid, white pith in the center that blends in easily with the wood.

winter twigs of box elder
winter twigs of box elder, pith exposed

The bark of young box elders has slight vertical ridges and is light brown or grey-brown, with dull orange, raised, wart-like lenticels. It is finely scaly. The scales and ridges become more rough and blocky as the tree ages. Mature bark is light brown to grey and heavily furrowed, with blocky segments breaking apart horizontally.

maturing bark of box elder

A few years ago I was introduced to a massive box elder in Boise, Idaho. It quickly became my favorite tree in the area, and I visit it frequently. For all the hate that box elder can receive, seeing its potential to create an experience that few other trees in the area offer can send all of that negative talk out the window. Visit something like it yourself sometime, and see what it can do for you.

massive box elder

More Winter Trees and Shrubs:

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Weeds of Boise Takes a GIS Course

Why has this blog been so quiet lately? There are plenty of excuses for that. It doesn’t really matter either way, but since we’re on the subject, one thing that has kept me occupied recently is being back in school. I’m working on a certificate in GIS, and I’m hoping to make some cool maps. More on that later perhaps.

For now, I thought I’d share one of my final projects. I figured it was a good excuse to use something I’ve already been working on – namely, Weeds of Boise – and apply it to one of my GIS classes. If you’ve been following Weeds of Boise, not much here will be new. Except the interactive maps!

Using observations from iNaturalist, I created three different maps of weeds found around Boise. I published them, using ArcGIS StoryMaps, alongside more of my usual pontificating about weeds and urban areas, etc. The maps aren’t perfect, but they opened up some interesting possibilities for what Weeds of Boise might become. Something to explore further in the future.

Until then, here is a link to the project if you’d like to take a look: Wild Urban Flora of Boise Idaho.

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Randomly Selected Botanical Terms: Phyllaries

Flowers in the aster family have one of the most recognizable shapes in botany – a circle with a series of petals surrounding it. If you were asked to draw a flower, there is a good chance your drawing would look something like a sunflower, a daisy, a cosmos, or an aster. It’s one of the most basic flower shapes, and yet it isn’t a single flower; it’s a pseudanthium – a false flower. This is because what might appear as a single flower is actually a collection of tens, hundreds, or even thousands of tiny flowers. This aggregation of flowers into a single compact unit is the reason the family was once given the name Compositae, and even now is often informally referred to as the composites.

Another reason why a flower in the aster family – or Asteraceae – might be the first thing you would draw is because it is the largest family of flowering plants, numbering up to 33,000 species worldwide. Chances are you’ve seen a few of them around. In the contiguous U.S. alone, there are more than 2400 species, and that doesn’t include the plethora of species brought in from regions across the world either intentionally (to be grown in our gardens) or unintentionally (as weeds). Of course, not all of the species in this family are going to have a typical sunflower-like flower head, but they do all have a specific type of inflorescence called a capitulum. Capitula are made up of densely packed, miniature flowers called florets, which are stalkless (or sessile) and arranged on a flattened central stem (or axis). There are at least four different types of florets in the aster family, but we’ll leave that discussion for another time.

In this post, we’re specifically interested in what is happening at the base or underside of the capitulum. All of the florets in a capitulum are held within a cup or bowl-shaped series of bracts called an involucre. Bracts are modified leaves, and this whorl of tightly held or loosely arranged bracts are initially found surrounding a developing flower bud. As the inflorescence opens, the involucre opens as well and its bracts persist at the base of the flower head. The bracts that make up the involucre are called phyllaries, and they vary in shape, number, and size depending on plant species. In fact, the features of phyllaries are so unique they are often relied on to help identify a plant in the aster family to genus, species, and infraspecies (variety, subspecies, etc.).

phyllaries of blanketflower (Gaillardia aristata)

When it comes to flowers in the aster family, there is more than meets the eye. After you take some time to appreciate the intricate beauty of its collection of florets, turn the flower head over and take in its phyllaries. They come in various colors, they can be hairy or smooth, their margins can be entire or adorned with hairs, teeth, etc., they can be flat and straight or they can curve outwards in interesting ways, their tips can be pointed, spine-tipped, rounded, or keel shaped. Phyllaries can be laid out very evenly, tightly overlapping each other like shingles on a roof (i.e. imbricate) or their arrangement can be slightly uneven and irregular (i.e. subimbricate). Use a hand lens to get a closer look at all of these features. As you get in the habit of observing the details of the involucre and its phyllaries, chances are each time you come across a flower in the aster family, you’ll find yourself flipping it over to get a look at its undercarriage. What will you find?

phyllaries of dandelion (Taraxacum officinale)
phyllaries of Mexican sunflower (Tithonia rotundifolia)
phyllaries of stemless four-nerve daisy (Tetraneuris acaulis)
phyllaries of hoary tansyaster (Dieteria canescens)
phyllaries of aromatic aster (Symphyotrichum oblongifolium ‘October Skies’)
phyllaries of curlycup gumweed (Grindelia squarrosa)

If phyllaries have piqued your interest and you’d like to learn more about plants in the family Asteraceae, I highly recommend getting your hands on the book, The Sunflower Family by Richard Spellenberg and Naida Zucker. It has a North American focus, but it’s a great place to start learning more about this massive plant family.

More Randomly Selected Botanical Terms:

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Another Year of Pollination: Viscin Threads

While we’re on the subject of pollen-gluing mechanisms, there is another material apart from pollenkitt that a limited number of flowering plant families use to link their pollen grains together. It functions, much like pollenkitt, by aiding in the attachment of pollen to visiting animals. However, unlike pollenkitt, it isn’t sticky, oily, or viscous, and is instead more like a series of threads. Viscin threads to be exact.

One of the major differences between pollenkitt and viscin threads is their composition. The lipid-rich coating that surrounds pollen grains, which we call pollenkitt, is derived from breakdown materials of an inner layer of the anther. It is added to pollen grains after they are formed and before the anther dehisces. Viscin threads are made up of sporopollenin, the same biopolymer that exine (the outer wall of a pollen grain) is composed of. Viscin threads have points of attachment on an outer layer of the exine called the ektexine. Unlike pollenkitt, viscin threads don’t add new color to pollen grains, nor do they contain scent compounds. Their thickness, length, abundance, and texture are dependent on the species of plant they are found on, much like pollenkitt varies in form and composition depending on species.

pollen strands of tufted evening primrose (Oenothera caespitosa)

Viscin threads evolved independently in three distantly related plant families. These include Onagraceae (the evening primrose family), Ericaceae (the heath family), and a subfamily in the pea family known as Caesalpinioideae (the peacock flower subfamily). Viscin threads are found in many, but not all, of the species in these three families. Some species in other plant families have what appear to be viscin threads but are actually ropy strands of pollenkitt, as they are composed of pollenkitt and not sporopollenin. Because they are made up of the same durable material as exine, viscin threads can be preserved in the fossil record. A paper published in Grana (1996) looked at the morphology of pollen grains with viscin threads from the Tertiary Period and concluded that “this advanced pollination syndrome using viscin threads as a pollen connecting agent” dates back to at least the Eocene and perhaps much earlier.

While pollenkitt’s stickiness adheres pollen grains together, viscin threads are more of a tangling device. Single pollen grains or pollen grain groupings called tetrads become tangled up together and then become entangled with a visiting insect, bird, or bat and carried away to a nearby flower. Disentanglement from the pollinator ideally happens when the threads are brushed against the sticky surface of a stigma. The viscin threads themselves vary by species and family. Micheal Hesse, in a paper published in Grana (1981), describes the threads in Onagraceae as “long, numerous, thin, and sculptured” with “knobs, furrows, etc.,” while those in Ericaceae are thin and smooth and those in Caesalpinioideae are thick and smooth.

smooth azalea, pink form (Rhododendron arborescens)

The length and size of tangled pollen masses also differ by species and can offer clues as to which pollinators visit which flowers. Research published in New Phytologist (2019) looked at the size of pollen thread tangles (PTT) in 13 different species of Rhododendron. They also noted which pollinators visited each species and how often they visited. The researchers found that species presenting pollen in small but abundant PTT were visited by bees, and those with large but few PTT were visited by birds and Lepidoptera (butterflies and moths). Bees also visited the flowers more frequently than birds and Lepidoptera. Bees collect and consume pollen. Between visits to anthers, they spend time grooming themselves, removing pollen clusters from their bodies and packing them into corbiculae (i.e. pollen baskets) for later*. Birds and Lepidoptera don’t groom pollen from their bodies and don’t collect it. In the authors terms, this “suggests pollinator-mediated selection on pollen packaging strategies.” Since flowers pollinated by bees lose much of their pollen in the process, they present it in smaller packages, and since flowers pollinated by birds and Lepidoptera are visited less frequently, their pollen packages are larger.

This is an example of the pollen presentation theory, and is something we will revisit as the Year of Pollination continues.

*This applies specifically to bee species that have corbiculae, and many bee species do not.

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Another Year of Pollination: Pollenkitt

Pollination in flowering plants is the process of moving pollen grains, which carry sperm cells, from the anthers to the stigmas of either the same flower or a separate flower. If things go well from there, sperm cells will be transported via pollen tubes into the ovaries where fertilization with egg cells can take place and seeds can form. Pollen grain development occurs within the anthers, and by the time the anthers dehisce – or split open – they are ready for transport.

In order to protect the enclosed sperm cells and aid in their movement, pollen grains consist of a series of layers that, among other things, help ensure safe travel. Two major layers are an internal layer called intine, composed largely of cellulose, and an external layer called exine, composed mainly of sporopollenin (a highly durable and complex biopolymer). In many flowering plants, especially those that rely on animals to help carry their pollen, an additional outer layer called pollenkitt is added to the pollen grains before anthers dehisce.

three different pollen grains (image credit: wikimedia commons/Asja Radja)

Pollenkitt is an oily, viscous, hydrophobic layer composed of lipids, carotenoids, flavonoids, proteins, and carbohydrates derived from the breakdown of an internal layer of the anther called the tapetum. Pollenkitt forms a sticky layer around the pollen grains and can add color to the pollen other than the typical yellow. The thickness of the pollenkitt and its composition is species specific. In fact, the look, size, and shape of pollen grains themselves are unique to each species and can even be used to help identify plants. Pollenkitt is found in almost all families of flowering plants and is particularly prevalent in species that are animal-pollinated. One exception is the mustard family (Brassicaceae), whose pollen grains are coated in a substance known as tryphine, which functions similar to pollenkitt but whose formation and composition differ enough to be considered separately.

dandelion pollen (image credit: wikimedia commons/Captainpixel)

The sticky nature of pollenkitt has numerous functions. For one, it helps pollen grains remain on anthers until an animal comes along to remove them. It also holds pollen grains together in clumps, helps pollen grains stick to insect (and other animal) pollinators during transport, and helps adhere them to stigmas when deposited. A paper published in Flora (2005) lists twenty possible functions for pollenkitt, many of which have been confirmed in certain species and some of which are hypothetical. In addition to functions having to do with pollen movement and placement, pollenkitt may also provide protection from water loss, UV radiation, and fungal and bacterial invasions. In species where pollen is offered as food to pollinating insects, pollenkitt is a more easily digestible food source than the pollen grain itself. Thanks to carotenoids, pollenkitt can make pollen more colorful, which may help attract pollinating insects, or, depending on the color, can also hide pollen from insect visitors.

Another important function of pollenkitt is to give pollen a scent. Odors can help encourage insect visitors or deter them, so depending on the situation, scented pollenkitt may be attracting pollinators or discouraging pollen consumers. In a study published in American Journal of Botany (1988), Heidi Dobson analyzed the chemical composition of 69 different species of flowering plants. She isolated numerous scent compounds in pollenkitt and suggested that “some of the chemicals in pollenkitt may … serve as identification cues to pollen-foraging bees.” Most of the species she analyzed were pollinated by bees (which consume pollen), but the few that were mainly pollinated by hummingbirds and butterflies tended to have fewer scent compounds. Since birds and butterflies are there for the nectar and not the pollen, it would make sense that the pollen of these plant species wouldn’t need to carry a scent.

bee collecting pollen (image credit: wikimedia commons)

In flowers that are wind-pollinated, the pollenkitt layer is either very thin or absent altogether. In this case, pollen grains need to be easily released from the anther and are better off when they aren’t sticking to other pollen grains. That way, they are free to be carried off in the breeze to nearby flowers. Some plant species are amphiphilous, meaning they can be both animal-pollinated and wind-pollinated, and according to the authors of the paper published in Flora (2005), pollenkitt layers in these species exhibit intermediate characteristics of both types of pollen grains, generally with thinner, less-sticky pollenkitt and more pollenkitt found within the cavities of the exine.

It’s clear that this unique pollen-glueing substance plays a critical role in the pollination process for many plant species. Considering that each species of plant has its own story to tell, there is still more to learn about the forms and functions that pollenkitt takes.

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This is the first in a series of posts in 2024 in which, once again, I am exploring the world of pollinators and pollination. You can read more about this effort in last month’s Year in Review post.

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Winter Trees and Shrubs: Sweetgum

Sweetgum is a distinctive tree at any time of year. It stands out among most other deciduous trees with its excurrent growth habit, which gives it a narrowly conical or pyramidal shape. Only in its advanced age does it begin to develop a more rounded and irregular form. Its leaves are star-shaped with either five or seven lobes and span between four and seven inches. Their glossy green color gives way to a wide range of colors in the fall, including yellow, orange, red, and maroon, remaining on the tree for several weeks depending on weather conditions. The fruits are particularly distinct, especially in the winter after the leaves have dropped. Woody balls made up of a series of capsules with pointed beaks hang from long stalks throughout the winter, windborne seeds having been released from small openings in the fall. Without even bothering to look at the twigs, seeing these durable, spiky balls hanging from branches (or feeling them under your feet) is a quick indication that you are looking at a sweetgum.

one of many fall colors found on sweetgum (Liquidambar styraciflua)

Liquidamabar styraciflua (one of the funnest botanical names to say) was previously placed in the family Hamamelidaceae but is now one of the few members of Altingiaceae. Its natural distribution is broad, covering a large portion of the eastern United States and west into Missouri then down into Texas, Mexico, and much of Central America. Outside of its natural range it has been widely planted as an ornamental, and there are several popular sweetgum cultivars currently in cultivation. Both the common and botanical names for this tree refer to the resin found in its bark, which historically has had many uses.

The winter twigs of sweetgum are stout, round, smooth, and yellow-brown to green or olive-green to brown-purple in color. They can also be glossy and feature a few scattered lenticels. Older twigs (or branchlets) are brown at first and then light grey with dark grey lenticels. They are occasionally adorned with corky wings similar to those of bur oak. Because the wings aren’t always present, it can be a fun thing to encounter when you are out looking at twigs.

corky wings on the branchlets of sweetgum

The winter buds of sweetgum twigs are egg-shaped and made up of bud scales with acute tips and ciliate margins. They are green to orange-brown or reddish in color and occasionally sticky. Lateral buds are alternately arranged, are much smaller than terminal buds, and are stalkless and flattened against the twig. They sit above a slightly raised leaf scar that is half-elliptical to triangular in shape and has three distinct vascular bundle traces. The pith of sweetgum twigs is solid, continuous, brownish, and irregularly shaped.

winter twig of sweetgum tree

Sweetgum bark is light to dark grey and is made up of a series of rough, vertically arranged, scaly ridges that become deeply furrowed with age. The mace-like fruits of sweetgum are about one inch wide and, at a glance, are similar in appearance to the seed balls of sycamore trees; however, sycamore seed balls easily break apart when compressed, while the pointed, woody capsules that make up a sweetgum ball are held firmly together and can hold their shape for long periods of time. When these “gumballs” collect on the ground below, they can become a hazard, especially where there is lots of foot traffic. Speaking from experience, they are also obnoxious when operating a mower. This polarizing feature has resulted in bad opinions of the sweetgum tree. Luckily, some people are out there defending it.

sweetgum bark
the persistent fruits of sweetgum

More Winter Trees and Shrubs:

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Tea Time: Fireweed

lf you’ve seen one fireweed, you’ve probably seen several. As an early successional species, growing in large numbers across a vast amount of space is kind of its thing. Any disturbance that leaves bare ground in its wake, such as a wildfire or a windstorm, gives fireweed the opportunity to colonize. It grows quickly and spreads via rhizomes, producing thousands of airborne seeds in the process, sending them off to continue colonization or contribute to soil seed banks in preparation for future disturbances. The role of plants like fireweed is vital – promptly covering bare ground to stave off erosion and acting as a nurse plant to new saplings destined to become the future forest. In a garden setting or in locations where aggressively spreading plants are discouraged, fireweed and its weedy behavior may be unwelcome, but in other contexts, its services are essential.

fireweed (Chamerion angustifolium)

Fireweed (Chamerion angustifolium) is a species in Onagraceae, commonly known as the evening primrose family. It has an impressive distribution, widespread across much of North America and Eurasia. This is owed largely to its adaptability. Deep shade and overly dry soil are two conditions that it does not tolerate well, otherwise it seems to grow in a wide variety of soil types, moisture levels, and sun exposures, particularly in areas where there is regular disturbance. Swaths of towering plants topped with rose-pink flower spikes make fireweed impossible to ignore and a favorite of wildflower enthusiasts.

Fireweed stems reach from three to nine feet tall and are rarely branched. Long, narrow, lance-shaped leaves are arranged alternately along the lengths of stems and give the plant a willow-like appearance, which explains another common name, rosebay willowherb. The undersides of leaves have a distinct venation pattern, in which the veins don’t reach the leaf margins, a feature that can help with identification.

distinct leaf veins of fireweed (Chamerion angustifolium)

A series of rose-pink to purple flowers top the stems of fireweed. Each flower has four sepals and four petals with eight stamens and a four-lobed stigma extending prominently from the center. Its long, narrow ovary can be confused for a flower stalk. Rich with nectar, fireweed flowers are a favorite of honeybee keepers. They are also edible, like much of the rest of the plant. Narrow, four-chambered capsules form in place of fertilized flowers and later split open to release abundant, small seeds with a tuft of fluff attached to each one to aid in wind dispersal.

Fireweed has a long history of being used as food and medicine. Stem fibers are also useful for making cord, and seed fluff is useful for weaving and padding. Certainly, fireweed’s abundance and ubiquity contribute to its utility. Having never eaten fireweed before, I decided that a good way to introduce it to my diet would be to make a tea. Fireweed leaves are commonly collected for tea and are said to make an excellent non-caffeinated replacement for black tea.

fireweed tea leaves

Making fireweed tea starts by stripping young leaves from fireweed stems. Recipes I encountered all called for fermenting the leaves before drying them. I did this by squeezing handfuls of leaves in my fists just enough to break and bruise them a bit and then packing them into a quart size Mason jar. I closed the lid tight and kept them in the jar for about five days, shaking it up a couple times a day supposedly to help prevent mold issues. After that, I dried the leaves on a baking sheet in the hot sun. From there, they are ready for making tea the same way you would make any other loose leaf tea, chopping the leaves up a bit before immersing them in hot water.

I found the taste of fireweed tea to be mild and pleasant. Despite several sources comparing it to black tea, I thought it was more similar to green tea. Sierra liked the smell more than the taste and wished it had honey in it. Compared to other teas I’ve tried in this short series of posts, this is definitely one of the better ones, and a tea I could see myself making again sometime.

More Tea Time Posts on Awkward Botany: