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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What Is a Plant, and Why Should I Care? part three

“If it wasn’t for the plants, and if it wasn’t for the invertebrates, our ancestors’ invasion of land could never have happened. There would have been no food on land. There would have been no ecosystems for them to populate. So really the whole ecosystem that Tiktaalik and its cousins were moving into back in the Devonian was a new ecosystem. … This didn’t exist a hundred million years before – shallow fresh water streams with soils that are stabilized by roots. Why? Because it took plants to do that – to make the [habitats] in the first place. So really plants, and the invertebrates that followed them, made the habitats that allowed our distant relatives to make the transition from life on water to life on land.” – Neil Shubin, author of Your Inner Fish, in an interview with Cara Santa Maria on episode 107 of her podcast, Talk Nerdy To Me

Plants were not the first living beings to colonize land – microorganisms have been terrestrial for what could be as long as 3.5 billion years, and lichens first formed on rocks somewhere between 550 and 635 million years ago – however, following in the footsteps of these other organisms, land plants paved the way for all other forms of terrestrial life as they migrated out of the waters and onto dry land.

The botanical invasion of land was a few billion years in the making and is worth a post of its own. What’s important to note at this point, is that the world was a much different place back then. For one, there was very little free oxygen. Today’s atmosphere is 21% oxygen; the first land plants emerged around 470 million years ago to an atmosphere that was composed of a mere 4% oxygen. Comparatively, the atmosphere back then was very carbon rich. Early plants radiated into numerous forms and spread across the land and, through processes like photosynthesis and carbon sequestration, helped to dramatically increase oxygen levels. A recent study found that early bryophytes played a major role in this process. The authors of this study state, “the progressive oxygenation of the Earth’s atmosphere was pivotal to the evolution of life.”

A recreation of a Cooksonia species - one of many early land plants. (photo credit: wikimedia commons)

A recreation of a Cooksonia species – one of many early land plants (photo credit: wikimedia commons)

The first land plants looked very different compared to the plants we are used to seeing today. Over the next few hundred million years plants developed new features as they adapted to life on land and to ever-changing conditions. Roots provided stability and access to water and nutrients. Vascular tissues helped transport water and nutrients to various plant parts. Woody stems helped plants reach new heights. Seeds offered an alternative means of preserving and disseminating progeny. Flowers – by partnering with animal life – provided a means of producing seeds without having to rely on wind, water, or gravity. And that’s just scratching the surface. Rooted in place and barely moving, if at all, plants appear inanimate and inactive, but it turns out they have a lot going on.

But what is a plant again? In part one and two, we listed three major features all plants have in common – multicellularity, cell walls composed of cellulose, and the ability to photosynthesize – and we discussed how being an autotroph (self-feeder/producer) sets plants apart from heterotrophs (consumers). Joseph Armstrong writes in his book, How the Earth Turned Green, “photosynthetic producers occupy the bottom rung of communities.” In other words, “all modern ecosystems rely upon autotrophic producers to capture energy and form the first step of a food chain because heterotrophs require pre-made organic molecules for energy and raw materials.”

So, why should we care about plants? Because if it wasn’t for them, there wouldn’t be much life on this planet to speak of, including ourselves.

Plants don’t just provide food though. They provide habitat as well. Plus they play major roles in the cycling of many different “nutrients,” including nitrogen, phosphorous, carbon, sulfur, etc. They are also a major feature in the water cycle. It is nearly impossible to list the countless, specific ways in which plants help support life on this planet, and so I offer two examples: moss and dead trees.

The diminutive stature of mosses may give one the impression that they are inconsequential and of little use. Not so. In her book, Gathering Moss, Robin Wall Kimmerer describes how mosses support diverse life forms:

There is a positive feedback loop created between mosses and humidity. The more mosses there are, the greater the humidity. More humidity leads inexorably to more mosses. The continual exhalation of mosses gives the temperate rain forest much of its essential character, from bird song to banana slugs. … Without mosses, there would be fewer insects and stepwise up the food chain, a deficit of thrushes.

Mosses are home to numerous invertebrate species. For many insects, mosses are a place to deposit their eggs and, consequentially, a place for their larvae to mature into adults. Banana slugs traverse the moss feeding on “the many inhabitants of a moss turf, and on the moss itself.” In the process they help to disperse the moss.

Moss is used as a nesting material by various species of birds, as well as squirrels, chipmunks, voles, bears, and other animals. Patches of moss can also function as “nurseries for infant trees.” In some instances, mosses inhibit seed germination, but they can also help protect seeds from drying out or being eaten. Kimmerer writes, “a seed falling on a bed of moss finds itself safely nestled among leafy shoots which can hold water longer than the bare soil and give it a head start on life.”

moss as nurse plant

Virtually all plants, from the tiniest tufts of grass to the tallest, towering trees have similar stories to tell about their interactions with other living things. Some have many more interactions than others, but all are “used” in some way. And even after they die, plants continue to interact with other organisms, as is the case with standing dead trees (a.k.a. snags).

In his book, Welcome to Subirdia, John Marzluff explains that when “hole creators” use dead and dying trees, they benefit a host of “hole users:”

Woodpeckers are natural engineers whose abandoned nest and roost cavities facilitate a great diversity of life, including birds, mammals, invertebrates, and many fungi, moss, and lichens. Without woodpeckers, birds such as chickadees and tits, swallows and martins, bluebirds, some flycatchers, nuthatches, wood ducks, hooded mergansers, and small owls would be homeless.

As plants die, they continue to provide food and habitat to a variety of other organisms. Eventually they are broken down to their most rudimentary components, and their nutrients are taken up and used by “new life.” Marzluff elaborates on this process:

Much of the ecological web exists out of sight – underground and in rotting wood. There, molds, bacteria, fungi, and a world of invertebrates convert the last molecules of sun-derived plant sugar to new life. These organisms are technically ‘decomposers,’ but functionally they are among the greatest of creators. Their bodies and chemical waste products provide us with an essential ecological service: soil, the foundation of terrestrial life.

Around 470 million years ago, plants found their way to land. Since then life of all kinds have made land their home. Plants helped lead the way. Today, plants continue their long tradition of supporting the living, both in life and in death.

Dung Moss (Revisited)

This is a revised version of a post that was originally published on January 14th, 2015. It includes excerpts from a chapter entitled, “Portrait of Splachnum,” in the book, Gathering Moss, by Robin Wall Kimmerer.

Certain plants, like corpse flowers and carrion flowers, emit foul odors when they bloom. The scent is akin to the smell of rotting flesh, hence their common names. The purpose of this repugnant act is to attract a specific group of pollinators: flies, carrion beetles, and other insects that are attracted to gross things. Though this particular strategy is rare, these aren’t the only plants that employ stinky smells to recruit such insects to aid in reproduction and dissemination. Consider dung mosses.

No moss is more fastidious in its choice of habitats than Splachnum. Absent from the usual mossy haunts, Splachnum is found only in bogs. Not among the commoners like Sphagnum that build the peaty hummocks, not along the margins of the blackwater pools. Splachnum ampullaceum occurs in one, and only one, place in the bog. On deer droppings. On white-tailed deer droppings. On white-tailed deer droppings which have lain on the peat for four weeks. In July.

At least three genera (SplachnumTetraplodon, and Tayloria) in the family Splachnaceae include species that go by the common name, dung moss. All Splachnum and Tetraplodon species and many species in the genus Tayloria are entomophilous. Entomophily is a pollination strategy in which pollen or spores are distributed by insects. Compare this to anemophily, or wind pollination, which is the common way that moss spores are distributed. In fact, dung mosses are the only mosses known to exhibit entomophily.

Dung Moss (photo credit: wikimedia commons)

Dung Moss (photo credit: wikimedia commons)

Before we go too much further, it’s important to understand how mosses differ from other plants. Mosses are in a group of non-vascular and non-flowering plants called bryophytes. Vascular tissues are the means by which water and nutrients are transported to and from plant parts. Lacking vascular tissues, water and nutrients are simply absorbed through the leaves and stems of mosses, which is why mosses are typically petite and prefer moist environments. Mosses also lack true roots and instead have rhizoids – threadlike structures that anchor the plants to their substrate of choice (such as dung).

Another major distinction between bryophytes and other plants is that bryophytes spend most of their life cycle as a haploid gametophyte rather than a diploid sporophyte. In most plants, the haploid gametophytes are the sperm (pollen) and egg cells; the sporophyte is everything else. In mosses, the familiar green, leafy structure is actually the gametophyte. The gametophyte houses sperm and egg cells, and when the egg is fertilized by sperm it forms a zygote that develops into the sporophyte structure which extends above the leafy gametophyte. A capsule at the top of the sporophyte contains spores which are eventually released and, upon finding themselves on a suitable substrate in a hospitable environment, germinate to produce new plants. The spore then is comparable to a seed in vascular, seed-bearing plants.

photo credit: wikimedia commons

photo credit: wikimedia commons

As stated earlier, the spores of most mosses are distributed by wind. Dung mosses, on the other hand, employ flies in the distribution of their spores. They attract the flies by emitting scents that only flies can love from an area on the capsule of the sporophyte called the apophysis. This area is often enlarged and brightly colored in yellow, magenta, or red, giving it a flower-like appearance which acts as a visual attractant. The smells emitted vary depending on the type of substrate a particular species of dung moss inhabits. Some dung mosses grow on the dung of herbivores and others on the dung of carnivores. Some even prefer the dung of a particular group of animals; for example, a population of Tetraplodon fuegiensis was found to be restricted to the feces and remains of foxes. However, dung is not the only material that dung mosses call home. Certain species grow on rotting flesh, skeletal remains, or antlers.

Splachnum ampullaceum inhabits the droppings of white-tailed deer. Had a wolf or coyote followed the scent of the deer into the bog, its droppings would been colonized by S. luteum. The chemistry of carnivore dung is sufficiently distinct from that of herbivores to support a different species. … Moose droppings have their own loyal follower. The family to which Splachnum belongs includes several other mosses with an affinity for animal nitrogen. Tetraplodon and Tayloria can be found on humus, but primarily inhabit animal remains such as bones and owl pellets. I once found an elk skull lying beneath a stand of pines, with the jawbone tufted with Tetraplodon.

Yellow Moosedung Moss (Splachnum luteum) has one of the largest and showiest sporophytes. (photo credit:

Yellow moosedung moss (Splachnum luteum) has one of the largest and showiest sporophytes. (photo credit:

The set of circumstances that converge to bring Splachnum into the world is highly improbable. Ripening cranberries draw the doe to the bog. She stands and grazes with ears alert, flirting with the risk of coyotes. Minutes after she has paused, the droppings continue to steam. … The droppings send out an invitation written in wafting molecules of ammonia and butyric acid. Beetles and bees are oblivious to this signal, and go on about their work. But all over the bog, flies give up their meandering flights and antennae quiver in recognition. Flies cluster on the fresh droppings and lap up the salty fluids that are beginning to crystallize on the surface of the pellets. Gravid females probe the dung and insert glistening white eggs down into the warmth. Their bristles leave behind traces from their earlier foraging trips among the day’s dung, delivering spores of Splachnum on their footprints.

The spores of dung mosses are small and sticky. When a fly visits these plants, the spores adhere to its body in clumps. The fly then moves on to its substrate of choice to lay its eggs, and the spores are deposited where they can germinate and grow into new moss plants. Flies that visit dung mosses receive nothing in return for doing so, but instead are simply “tricked” into disseminating the propagules. The story is similar with corpse flowers and carrion flowers; flies are drawn in by the smells and recruited to transmit pollen while receiving no nectar reward for their work.

There are 73 species in the Splachnaceae family, and nearly half of these species are dung mosses. Most are found in temperate habitats in both the northern and southern hemispheres, with a few species occurring in the mountains of subtropical regions. They can be found in both wet and relatively dry habitats. Dung mosses are generally fast growing but short lived, with some lasting only about 2 years. It isn’t entirely clear how and why mosses in this family evolved to become entomophilous, but one major benefit of being this way is that their spores are reliably deposited on suitable habitat.

Since Splachnum can grow only on droppings, and nowhere else, the wind cannot be trusted with dispersal. Escape of the spores is successful only if they have both a means of travel and a reserved ticket for a particular destination. In the monotonous green of the bog, flies are attracted to the cotton candy colors of Splachnum, mistaking them for flowers. Rooting about in the moss for non-existent nectar the flies become coated with the sticky spores. When the scent of fresh deer droppings arrives on the breeze, the flies seek it out and leave Splachnum-coated footprints in the steaming dung.

Sporophytes of Splachnum vasculosum (photo credit:

Sporophytes of Splachnum vasculosum (photo credit:


Koponen, A. 2009. Entomophily in the Splachnaceae. Botanical Journal of the Linnean Society 104: 115-127.

Marino, P., R. Raguso, and B. Goffinet. 2009. The ecology and evolution of fly dispersed dung mosses (Family Splachnaceae): Manipulating insect behavior through odour and visual cues. Symbiosis 47: 61-76.

Ethnobotany: Cattails

“If you ever eat cattails, be sure to cook them well, otherwise the fibers are tough and they take more chewing to get the starchy food from them than they are worth. However, they taste like potatoes after you have been eating them for a couple weeks, and to my way of thinking are extremely good.”  – Sam Gribley in My Side of the Mountain by Jean Craighead George


Illustration by Franz Anthony (

Ask anyone to list plants commonly found in American wetlands, and you can guarantee that cattails will make the list nearly every time. Cattails are widespread throughout the Northern Hemisphere. They are so successful, that it is hard to picture a wetland without them. In her book, Braiding Sweetgrass, Robin Wall Kimmerer discusses this well known association:

Cattails grow in nearly all types of wetlands, wherever there is adequate sun, plentiful nutrients, and soggy ground. Midway between land and water, freshwater marshes are among the most highly productive ecosystems on earth, rivaling the tropical rainforest. People valued the supermarket of the swamp for the cattails, but also as a rich source of fish and game. Fish spawn in the shallows; frogs and salamanders abound. Waterfowl nest here in the safety of the dense sward, and migratory birds seek out cattail marshes for sanctuary on their journeys.

The two most abundant species of cattails in North America are Typha latifolia (common cattail) and Typha angustifolia (narrow leaf cattail). T. angustifolia may have been introduced from Europe. The two species also hybridize to form Typha x glauca. There are about 30 species in the genus Typha, and they share the family Typhaceae with just one other genus. The common names for cattail are nearly as abundant as the plant itself: candlewick, water sausage, corn dog plant, cossack asparagus, reedmace, nailrod, cumbungi, etc., etc.

Cattails have long, upright, blade-like leaves. As they approach the base of the plant, the leaves wrap around each other to form a tight bundle with no apparent stem. As Kimmerer puts it, this arrangement enables the plants to “withstand wind and wave action” because “the collective is strong.” Flowers appear on a tall stalk that reaches up towards the tops of the leaves. The inflorescence is composed of hundreds of separate male and female flowers. Male flowers are produced at the top of the stalk and female flowers are found directly below them. In the spring, the male flowers dump pollen down onto the female flowers, and wind carries excess pollen to nearby plants, producing what looks like yellow smoke.

After pollination, the male flowers fade away, leaving the female flowers to mature into a seed head. Just like the flowers, the seeds are small and held tightly together, maintaining the familiar sausage shape. Each seed has a tuft of “hair” attached to it to aid in wind dispersal. In The Book of Swamp and Bog, John Eastman writes about the abundant seeds (“an estimated average of 220,000 seeds per spike”) of cattail: “A quick experiment, one that Thoreau delighted to perform, demonstrates how tightly the dry seeds are packed in the spike – pull out a small tuft and watch it immediately expand to fill your hand with a downy mass.”

cattails bunch

cattail fluff

Because cattails spread so readily via rhizomes, prolific airborne seeds mostly serve to colonize new sites, away from the thick mass of already established cattails. The ability to dominate vast expanses of shoreline gives cattails an invasive quality that often results in attempts at removal. Various human activities may be aiding their success. Regardless, they provide food and habitat to numerous species of insects, spiders, birds, and mammals. A cattail marsh may not be diverse plant-wise, but it is teeming with all sorts of other life.

Ethnobotanically speaking, it is hard to find many other species that have as many human uses as cattails. For starters, nearly every part of the plant is edible at some point during the year. The rhizomes can be consumed year-round but are best from fall to early spring. They can be roasted, boiled, grated, ground, or dried and milled into flour. Starch collected from pounding and boiling the rhizomes can be used as a thickener. In the spring, young shoots emerging from the rhizomes and the tender core of the leaf bundles can be eaten raw or cooked and taste similar to cucumber. Young flower stalks can be boiled and eaten like corn on the cob and taste similar to artichoke. Pollen, which is high in protein, can be mixed with flour and used to make pancakes and baked goods, among other things. The seeds can be ground into flour or pressed to produce cooking oil.

Cattail leaves can be used to make cords, mats, baskets, thatch, and many other things. Kimmerer writes about the excellent wigwam walls and sleeping mats that weaved cattail leaves make:

The cattails have made a suburb material for shelter in leaves that are long, water-repellent, and packed with closed-cell foam for insulation. … In dry weather, the leaves shrink apart from one another and let the breeze waft between them for ventilation. When the rains come, they swell and close the gap, making the [wall] waterproof. Cattails also make fine sleeping mats. The wax keeps away moisture from the ground and the aerenchyma provide cushioning and insulation.

The fluffy seeds make great tinder for starting fires, as well as excellent insulation and pillow and mattress stuffing. The dry flower stalks can be dipped in fat, lit on fire, and used as a torch. Native Americans used crushed rhizomes as a poultice to treat burns, cuts, sores, etc. A clear gel is found between the tightly bound leaves of cattail. Kimmerer writes, “The cattails make the gel as a defense against microbes and to keep the leaf bases moist when water levels drop.” The gel can be used like aloe vera gel to soothe sunburned skin.

Eastman rattles off a number of commercial uses for cattail: “Flour and cornstarch from rhizomes, ethyl alcohol from the fermented flour, burlap and caulking from rhizome fibers, adhesive from the stems, insulation from the downy spikes, oil from the seeds, rayon from cattail pulp, …” To conclude his section on cattails he writes, “With cattails present, one need not starve, freeze, remain untreated for injury, or want for playthings.”

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