The Wonderful World of Plantlets, Bulbils, Cormlets, Tubercles, and Gemmae

Probably the most well known strategy that plants have for dispersal is by way of seeds. Seeds are plants in embryo, and new generations of plants are born when seeds, released from their parent plants, find suitable locations to germinate. But one of the most amazing things about plants in general is that they have the ability to reproduce in a variety of different ways, and many plant species are not limited to seeds as their only means of dispersal. A paper by Scott Zona and Cody Coyotee Howard, published in Flora (February 2022), introduces us to the intriguing world of aerial vegetative diaspores – just one of the many ways that plants have to get around.

A diaspore is a plant structure that facilitates dispersal. Seeds are diaspores, as are spores, which are produced by non-seed bearing plants like mosses and ferns. If you’ve ever planted bulbs, you’ve handled another type of diaspore. Bulbs and corms, which many spring flowering plants are grown from, form little offshoots called bulblets and cormels that, when detached from their parent structure, can grow into new individuals. These vegetative diaspores are produced below ground. Aerial vegetative diaspores, on the other hand, are formed on above ground plant parts. This clunky term encompasses a number of different structures that are often simply called bulbils, which Zona and Howard explain is used as “a catch-all term that obscures their morphological identity.”

Compiling a list of plant species that feature aerial vegetative diaspores is a difficult task when plant descriptions from various sources use a broad selection of terminology for the same or similar plant parts. To help complete this task, Zona and Howard defined five distinct types of aerial vegetative diaspores – plantlets, bulbils, cormlets, tubercles, and gemmae – and came up with a list of 252 taxa that are known to feature at least one of these structures.

plantlets on the leaf margin of Kalanchoe daigremontiana (wikimedia commons; Aurélien Mora)

Plantlets are miniature plants attached to another plant. Once mature, they have clearly visible leaves, stems, and roots (or root initials) and are non-dormant, meaning they are ready to grow on their own as soon as they’re given the opportunity. The tiny plants borne along the margins of the leaves of mother of thousands (Kalanchoe daigremontiana) is a great example of a plantlet.

A bulbil consists of a shortened stem surrounded by scale leaves modified for food and water storage. Sometimes root initials are visible at the base of the bulbil. Bulbils remain dormant until they are dispersed and conditions are suitable for growth. When bulbils start growing but remain attached to the plant, they become a plantlet. A good example of a bulbil can be found on bulbous bluegrass (Poa bulbosa).

Cormlets are comprised of stem tissue and, like plantlets and bulbils, have a single axis of polarity. They have highly reduced scale leaves and are dormant at dispersal. Bulbil bugle lily (Watsonia meriana), despite its misleading common name, is a good example of a plant that produces cormlets.

Tubercles are made up of swollen stem tissue and, like tubers (their underground counterparts), have multiple shoot buds and multiple axes of polarity (meaning there is no right side up like there is in plantlets, bulbils, and cormlets). They lack scale leaves and are dormant at dispersal. Air potato (Dioscorea bulbifera) is an example of a tubercle-producing plant. As you might guess from the common name, potato-like structures are produced aerially on this vining plant that was introduced to North America from Africa and is now invasive in Florida.

A gemma is a tiny cluster of undifferentiated cells. Gemmae are non-dormant and lack polarity. They are the smallest and least common form of aerial vegetative diaspore and can be found on Drosera pygmaea, a species of sundew native to parts of Australia and New Zealand.

Drosera pygmaea (wikimedia commons; Björn S…)

Zona and Howard’s list of plants with aerial vegetative diaspores is the most comprehensive list to date – although it is undoubtedly and understandably missing some – and includes representatives from 42 plant families and 21 plant orders. Plantlets are the most common form of aerial vegetative diaspore at 116 taxa, with bulbils coming in second at 72. Cormlets and tubercles are less common, with 25 and 16 taxa respectively. Their paper includes the full list and offers further information about many of the species listed. It’s worth taking time to explore and is a valuable resource for anyone interested in the topic. In addition, their discussion section highlights a number of questions that warrant further investigation.

Questions surrounding reproductive strategies and the dispersal of aerial vegetative diaspores are particularly interesting. Because these structures are vegetative, they are essentially clones of the parent plant, meaning there is no genetic mixing as occurs when seeds are produced. This can be an advantage when sexual reproduction isn’t possible due to lack of pollinators, environmental restrictions, or chromosomal/polyploidy anomalies. It also assures that new individuals are pre-adapted to the site, and it can help a species colonize an area quickly. This ability to rapidly colonize explains why several of the species on Kona and Howard’s list are known to be invasive in parts of the world outside of their native range.

A species that produces both seeds and aerial vegetative diaspores may have an advantage when it comes to dispersal since both types of diaspores have their strengths. Seeds can remain dormant in the soil and are likely to persist in the environment longer than vegetative diaspores, but vegetative diaspores can be produced without relying on pollinators and can establish new individuals quickly. The modes of dispersal between the two can also vary. Seeds can be dispersed by wind or carried away by animals, while vegetative diaspores often rely solely on gravity to get around. One exception is hitchhiker elephant ear (Remusatia vivipara), whose bulbils are equipped with tiny hooks that cling to animal fur and are transported in a similar manner to burs.

hooked bulbils of hitchhiker elephant ear (Remusatia vivipara) (wikimedia commons; Dinesh Valke)

When the advantages of aerial vegetative diaspores are considered, it is a wonder that we don’t see them more often. Many plants can be easily propagated by taking stem, leaf, and/or root cuttings and placing them in conditions that favor adventitious root and shoot growth. This may suggest that dormant genetic pathways for producing vegetative diaspores exist in most plants. Or maybe not. Genetic studies of species that feature these structures are needed in order to understand why they are found in some species and not others. Kona and Howard leave us with a slew of research questions like this, and it’s a topic I’ll continue to check in on.

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Burr Tongue, or The Weed That Choked the Dog

It is said that the inspiration for Velcro came when Swiss inventor, George de Mestral, was removing the burrs of burdock from his dog’s coat, an experience we had with Kōura just days after adopting her. I knew that common burdock was found on our property, and I had made a point to remove all the plants that I could easily get to. However, during Kōura’s thorough exploration of our yard, she managed to find the one plant I had yet to pull due to its awkward location behind the chicken coop.

I knew when I saw the clump of burrs attached to her hind end that we were going to spend the evening combing them out of her fur. However, not long after that we discovered that Kōura had already started the process and in doing so had either swallowed or inhaled some. What tipped us off was her violent hacking and gagging as she moved frantically around the living room. She was clearly distraught, and so were we. Recognizing that she had probably swallowed a burr, we made a quick decision to take her to an emergency vet. This was our unfortunately timed (this happened on Christmas Eve) introduction to burr tongue and all the frightening things that can happen when a dog swallows burdock burrs.

The roots, shoots, and leaves of both greater burdock (Arctium lappa) and common burdock (Arctium minus) are edible, which I have already discussed in an Eating Weeds post. The burrs, on the other hand, are clearly not. While sticking to the fur of animals and the clothing of people is an excellent way for a plant to get their seeds dispersed, the sharp, hooked barbs that facilitate this are not something you want down your throat. When this occurs, the natural response is to try to hack them up, which Kōura was doing. Salivating heavily and vomiting can also help. In many cases, this will be enough to eliminate the barbs. However, if they manage to work their way into the soft tissues of the mouth, tongue, tonsils, or throat and remain there, serious infection can occur.

burr of common burdock (Arctium minus)

A paper published in The Canadian Veterinary Journal in 1973 describes the treatment for what is commonly known as burr tongue and technically referred to as granular stomatitis. The paper gives an account of what can happen when “long-haired breeds of dogs … run free in areas where [burdock] grows” and the hooked scales of the burrs consequently “penetrate the mucous membrane of the mouth and tongue.” Dogs with burrs imbedded in their mouths may start eating less or more slowly, drinking more water, and drooling excessively. As infection progresses, their breath can start to stink. A look inside the mouth and at the tongue will reveal lesions where the burrs have embedded themselves. Treatment involves putting the dog under anesthesia, scraping away the infected tissue, and administering antibiotics. Depending on the severity of the lesions, scar tissue can form where the barbs were attached.

To prevent infection from happening in the first place, a veterinarian can put the dog under anesthesia and use a camera inside the dog’s mouth and throat to search for pieces of the burr that may have gotten lodged. There is no guarantee that they will find them all or be able to remove them, and so the dog should be monitored over the next several days for signs and symptoms. At our veterinary visit, the vet also warned us that if any burrs were inhaled into the lungs, they could cause a lung infection, which is another thing to monitor for since it would be practically impossible for an x-ray or a camera to initially find them.

Luckily, now more than three weeks later, Kōura appears to be doing fine, and the offending burdock has been taken care of. One thing is for sure, as someone who is generally forgiving of weeds, burdock is one weed that will not be permitted to grow at Awkward Botany Headquarters.

For more adventures involving Kōura, be sure to follow her on Instagram @plantdoctordog.

The Serotinous Cones of Lodgepole Pine

Behind the scales of a pine cone lie the seeds that promise future generations of pine trees. Even though the seeds are not housed within fruits as they are in angiosperms (i.e. flowering plants), the tough scales of pine cones help protect the developing seeds and keep them secure until the time comes for dispersal. In some species, scales open on their own as the cone matures, at which point winged seeds fall from the tree, taking flight towards their new homes. In other species, the scales must be pried open by an animal in order to free the seed. A third group of species have what are called serotinous cones, the scales of which are sealed shut with resin. High temperatures are required to soften the resin and expose the seeds.

Serotinous cones are a common trait of pine species located in regions where wildfire naturally and regularly occurs. One such species is lodgepole pine (Pinus contorta), which is found in abundance in forests across much of western North America. Lodgepole pine is a thin-barked tree species that burns easily and is often one of the first plants to recolonize after a stand-replacing wildfire. There are 3 or 4 subspecies of lodgepole pine. The one with the largest distribution and the one that most commonly exhibits serotinous cones is P. contorta subsp. latifolia, which occurs throughout the Rocky Mountains, north into the Yukon, and just west of the Cascade Range.

needles of lodgpole pine (Pinus contorta)

Lodgepole pine grows tall and straight, generally maxing out at around 80 feet tall. Its needles are about two and a half inches long, are borne in bundles of two, and tend to twist away from each other, which is one explanation for the specific epithet, contorta. Its cones are egg-shaped with asymmetrical bases, measuring less than two inches long with prickly tips at the ends of each scale. The seeds of lodgepole pine are tiny with little, papery wings that aid in dispersal. The cones can remain attached to the tree for 15-20 years (sometimes much longer), and the seeds remain viable for decades. In non-serotinous cones, the scales start opening on their own in early autumn. Serotinous cones require temperatures of 45-50°C (113-122°F), to release the resin bond between the scales. Some cones that happen to fall from the tree can open when exposed to particularly warm temperatures on the ground. Otherwise, it takes fire to free the seeds.

Serotinous cones aren’t a guarantee, and the percentage of trees with serotinous cones compared to those with non-serotinous cones varies widely across the range of lodgepole pine, both in space and in time. One reason for this is that trees with serotinous cones don’t develop them until they reach a certain age, generally around 20-30 years old, or perhaps as old as 50 or 60. The cones of young trees are all non-serotinous. But some trees never develop serotinous cones at all. Serotiny is a genetic trait, and there are various factors that either select for or against it. A number of factors are at play simultaneously over the life of a tree and across a population of trees, so it is difficult to determine exactly why the percentage of serotinous cones is so variable across the range of the species. What follows are a few potential explanations for this phenomenon.

closed cone of lodgepole pine (Pinus contorta)

As a fire-adapted, pioneer species, lodgepole pine has evolved to live in environments where fire is predictably common. Serotinous cones help ensure that a population won’t be wiped out when a massive wildfire comes through. After the fire has passed and the seeds are released, lodgepole pine can quickly repopulate the barren ground. As long as fire occurs within the lifespan of a population of similarly aged trees, it is advantageous for the majority of individuals to maintain their serotinous trait. If the population is located in an area that historically does not see much fire, serotinous cones may be a disadvantage and can have adverse effects on the longevity of that population.

A study published in Ecology in 2003 looked at the influence that the frequency of fire has on lodgepole pine stands found at low and high elevations in Yellowstone National Park. At lower elevations, where summer temperatures are warmer and precipitation is relatively minimal, fires occur more frequently compared to higher elevations, which tend to be cooler and wetter. The researchers found that at lower elevations when fires occurred at short intervals (less than 100 years between each fire), lodgepole pine was slower to repopulate compared to longer intervals. This suggests that the percentage of serotiny found in stands that experienced short fire intervals was low, and that stands with long fire intervals exhibit a higher percentage of serotiny. After all, as mentioned above, lodgepole pines don’t start developing serotinous cones until later in life.

At higher elevations, where fire occurs less frequently, lodgepole pines were found to have a low percentage of serotinous cones regardless of the age of the stand. Because the trees at high elevations are more likely to die of old age rather than fire, maintaining serotinous cones would be a disadvantage. Open cones are preferred. Thus, at least in this study, a greater percentage of serotinous cones was found in lodgepole pines at lower elevations compared to those at higher elevations. Latitude, elevation, mountain pine beetle attacks, and other environmental factors have all been used to explain differences in serotiny. However, the factor that seems to have the greatest influence is the frequency of fire. As James Lotan writes in a 1976 report: “A high degree of cone serotiny would be expected where repeated, high-intensity fires occur. Where forest canopies are disrupted by factors other than fire, open cones annually supply [seed] for restocking disturbances such as windfalls.”

That being said, one other factor does appear to play a critical role in whether or not lodgepole pines produce serotinous cones, and that is seed predation by squirrels. In a paper published in Ecology in 2004, researchers wondered why the percentage of serotinous cones wasn’t even higher in populations where fire reliably occurred during the lifetime of the stand. To help answer this question they looked at the activities of pine squirrels, which are the main seed predator of lodgepole pine seeds. Pine squirrels visit the canopy of lodgepole pines and consume the seeds found in serotinous cones. Because non-serotinous cones quickly shed their seeds, serotinous cones are a more reliable and accessible food source, and because pine squirrels are so effective at harvesting the seeds of serotinous cones, the researchers concluded that, “in the presence of pine squirrels, the frequency of serotiny is lower and more variable, presumably reflecting,” among a variety of other factors, “the strength of selection exerted by pine squirrels.”

A study published in PNAS in 2014 added evidence to this conclusion. While acknowledging that fire plays a major role in the frequency of serotinous cones, the researchers asserted that “squirrels select against serotiny and that the strength of selection increases with increasing squirrel density.” However, despite making it easier for squirrels to access their seeds, lodgepole pines maintain a degree of serotinous cones, since clearly their main advantage is retaining a canopy-level seed bank from which seeds are released after a fire and by which a new generation of lodgepole pines is born.

open cones of lodgepole pine (Pinus contorta)

Further Reading and Viewing:

Meet Erigeron linearis

Erigeron is a genus of herbaceous, flowering plants consisting of between 390 and 460 species and is a member of the aster/sunflower family (Asteraceae). Plants in this genus are annuals, biennials, or perennials and are mainly found in temperate regions around the world. At least 163 species occur in the contiguous United States. Erigeron diversity is particularly high in western states; however, each state is home to at least one Erigeron species.

A common name for plants in this genus is fleabane. This name comes from an outdated belief that the plants can be used to repel or poison fleas, flies, gnats, and other tiny insects, a belief for which there is no evidence. In Ancient Greek, the name Erigeron is said to mean something akin to “old man in the early morning,” likely referring to the appearance of the seed heads which look like little tufts of white hair. Some Erigeron species are also commonly referred to as daisies.

desert yellow fleabane (Erigeron linearis)

One species of Erigeron that I would like you to meet is Erigeron linearis. While most of the plants in this genus have flowers that are white, pink, or various shades of purple, E. linearis is a yellow-flowered species, hence the common name, desert yellow fleabane. Another common name for this plant is narrow leaved fleabane, a reference to its linear leaves. E. linearis is a small plant with a prominent taproot that reaches up to 20 centimeters tall and forms a leafy, rounded mat or cushion of whitish or gray-green, alternately arranged leaves. The white appearance is due to numerous, fine, appressed hairs on the leaves and stems. Flower stalks are produced in abundance in late spring through early summer and are mostly leafless. They reach above the mound of leaves and are each topped with at least one flower head, which nods at first, but then straightens out as the flowers open. Each flower head is about 2 centimeters wide and is typical of plants in the sunflower family, with a cluster of deep yellow disc florets in the center, surrounded by ray florets that are lighter in color. Both disc and ray florets are fertile; however, the disc florets have both “male” (stamens) and “female” (pistils) flower parts, while the ray florets have only “female” parts. The involucre, which sits at the base of the flowers, is egg-shaped or hemispheric and made up of a series of tiny, fuzzy bracts called phyllaries.

the flower head of desert yellow fleabane (Erigeron linearis)

The fruit of Erigeron linearis is called a cypsela, an achene-like fruit that is characteristic of plants in the sunflower family. The fruits are miniscule and topped with a pappus composed of short outer bristles and longer, pale, inner bristles. The two types of pappus bristles (or double pappus) must be the reason for the scientific name this species was originally given in 1834, Diplopappus linearis. While the seeds of more than 80% of flowering plant species found in dryland regions exhibit some form of dormancy, a study published in Plant Biology (2019), found that E. linearis is one of the few species with non-dormant seeds. This means that for those of us interested in growing plants native to the Intermountain West, E. linearis is a pretty easy one to grow and is a great addition to water-wise gardens, pollinator gardens, and rock gardens.

Erigeron linearis seedling

Erigeron linearis is distributed across several western states and into Canada. It is found in northern California, eastern Oregon and Washington, southern British Columbia, across Idaho and east into southern Montana, western Wyoming and northwestern Utah. It is found at low to moderate elevations in open, rocky foothills, grasslands, sagebrush steppe, and juniper woodlands. It prefers well-drained soils and full sun. It is one of many interesting plants found on lithosols (also known as orthents), which are shallow, poorly develop soils consisting of partially weathered rock fragments. In the book Sagebrush Country, Ronald Taylor calls lithosols “the rock gardens of the sagebrush steppe,” and refers to E. linearis and other members of its genus as “some of the more colorful components of the lithosol gardens.” E. linearis is a food source for pronghorn, mule deer, and greater sage-grouse, and the flowers are visited by several species of bees and butterflies. The plant is also a larval host for sagebrush checkerspots.

desert yellow fleabane (Erigeron linearis)

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Drought Tolerant Plants: Blue Flax

“Lewis’s prairie flax is a pretty garden ornamental suited to hot, dry sites. Each morning delicate sky blue flowers open on slender arching stems, only to fall off in the afternoon and be replaced by others the next morning. In spite of its fragile appearance, it is quite sturdy and may put out a second flush of blossoms on new growth in late summer.”Common to the This Country: Botanical Discoveries of Lewis and Clark by Susan H. Munger

When selecting plants for a waterwise garden, it is imperative that at least a portion of the plants are easy to grow and maintain and are adapted to a wide variety of conditions. This will ensure a more successful garden, both functionally and aesthetically. Luckily, there are a number of drought-tolerant plants that pretty much anyone can grow without too much trouble. Blue flax, in my opinion, is one such plant.

You may be familiar with flax as a culinary plant, known for its edible seeds which are used to make flour (i.e. meal) and oil. Or perhaps you’ve used linseed oil, a product of flax seeds, to protect wooden, outdoor furniture or in other wood finishing projects. You may also think of linen when you think of flax; and you should, because linen is a textile made from the fibrous stems of the flax plant. All of these products generally come from a domesticated, annual flax known as Linum usitatissimum – a species that has been of benefit to humans for millenia. Various species of flax have also been planted for erosion control, fire breaks, forage for livestock, and in pollinator-friendly gardens. Flax seeds, a common ingredient in bird seed mixes, provide food for birds and other small animals. All this to say, humans and flax share a long history together, and it deserves a place in your garden.

The flax species profiled here is actually two species: Linum lewisii and Linum perenne. That’s because these two species look nearly identical and are both used as garden ornamentals and in wildflower seed mixes. They are also both known as blue flax, among myriad other common names. Due to their similiarity, L. lewisii is considered by some to be a subspecies of L. perenne.

Linum lewisii is found across western North America and received its name after being collected by a member of the Lewis and Clark Expedition. The plant collection was brought back from the expedition and determined to be new to western science. It was described and named by Frederick Pursh. Linum perenne is a European species which was introduced to North America as an ornamental and has since become widely naturalized. In 1980, a naturalized selection of L. perenne was released for use in restoration plantings under the cultivar name ‘Appar’ with the understanding that it was L. lewisii. A genetic study later revealed that the cultivar was instead L. perenne. The study also provided evidence that “North American Lewis flax and European perennial blue flax are reproductively isolated,” suggesting that they are indeed two separate species.

Despite being separate species, telling them apart can be a challenge. Blue flax plants grow from a taproot and woody base and are multistemmed, reaching two to three feet tall. The stems are thin yet stringy, wiry, and not easily torn, which helps explain why flax is such a good plant for making textiles. Short, slender leaves are alternately arranged along the length of the stems, while flower buds form at the ends of stems in loose clusters. Flowers bloom early in the day and are spent by the afternoon. They are 5-petaled, saucer-shaped, and a shade of blue – from whitish blue to deep blue – depending on the plant. Small, round, 10-chambered seed capsules form in the place of flowers, each chamber housing one or two flat, shiny, dark brown seeds. Flowers bloom daily in succession up towards the ends of stems even as the fruits of spent flowers lower on the stalk mature.

seed capsules of blue flax

A close look at their flower parts is really the only way you might be able to tell these two species apart. Blue flax flowers have five stamens topped with white anthers and five styles topped with little, yellow stigmas. The flowers of L. lewisii are homostlyous, which means their styles are all the same length and are generally taller than or about the same height as the stamens. The flowers of L. perenne are heterostylous, which means their flowers can either have styles that are much longer than their stamens or stamens that are much longer than their styles. Each plant in a population of L. perenne has either all long-styled flowers or all short-styled flowers. In a mixed population of L. perenne and L. lewisii, separating the long-styled L. perenne plants from the L. lewisii plants presents a challenge (at least for me).

long-styled blue flax flower
short-styled blue flax flower

Due to the similarity of these two species, it’s easy to see how the plants or seeds of blue flax could easily be mislabeled and sold as one species even though they are the other species. This could be a problem in a restoration planting where seed source and identity is critical, but in your garden, it’s really no big deal. Both species are great for waterwise and pollinator gardens. They are equally beautiful and easy to grow and care for. If nothing else, perhaps the challenge in identifying them will encourage you to take a closer look at your flowers and familiarize yourself with their tinier parts – an act all of us amateur botanists could stand to do more often.

The photos in this post were taken at Idaho Botanical Garden in Boise, Idaho.

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To Fruit or Not to Fruit – The Story of Mast Seeding

Perennial plants that are able to reproduce multiple times during their lifetime don’t always yield the same amount of seeds each time they reproduce. For some of these plants, there is a stark difference between high-yield years and low-yield years, with low-yield years outnumbering the occasional high-yield years. In years when yields are high, fruit production can seem excessive. This phenomenon is called masting, or mast seeding, and it takes place at the population level. That is, during a mast year, virtually all individuals in a population of a certain species synchronously produce a bumper crop of seeds.

Plants of many types can be masting species. Bitterroot milkvetch (Astragalus scaphoides) and a tussock grass known as Chionochloa pallens are masting species, for example. However, this behavior is most commonly observed in trees, notably nut producing trees like oaks, beeches, and pecans. As you might imagine, the boom and bust cycles of mast seeding plant populations can have dramatic ecological effects. Animals that eat acorns, for example, are greeted with a veritable buffet in a mast year, which can increase their rate of reproduction for a spell. Then, in years when acorns are scarce, the populations of those animals can plummet.

How and why masting happens is not well understood. It is particularly baffling because masting populations can cover considerably large geographic areas. How do trees covering several square miles all “know” that this is the year to really go for it? While a number of possible explanations have been explored, there is still much to learn, especially since so many different species growing in such varied environments exhibit this behavior.

A popular explanation for mast seeding is predator satiation. The fruits and seeds of plants are important food sources for many animals. When a population of plants produces fruit in an unusually high abundance, its predators won’t possibly be able to eat them all. At least a few seeds will be left behind and can sprout and grow into new plants. By satiating their predators they help ensure the survival of future generations. However, even if a plant species has evolved to behave this way, it still doesn’t explain how all the plants in a particular population seem to know when it’s time for another mast year.

Predator satiation is an example of an economy of scale, which essentially means that individual plants benefit when the population acts as a whole. Another economy of scale that helps explain masting is pollen coupling. This has to do with the timing of flowering in cross pollinating species. If individuals flower out of sync with one another, the opportunities for cross pollination are limited. However, if individuals in a population flower simultaneously, more flowers will be pollinated which leads to increased fruit and seed production.  For this to happen, there are at least two factors that come into play. First, the plants have to have enough resources to flower. Making flowers is expensive, and if the resources to do so (like carbon, nitrogen, and water) aren’t available, it won’t happen. Second, weather conditions have to work in their favor. Timing of flowering depends, not only on daylength, but on temperature, rainfall, and other local weather conditions. If individuals across a population aren’t experiencing similar weather, the timing of their flowering may be off.

pollen-producing (male) flowers of pecan (Carya illinoinensis) — via wikimedia commons; Clemson University

Resource matching and resource budgeting are other proposed explanations for masting. Since plants can only use the resources available to them for things like growth and reproduction, they vary each year in how much growing or reproducing they do. Theoretically, if plants in a population are all going to flower in the same year, they all have to have access to a similar amount of resources. Often, the year following a mast year, there is a significant drop in fruit production, as though the plants have used up all of their available resources for reproduction and are taking a break. Some hypothesize that masting is a result of resource storage, and that plants save up resources for several years until they have what they need for yet another big year.

Another thing to consider is how plant hormones might play a role in masting. Gene expression and environmental cues both result in hormonal responses in plants. As Bogdziewicz, et al. write in Ecology Letters (2020), “if hormones and the genes that control them are hypersensitive to an environmental signal, masting can be at least partially independent of resource- and pollen-based mechanisms.” This and other potential explanations for masting are, at this point, largely theoretical. In their paper, Bogdziewicz, et al. propose a number of ways that theoretical predictions can be experimentally tested. If the “research agenda” outlined in their paper is carried out, they believe it will “take the biology of masting from a largely observational field of ecology to one rooted in mechanistic understanding.”

In her book, Braiding Sweetgrass, Robin Wall Kimmerer proposes an additional explanation for the mechanisms behind masting – the trees are talking to one another. Not in the way that you and I might converse, but rather by sending signals through the air via pheromones and underground via complex fungal networks. There is already evidence for this behavior when it comes to plants defending themselves from predators and in sharing resources, so why not in planning when to reproduce? As Kimmerer writes regarding masting, “the trees act not as individuals, but somehow as a collective.” The question now is how.

seedlings of European beech (Fagus sylvatica), a mast-seeding species — via wikimedia commons; user: Beentree

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Winter Trees and Shrubs: Northern Catalpa

The names of plants often contain clues that can either help with identification or that tell something about the plant’s history or use. The name, catalpa, is said to be derived from the Muscogee word, katałpa, meaning “winged head,” presumably referring to the tree’s winged seeds. Or maybe, as one writer speculates, it refers to the large, heart-shaped, floppy leaves that can make it look like the tree is “ready to take flight.” Or perhaps it’s a reference to the fluted, fused petals of the tree’s large, tubular flowers. I suppose it could mean any number of things, but I’m sticking with its seeds, which are packed by the dozens in the tree’s long, slender, bean-like fruits. The seeds are flat, pale brown, and equipped with paper thin, fringed appendages on either side that assist in wind dispersal – wings, in other words.

winged seeds of northern catalpa (Catalpa speciosa)

Catalpa speciosa, or northern catalpa, is a relatively fast growing, short-lived tree native to the Midwest and one of only two species in the genus Catalpa found in the United States. Its distribution prior to the arrival of Europeans appears to have been restricted to a portion of the central Mississippi River valley, extending west into Arkansas, east into Tennessee, and north into Illinois and Indiana. It has since been widely planted outside of its native range, naturalizing in areas across the Midwest and eastern US. Early colonizers planted northern catalpa for use as fence posts, railroad ties, and firewood. Its popularity as an ornamental tree is not what it once was a century ago, but it is still occasionally planted in urban areas as a shade tree. Its messiness – littering the ground below with large leaves, flowers, and seed capsules – and its tendency to spread outside of cultivation into natural areas are reasons why it has fallen out of favor with some people.

The oval to heart-shaped, 8 to 12 inch long leaves with long petioles rotting on the ground below the tree are one sure sign that you’ve encountered a catalpa in the winter time. The leaves are some of the first to fall at the end of the growing season, briefly turning an unmemorable yellow before dropping.

leaf of northern catalpa (Catalpa speciosa) in the winter with soft hairs on the underside still visible

The leaf arrangement on northern catalpa is whorled and sometimes opposite. The twigs are easy to identify due to several unique features. They are stout, round, and grayish brown with prominent lenticels. The leaf scars are large, rounded, and raised up on the twig, looking a bit like little suction cups. They are arranged in whorls of three, with one scar considerably smaller than the other two. A series of bundle traces inside the scar form an ellipse. The leaf buds are tiny compared to the scar and are protected by loose, pointed, brown bud scales. Northern catalpa twigs lack a terminal bud. In the winter, seed capsules or the stalk of an old inflorescence often remain attached to the terminal end of the twig. The pith inside of the twig is thick, white, and solid.

twig of northern catalpa (Catalpa speciosa)

pith inside twig of northern catalpa (Catalpa speciosa)

Another common name for Catalpa speciosa is cigar tree, a name that comes from its up to 18 inch long, cigar-like seed capsules that hang from the otherwise naked tree throughout the winter. The sturdy, cylindrical pod starts out green in the summer and turns dark brown by late fall. Seed pods that haven’t fallen or already split open will dehisce in the spring time, releasing their papery seeds to the wind.

fruits of northern catalpa (Catalpa speciosa) hanging from the tree in the winter

The young bark of northern catalpa is thin and easily damaged. As it matures, it becomes furrowed with either scaly ridges or blocky plates. Mature trees are generally twisted at the base but otherwise grow straight, reaching 30 to 60 feet tall (sometimes taller) with an open-rounded to narrow-oval crown.

maturing bark of northern catalpa (Catalpa speciosa)

Northern catalpa is one of the last trees to leaf out in the spring. In late spring or early summer, 10 inch long clusters of white, tubular flowers are produced at the tips of stems. Before the flowers open, they look a bit like popped popcorn, reminding me of a song from my childhood (which I will reluctantly leave right here). The margins of its trumpet-shaped petals are ruffled and there is yellow, orange, and/or purple spotting or streaking on the inside of the tubes.

flower of northern catalpa (Catalpa speciosa) just before it opens

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Winter Trees and Shrubs: Netleaf Hackberry

Boise, Idaho is frequently referred to as the City of Trees despite being located in a semiarid region of the Intermountain West known as the sagebrush steppe where few trees naturally grow. It earns this moniker partly because the name Boise is derived from the river that runs through it (the Boise River), which was named La Rivere Boisse, or The Wooded River, by early French trappers. Although it flows through a largely treeless landscape, The Wooded River was an apt name on account of the wide expanse of cottonwoods and willows that grew along its banks. The fervent efforts of early colonizers to plant trees in large numbers across their new city also helped Boise earn the title, City of Trees. Today, residents continue the legacy of planting trees, ensuring that the city will remain wooded for decades to come.

As is likely the case for most urban areas, the majority of trees being planted in Boise are not native to the region. After all, very few tree species are. However, apart from the trees that flank the Boise River, there is one tree in particular that naturally occurs in the area. Celtis reticulata, commonly known as netleaf hackberry, can be found scattered across the Boise Foothills amongst shrubs, bunchgrasses, and wildflowers, taking advantage of deep pockets of moisture found in rocky outcrops and draws.

