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)

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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:

Book Review: The Gyroscope of Life

Gyroscopes are entertaining toys and incredibly useful tools. They retain their balance and resist changes to their orientation as long as their flywheel is spinning. As the flywheel slows or stops, the gyroscope wobbles out of control and ultimately quits. Considering their design and function, it’s easy to find parallels between gyroscopes and living systems. Consistent energy inputs keep living things alive. Changes can bring imbalance; major disruptions can lead to death. There is a reason we often describe the natural world as a sort of balancing act. It is the work of an ecologist to make sense of this balancing act. The better we understand it, the more equipped we are to protect it and operate responsibly within it.

It is through this lens that David Parrish writes about the biological world in The Gyroscope of Life, a book that Parrish refers to as “a love song to the field of biology.” Parrish has spent much of his life observing and studying the natural world and, as professor emeritus of Crop and Soil Environmental Sciences at Virginia Tech, undoubtedly shared much of what he presents in his book with countless students over the years. The Gyroscope of Life reads like part memoir and part last lecture, and is the work of someone who has an obvious passion for science and nature.

Parrish spends the first few chapters of his book writing mostly about his life and how he came to be a biologist. He acknowledges his privelege – “born male, white, and American in an era where each of those attributes provided me major advantages” –  having essentially been placed on third base from the start, “well down the third base line.” An aspiring zoologist turned botanist, he spent his early years in graduate school studying seeds and seed dormancy. It’s a topic that obviously interests him, as several pages of the book are spent considering what’s going on inside of a seed. “Seeds provide the widest-spread examples of suspended life,”  Parrish says. Are they alive or dead or neither?

Two additional, major life events play a prominent role in the arc of Parrish’s book. One being his break from organized religion and the other his battle with advanced prostate cancer. He grew up in an orthodox Christian home with a very literal understanding of the Bible. His education put him at odds with what he was taught growing up about (among other things) the age of the earth and its creation. Eventually he came to understand that science and religion “exist in separate non-overlapping spheres – the physical and the metaphysical.” He doesn’t necessarily see science and religion as being inherently at odds with each other, but his understanding of science makes it difficult to “find resonance in religion” due to the “cacophony of dissonance” it offers.

In addressing his prostate cancer, Parrish underwent an operation that gave him a newfound perspective on gender. Freed from “testosterone poisoning,” he was able to more fully consider sex and gender from a biological perspective, which he says he had been doing for decades prior to the operation. He spends a good portion of the book “demystifying sex and gender.” One compelling example he offers involves avocado flowers, which actually change gender over time, a phenomenon known as synchronous dichogamy.

avocado flowers (Persea americana) via wikimedia commons

Over the course of its pages, The Gyroscope of Life covers a significant number of topics in the fields of biology and ecology. It’s a relatively short book, but as it careens through such wide-ranging material, it does so in an approachable and suprisingly succint manner. Parrish’s sense of humor, which doesn’t waver despite how bleak the discussion sometimes gets, helps carry the story along and keeps things interesting. Parrish covers evolution (“[Biologists] argue that, if evolution didn’t happen, it should.”), taxonomy (“the name for naming things”) and sytematics, ecological niches (“[humans] are essentially living niche-free and ecosystemless”), domestication, and so much more. The last chapter is spent discussing agroecosystems (“the organisms and abiotic environment that interact in a human-managed agricultural setting”), a topic he spent much of his career studying.

The underlying message of this book, as I see it, is a simultaneous celebration for life on earth and a concern for the direction things are going considering how humans have managed things. Parrish has some admonition for humans in light of how we’ve treated our home planet, but he isn’t too heavy-handed about it. Overall, reading the book felt like sitting in on a lecture given by a friendly and dynamic professor who has obviously given a lot of thought to what he has to say.

Check out the following video to see David Parrish describe the book in his own words.

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The Hidden Flowers of Viola

Violas keep a secret hidden below their foliage. Sometimes they even bury it shallowly in the soil near their roots. I suppose it’s not a secret really, just something out of sight. There isn’t a reason to show it off, after all. Showy flowers are showy for the sole purpose of attracting pollinators. If pollinators are unnecessary, there is no reason for showy flowers, or to even show your flowers at all. That’s the story behind the cleistogamous flowers of violas. They are a secret only because unless you know to look for them, you would have no idea they were there at all.

Cleistogamy means closed marriage, and it describes a self-pollinating flower whose petals remain sealed shut. The opposite of cleistogamy is chasmogamy (open marriage). Most of the flowers we are familiar with are chasmogamous. They open and expose their sex parts in order to allow for cross-pollination (self-pollination can also occur in such flowers). Violas have chasmogamous flowers too. They are the familiar five-petaled flowers raised up on slender stalks above the green foliage. Cross-pollination occurs in these flowers, and seed-bearing fruits are the result. Perhaps as a way to ensure reproduction, violas also produce cleistogamous flowers, buried below their leaves.

an illustration of the cleistogamous flower of Viola sylvatica opened to reveal its sex parts — via wikimedia commons

Flowers are expensive things to make, especially when the goal is to attract pollinators. Colorful petals, nectar, nutritious pollen, and other features that help advertise to potential pollinators all require significant resources. All this effort is worth it when it results in the ample production of viable seeds, but what if it doesn’t? Having a method for self-pollination ensures that reproduction will proceed in the absence of pollinators or in the event that floral visitors don’t get the job done. A downside, of course, is that a seed produced via self-pollination is essentially a clone of the parent plant. There will be no mixing of genes with other individuals. This isn’t necessarily bad, at least in the short term, but it has its downsides. A good strategy is a mixture of both cross- and self-pollination – a strategy that violas employ.

The cleistogamous flowers of violas generally appear in the summer or fall, after the chasmogamous flowers have done their thing. The fruits they form split open when mature and deposit their seeds directly below the parent plant. Some are also carried away by ants and dispersed to new locations. Seeds produced in these hidden flowers are generally superior and more abundant compared to those produced by their showy counterparts. People who find violas to be a troublesome lawn weed – expanding far and wide to the exclusion of turfgrass – have these hidden flowers to blame.

That being said, there is a defense for violas. In the book The Living Landscape by Rick Darke and Doug Tallamy, Tallamy writes: “Plants such as the common blue violet (Viola sororia), long dismissed by gardeners as a weed, can be reconstituted as desirable components of the herbaceous layer when their ecosystem functionality is re-evaluated. Violets are the sole larval food source for fritillary butterflies. Eliminating violets eliminates fritillaries, but finding ways to incorporate violets in garden design supports fritillaries.”

sweet violet (Viola odorata)

In my search for the cleistogamous flowers of viola, I dug up a sweet violet (Viola odorata). I was too late to catch it in bloom, but the product of its flowers – round, purple, fuzzy fruits – were revealed as I uprooted the plant. Some of the fruits were already opening, exposing shiny, light brown seeds with prominent, white elaiosomes, there to tempt ants into aiding in their dispersal. I may have missed getting to see what John Eastman calls “violet’s most important flowers,” but the product of these flowers was certainly worth the effort.

Fruits formed from the cleistogamous flowers of sweet violet (Viola odorata)

Up close and personal with the fruit of a cleistogamous flower

The seeds (elaiosomes included) produced by the cleistogamous flower of sweet violet (Viola odorata)

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Revisiting the Moon Tree

I first learned about Moon Trees in the fall of 2015. One of the trees – a loblolly pine – had been planted at an elementary school just down the street from where I was living at the time. It wasn’t a new thing – it was planted back in 1977, during the period when most other Moon Trees where being planted around the country and the world – but because it wasn’t doing too well, it was in the news. Members of the community, concerned about its long-term survival, were pitching in to help keep it alive. Once I was made aware of it, I also became concerned and decided to go check on it. I even wrote a post about it, which you can read here.

Now that nearly 5 years have passed, I figured I should go check on it again. I hadn’t heard any more news about it, so I assumed it was still hanging in there, but who knows? Maybe not. Since I was going to be on that side of town for Father’s Day, I made plans to stop by. My dad hadn’t seen the tree yet, so he decided to join me.

As we approached Lowell Elementary on our bikes, I was half-expecting the tree to be gone. It was in pretty sad shape when the community stepped in to help it. Braced for this possibility, I anxiously peered down the street as we biked closer. When the tree came into view, I felt relief and announced, “There it is!”

All this time later, it still looks a little rough. The majority of its bark remains largely obscured by crusty, dried up sap, and its canopy isn’t as full as it likely would be if it was a picture of health. But it’s alive and, surprisingly enough, still growing taller, reaching for the moon.

Any loblolly pine would feel out of place in Idaho – it’s a species whose distribution spans the southwest region of the United States, which is starkly different from the northwest – however, this individual in particular is an anomaly. The seed it sprouted from took a journey into space, circled the moon a number of times and then, as a sapling, was planted in Idaho (of all places). Now, over 40 years later, it stands as a symbol of resilience. Something we could all use right now, I’m sure.

This sign was installed shortly after my original Moon Tree post.

Boise, Idaho’s Moon Tree in June 2020

My dad by the Moon Tree in Boise, Idaho

Me by the Moon Tree in Boise, Idaho

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Was a Moon Tree planted near you? Is it still around? Tell us about it in the comment section below.

 

Dispersal by Bulbils – A Bulbous Bluegrass Story

The main way that a plant gets from place to place is in the form of a seed. As seeds, plants have the ability to travel miles from home, especially with the assistance of outside forces like wind, water, and animals. They could also simply drop to the ground at the base of their parent plant and stay there. The possibilities are endless, really.

But what about plants that don’t even bother making seeds? How do they get around? In the case of bulbous bluegrass, miniature bulbs produced in place of flowers function exactly like seeds. They are formed in the same location as seeds, reach maturity and drop from the plant just like seed-bearing fruits, and are then dispersed in the same ways that seeds are. They even experience a period of dormancy similar to seeds, in that they lie in wait for months or years until the right environmental conditions “tell” them to sprout. And so, bulbils are basically seeds, but different.

bulbous bluegrass (Poa bulbosa)

Bulbous bluegrass (Poa bulbosa) is a Eurasian native but is widely distributed outside of its native range having been repeatedly spread around by humans both intentionally and accidentally. It’s a short-lived, perennial grass that can reach up to 2 feet tall but is often considerably shorter. Its leaves are similar to other bluegrasses – narrow, flat or slightly rolled, with boat-shaped tips and membranous ligules – yet the plants are easy to distinguish thanks to their bulbous bases and the bulbils that form in their flower heads. Their bulbous bases are actually true bulbs, and bulbous bluegrass is said to be the only grass species that has this trait. Just like other bulb-producing plants, the production of these basal bulbs is one way that bulbous bluegrass propagates itself.

basal bulbs of bulbous bluegrass

Bulbous bluegrass is also propagated by seeds and bulbils. Seeds form, like any other plant species, in the ovary of a pollinated flower. But sometimes bulbous bluegrass doesn’t make flowers, and instead modifies its flower parts to form bulbils in their place. Bulbils are essentially tiny, immature plants that, once separated from their parent plant, can form roots and grow into a full size plant. The drawback is that, unlike with most seeds, no sexual recombination has occurred, and so bulbils are essentially clones of a single parent.

The bulbils of bulbous bluegrass sit atop the glumes (bracts) of a spikelet, which would otherwise consist of multiple florets. They have dark purple bases and long, slender, grass-like tips. Bulbils are a type of pseudovivipary, in that they are little plantlets attached to a parent plant. True vivipary occurs when a seed germinates inside of a fruit while still attached to its parent.

Like seeds, bulbils are small packets of starch and fat, and so they are sought ought by small mammals and birds as a source of food. Ants and small rodents are said to collect and cache the bulbils, which is one way they get dispersed. Otherwise, the bulbils rely mostly on wind to get around. They then lie dormant for as long as 2 or 3 years, awaiting the ideal time to take root.

bulbils of bulbous bluegrass

Bulbous bluegrass was accidentally brought to North America as a contaminant in alfalfa and clover seed. It was also intentionally planted as early as 1907 and has been evaluated repeatedly by the USDA and other organizations for use as a forage crop or turfgrass. It has been used in restoration to stabilize soils and reduce erosion. Despite numerous trials, it has consistently underperformed mainly due to its short growth cycle and long dormancy period. It is one of the first grasses to green up in the spring, but by the start of summer it has often gone completely dormant, limiting its value as forage and making for a pretty pathetic turfgrass. Otherwise, it’s pretty good at propagating itself and persisting in locations where it hasn’t been invited and is now mostly considered a weed – a noxious one at that according to some states. Due to its preference for dry climates, it is found most commonly in western North America.

In its native range, bulbous bluegrass frequently reproduces sexually. In North America, however, sexual reproduction is rare, and bulbils are the most common method of reproduction. Prolific asexual reproduction suggests that bulbous bluegress populations in North America should have low genetic diversity. Researchers set out to examine this by comparing populations found in Washington, Oregon, and Idaho. Their results, published in Northwest Science (1997), showed a surprising amount of genetic variation within and among populations. They concluded that multiple introductions, some sexual reproduction, and the autopolyploidy nature of the species help explain this high level of diversity.

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Interested in learning more about how plants get around? Check out the first issue of our new zine Dispersal Stories.

Winter Interest in the Lower Boise Foothills

The Boise Foothills, a hilly landscape largely dominated by shrubs and grasses, are a picturesque setting any time of the year. They are particularly beautiful in the spring when a wide array of spring flowering plants are in bloom, and then again in late summer and early fall when a smaller selection of plants flower. But even when there aren’t flowers to see, plants and other features in the Foothills continue to offer interest. Their beauty may be more subtle and not as immediately striking as certain flowers can be, but they catch the eye nonetheless. Appeal can be found in things like gnarled, dead sagebrush branches, lichen covered rocks, and fading seed heads. Because the lower Boise Foothills in particular have endured a long history of plant introductions, an abundance of weeds and invasive plants residing among the natives also provide interest.

This winter has been another mild one. I was hoping for more snow, less rain, and deeper freezes. Mild, wet conditions make exploring the Foothills difficult and ill-advised. Rather than frozen and/or snow covered, the trails are thick with mud. Walking on them in this state is too destructive. Avoiding trails and walking instead on trail side vegetation is even more destructive, and so Foothills hiking is put on hold until the ground freezes or the trails dry out. This means I haven’t gotten into the Foothills as much as I would like. Still, I managed to get a few photos of some of the interesting things the lower Boise Foothills have to offer during the winter. What follows is a selection of those photos.

snow melting on the fruit of an introduced rose (Rosa sp.)

fading seed heads of hoary tansyaster (Machaeranthera canescens)

samaras of box elder (Acer negundo)

snow on seed heads of yarrow (Achillea millefolium)

gall on introduced rose (Rosa sp.)

sunflower seed heads (Helianthus annuus)

sunflower seed head in the snow (Helianthus annuus)

snow falling in the lower Boise Foothills

fading seed heads of salsify (Tragopogon dubius)

lichen on dead box elder log

seed head of curlycup gumweed (Grindelia squarrosa)

lichen and moss on rock in the snow

fruits of poison ivy (Toxicodendron radicans)

See Also: Weeds and Wildflowers of the Boise Foothills (June 2015)

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The first issue of our new zine, Dispersal Stories, is available now. It’s an ode to traveling plants. You can find it in our Etsy Shop