The western edge of netleaf hackberry’s range extends to the northwest of Boise into Washington, west into Oregon, and down into California. The majority of its range is found south of Idaho, across the Southwest and into northern Mexico, then east into the prairie regions of Kansas and Oklahoma. Previously placed in the elm family, it is now considered a member of the family Cannabaceae (along with hemp and hops). It’s a relatively small, broad tree (sometimes a shrub) with a semi-rounded crown. It grows slowly, is long-lived, and generally has a gnarled, hardened, twisted look to it. It’s a tough tree that has clearly been through a lot.

The leaves of Celtis reticulata are rough, leathery, and oval to lance shaped with serrate or entire leaf margins. Their undersides have a distinct net-like pattern that gives the tree its common name. A very small insect called a hackberry psyllid lays its eggs inside the leaf buds of netleaf hackberries in the spring. Its larvae develop inside the leaf, feeding on the sugars produced during photosynthesis, and causing nipple galls to form in the leaves. It’s not uncommon to see a netleaf hackberry with warty-looking galls on just about every leaf. Luckily, the tree doesn’t seem to be bothered by this.

fallen leaves of netleaf hackberry (Celtis reticulata) with nipple galls

The fruit of netleaf hackberry is a pea-sized drupe that hangs at the end of a pedicel that is 1/4 to 1/2 inch long. Its skin is red-orange to purple-brown, and its flesh is thin with a large seed in the center. The fruits, along with a few random leaves, persist on the tree throughout the winter and provide food for dozens of species of birds and a variety of mammals.

persistent fruit of netleaf hackberry (Celtis reticulata)

Celtis reticulata is alternately branched. Its twigs are slender, zig-zagging, and often curved back towards the trunk. They are reddish-brown with several pale lenticels and have sparse, fine, short hairs that are hard to see without a hand lens. The leaf scars are small, half-round, and raised up from the twig. They have three bundle scars that form a triangle. The buds are triangle-shaped with fuzzy bud scales that are slightly lighter in color than the twig. The twigs are topped with a subterminal bud, and the pith (the inner portion of the twig) is either chambered or diaphragmed and difficult to see clearly without a hand lens. 

twigs of netleaf hackberry (Celtis reticulata)

The young bark of netleaf hackberry is generally smooth and grey, developing shallow, orange-tinged furrows as it gets older. Mature bark is warty like its cousin, Celtis occidentals, and develops thick, grey, corky ridges. Due to its slow growth, the bark can be retained long enough that it becomes habitat for extensive lichen colonies.

bark of young netleaf hackberry (Celtis reticulata)

bark of mature netleaf hackberry (Celtis reticulata)

Netleaf hackberry is one of the last trees to leaf out in the spring, presumably preserving as much moisture as possible as it prepares to enter another scorching hot, bone-dry summer typical of the western states. Its flowers open around the same time and are miniscule and without petals. Their oversized mustache-shaped, fuzzy, white stigmas provide some entertainment for those of us who take the time to lean in for a closer look.

spring flowers of netleaf hackberry (Celtis reticulata)

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Photos of netleaf hackberry taken at Idaho Botanical Garden in Boise, Idaho.

Winter Trees and Shrubs: Eastern Redbud

Botanizing doesn’t have to end when the leaves fall off the trees and the ground goes frozen. Plants may stop actively growing during this time, but they are still there. Some die back to the soil level and spend the entire winter underground, leaving behind brown, brittle shells of their former selves. Others, particularly those with woody stems, maintain their form (although many of them leafless) as they bide their time while daylength dips and rises again, bringing with it the promise of warmer weather. Plants that leave us with something to look at during the winter can still be identified. Without foliage or flowers to offer us clues, we rely instead on branches, bark, and buds to identify woody species. In some cases, such features may even be more helpful in determining a certain species than their flowers and foliage ever were. Either way, it’s a fun challenge and one worth accepting if you’re willing to brave the cold, hand lens and field guides in tow.

In this series of posts I’ll be looking closely at woody plants in winter, examining the twigs, buds, bark, and any other features I come across that can help us identify them. Species by species, I will learn the ropes of winter plant identification and then pass my findings along to you. We’ll begin with Cercis canadensis, an understory tree commonly known as eastern redbud.

Eastern redbud is distributed across central and eastern North America, south of southern Michigan and into central Mexico. It is also commonly grown as an ornamental tree outside of its native range, and a number of cultivars have been developed for this purpose. Mature trees reach up to 30 feet and have short trunks with wide, rounded crowns. Its leaves are entire, round or heart-shaped, and turn golden-yellow in the fall. Gathered below the tree in winter, the leaves maintain their shape and are a light orange-brown color.

fallen leaf of eastern redbud (Cercis canadensis)

Eastern redbud is alternately branched with slender, zig-zagging twigs that are dark reddish-brown scattered with several tiny, light-colored lenticels. Older sections of branches are more grey in color. Leaf scars (the marks left on twigs after leaves fall) are a rounded triangle shape and slightly raised with thin ridges along each side. The top edge of the leaf scar is fringed, which I found impossible to see without magnification. Leaf buds are egg-shaped and 2-3 mm in length with wine-red bud scales that are glabrous (smooth) with slightly white, ciliate margins. Descriptions say there are actually two buds – one stalked and one sessile. If the second bud is there, it’s miniscule and obscured by the leaf scar. I haven’t actually been able to see one. Twigs lack a terminal bud or have a tiny subterminal bud that points off to one side. The pith of the twigs is rounded and pale pink. Use sharp pruners or a razor blade to cut the twig in half lengthwise to see it.

twig and buds of eastern redbud (Cercis canadensis)

Bark is helpful in identifying woody plants any time of year, but is especially worth looking at during the winter when branches have gone bare. The bark of young eastern redbud is grey with orange, furrowed streaks running lengthwise along the trunk. In mature trees, the bark is gray, scaly, and peels to reveal reddish-brown below.

bark of young eastern redbud (Cercis canadensis)

bark of mature eastern redbud (Cercis canadensis)

Eastern redbud is in the bean family (Fabaceae) and its flowers and fruits are characteristic of plants in this family. Fruits can persist on the tree throughout the winter and are another way to identify the tree during the off-season. Seed pods are flat, dark red- or orange-brown, and up to 2.5 inches long with four to ten seeds inside. The seeds are flat, round, about 5 millimeters long, and ranging in color from orange-brown to black.

persistent fruits of eastern redbud (Cercis canadensis)

seeds of eastern redbud (Cercis canadensis)

Eastern redbud flowers in early spring before it has leafed out. Clusters of bright pink flowers form on old branches rather than new stems and twigs. Sometimes flowers even burst right out of the main trunk. This unique trait is called cauliflory.

cauliflory on eastern redbud (Cercis canadensis)


Photos of eastern redbud taken at Idaho Botanical Garden in Boise, Idaho.

Seed Shattering Lost – The Story of Foxtail Millet

For a plant to disperse its seeds, it must first let go of them. Sounds obvious, but it is a key step in the dispersal process and an act that is actually coded in a plant’s DNA. As fruits ripen and seeds mature, an abscission layer is formed that separates the seed-bearing fruits from the plant. No longer attached to their parents, seeds are left to their own devices. If all goes well, they will find themselves in a suitable location where they can germinate and grow into a whole new plant, fully equipped to make seed babies of their own.

The releasing of mature seeds is known as shattering, a term most commonly used in reference to grasses and plants with dehiscent seed pods (i.e. fruits that split open when ripe, such as those in the bean and mustard families). In grasses, seeds form along a central stem called a rachis. As the seeds ripen, they separate from the rachis and drop from the plant. In some cases, the rachis is brittle and a section of it breaks off with each seed that falls.

Seed shattering is not a desirable trait when it comes to food crops. It’s easy to see how yields can be poor if seeds disperse before they are harvested. Thus, an essential step in domesticating certain agricultural crops was selecting plants that lacked this particular trait. Instead of dropping mature seeds, such plants hold on to them, making them easy to collect. A simple and naturally occurring mutation in the genes of these plants allowed early farmers to select varieties that were more ideal for agriculture than their wild progenitors.

Genetic studies of agricultural crops have located genes in a number of species that code for seed shattering, confirming that domestication in many cases involved selecting plants with this gene turned off. A recent study, published in Nature Biotechnology (October 2020), took a different route in locating this gene, looking instead at a weedy, wild relative of a crop that was domesticated at least 8000 years ago. Green foxtail (Setaria viridis) is the wild antecedent of foxtail millet (Setaria italica), a crop that, while still commonly grown for food in parts of Asia, is mostly grown for hay, silage, and bird seed in North America. Recently, interest in foxtail millet and other millets (a term used to refer to the grains of several different species of grasses) is on the rise due to the ability of these crops to tolerate drought and heat.

Illustration of three Setaria species from Selected Weeds of the United States (Agriculture Handbook No. 366) published in 1970

Setaria viridis is an abundant, widespread weed adapted to human disturbance. It’s of Eurasian origin but has been present in North America since the early 1800’s and was likely introduced both intentionally and accidentally. It’s an annual grass with prominent, bristly flowerheads that are easily recognizable and the reason for its common name, green foxtail. A handful of other closely related, similar-looking species are also common weeds in North America. Due to useful traits including its short life cycle, small genome, and self-fertility, S. viridis has been used frequently as a model species to carry out a variety of scientific studies. The study referred to above aimed to further enhance the use of green foxtail, particularly when it comes to crop science.

Researchers traveled across the United States collecting nearly 600 samples of green foxtail in order to better understand its genome. They found that the North American population of green foxtail is composed of multiple introductions and that, as the species has followed humans around, it has quickly adapted to diverse climates found across the continent. In examining the genome, they were able to identify the genetic underpinnings for three traits that have importance to agriculture: response to climate, leaf angle (which is used as a predictor of yield in grain crops), and seed shattering.

foxtail millet (Setaria italica) via wikimedia commons

The seed shattering gene – which the researchers named Less Shattering 1 (SvLes1) – was an especially interesting discovery. When compared to the orthologous gene found in foxtail millet, they found that a frameshift mutation had caused a disruption in the gene, turning it off. Using CRISPR (a gene editing tool) they were able to recreate a similar interruption in green foxtail, which resulted in a loss of seed shattering similar to that of foxtail millet. It became clear that selecting plants with this mutation was an essential step in the domestication of this ancient grain.

An excerpt about seed shattering from Fruit from the Sands by Robert N. Spengler III: 

In many of the world’s domesticated grains, especially those from the founder crops of southwest Asia (i.e. wheat and barley), the earliest phenotypical trait of domestication that archaeobotanists look for is a tough rachis, the small stem by which an individual grain or small cluster of grains is attached to the ear. In their wild form, most grains are programmed to detach easily after the grain ripens; however, in domesticated cereals, the grains remain attached to the ear throughout the harvesting process. This change is an inadvertent result of human harvesting with sickles: as people reap their harvest, the grains with a brittle rachis are dropped and those with a tough rachis are collected, stored, and replanted for successive harvests.

Further Reading: