A Few Fun Facts About Pollen

Sexual reproduction in vascular plants requires producing and transporting pollen grains – the male gametophytes or sperm cells of a plant. These reproductive cells must make their way to the egg cells in or order to form seeds – plants in embryo. The movement of pollen is something we can all observe. It’s happening all around us on a regular basis. Any time a seed-bearing plant (also known as a spermatophyte) develops mature cones or flowers, pollen is on the move. Pollen is a ubiquitous and enduring substance and a fascinating subject of study. In case you don’t believe me, here are a few fun facts.

Bee covered in pollen – photo credit: wikimedia commons

Pollen is as diverse as the species that produce it. Pollen grains are measured in micrometers and are so tiny that the only reason we can see them with the naked eye is because they are often found en masse. Yet they are incredibly diverse in size, shape, and texture, and each plant species produces its own unique looking pollen. With the help of a good microscope, plants can even be identified simply by looking at their pollen. See images of the pollen grains of dozens of plant species here and here.

Pollen helps us answer questions about the past. Because pollen grains are so characteristic and because their outer coating (known as exine) is so durable and long-lasting, studying pollen found in sediments and sedimentary rocks helps us discover all sorts of things about deep time. The study of pollen and other particulates is called palynology. Numerous disciplines look to palynology to help them answer questions and solve mysteries. Its even used in forensics to help solve crimes. Criminals should be aware that brushing up against a plant in bloom may provide damning evidence.

Pollen oddities. While all pollen is different, some plants produce particularly unique pollen. The pollen grains of plants in the orchid and milkweed families, for example, are formed into united masses called pollinia. Each pollinium is picked up by pollinators and transferred to the stigmas of flowers as a single unit. A number of other species produce other types of compound pollen grains. The pollen grains of pines and other conifers are winged, and the pollen grains of seagrass species, like Zostera spp., are filamentous and said to hold the record for longest pollen grains.

The pollinia of milkweed (Asclepias spp.) look like the helicopter-esque fruits of maple trees. photo credit: wikimedia commons

Pollen tube oddities. In flowering plants, when pollen grains reach the stigma of a compatible flower, a vegetative cell within the grain forms a tube in order to transport the regenerative cells into the ovule. This tube varies in length depending on the length of the flower’s style. Because corn flowers produce such long styles (also know as corn silk), corn pollen grains hold the record for longest pollen tube, which can measure 12 inches or more. Species found in the mallow, gourd, and bellflower families produce multiple pollen tubes per pollen grain. Hence, their pollen is said to be polysiphonous.

Pollen is transported in myriad ways. Plants have diverse ways of getting their pollen grains where they need to be. Anemophilous plants rely on wind and gravity. They produce large quantities of light-weight pollen grains that are easily dislodged. Most of this pollen won’t make it, but enough of it will to make this strategy worth it. Hydrophilous plants use water and, like wind pollinated plants, may produce lots of pollen due to the unpredictably of this method. Some hydrophilous plants transport their pollen on the surface of the water, while others are completely submerged during pollination.

Employing animals to move pollen is a familiar strategy. Entomophily (insect pollination) is the most common, but there is also ornithophily (bird pollination) and chiropterophily (bat pollination), among others. Plants that rely on animals for pollination generally produce pollen grains that are sticky and nutritious. They attract animals using showy flowers, fragrance, and nectar. The bodies of pollinating insects have modifications that allow them to collect and transport pollen. Certain bees, like honey bees and bumblebees, have pollen baskets on their hind legs, while other bees have modified hairs called scopae on certain parts of their bodies.

Pollen is edible. Some animals – both pollinating and non-pollinating – use pollen as a food source. Animals that eat pollen are palynivores. Bees, of course, eat pollen, but lots of other insects do, too. Even some spiders, which are generally thought of as carnivores, have been observed eating pollen that gets trapped in their webs.

Pollen is thought to be highly nutritious for humans as well, and so, along with being taken as a supplement, it is used in all sorts of food products. To collect pollen, beekeepers install pollen traps on their beehives that strip incoming worker bees of their booty. Pollen from various wind pollinated plants, like cattails and pine trees, are also collected for human consumption. For example, a Korean dessert called dasik is made using pine pollen.

pine pollen – photo credit: wikimedia commons

Pollen makes many people sick. Hay fever is a pretty common condition and is caused by an allergy to wind-borne pollen. This condition is also known as pollinosis or allergic rhinitis. Not all flowering plants are to blame though, so here is a list of some of the main culprits. Because so many people suffer from hay fever, pollen counts are often included in weather reports. Learn more about what those counts mean here.

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When Urban Pollinator Gardens Meet Native Plant Communities

Public concern about the state of bees and other pollinating insects has led to increased interest in pollinator gardens. Planting a pollinator garden is often promoted as an excellent way for the average person to help protect pollinators. And it is! However, as with anything in life, there can be downsides.

In many urban areas, populations of native plants remain on undeveloped or abandoned land, in parks or reserves, or simply as part of the developed landscape. Urban areas may also share borders with natural areas, the edges of which are particularly prone to invasions by non-native plants. Due to human activity and habitat fragmentation, many native plant populations are now threatened. Urban areas are home to the last remaining populations of some of these plants.

Concern for native plant populations in and around urban areas prompted researchers at University of Pittsburgh to review some of the impacts that urban pollinator gardens may have and to develop a “roadmap for research” going forward. Their report was published earlier this year in New Phytologist.

Planting a wildflower seed mix is a simple way to establish a pollinator garden, and such mixes are sold commercially for this purpose. Governmental and non-governmental organizations also issue recommendations for wildflower, pollinator, or meadow seed mixes. With this in mind, the researchers selected 30 seed mixes “targeted for urban settings in the northeastern or mid-Atlantic USA” to determine what species are being recommended for or commonly planted in pollinator gardens in this region. They also developed a “species impact index” to assess “the likelihood a species would impact remnant wild urban plant populations.”

A total of 230 species were represented in the 30 seed mixes. The researchers selected the 45 most common species for evaluation. Most of these species (75%) have generalized pollination systems, suggesting that there is potential for sharing pollinators with remnant native plants. Two-thirds of the species had native ranges that overlapped with the targeted region; however, the remaining one-third originated from Europe or western North America. The native species all had “generalized pollination systems, strong dispersal and colonization ability, and broad environmental tolerances,” all traits that could have “high impacts” either directly or indirectly on remnant native plants. Other species were found to have either high dispersal ability but low chance of survival or low dispersal ability but high chance of survival.

This led the researchers to conclude that “the majority of planted wildflower species have a high potential to interact with native species via pollinators but also have the ability to disperse and survive outside of the garden.” Sharing pollinators is especially likely due to super-generalists like the honeybee, which “utilizes flowers from many habitat types.” Considering this, the researchers outlined “four pollinator-mediated interactions that can affect remnant native plants and their communities,” including how these interactions can be exacerbated when wildflower species escape gardens and invade remnant plant communities.

photo credit: wikimedia commons

The first interaction involves the quantity of pollinator visits. The concern is that native plants may be “outcompeted for pollinators” due to the “dense, high-resource displays” of pollinator gardens. Whether pollinator visits will increase or decrease depends on many things, including the location of the gardens and their proximity to native plant communities. Pollinator sharing between the two has been observed; however, “the consequences of this for effective pollination of natives are not yet understood.”

The second interaction involves the quality of pollinator visits. Because pollinators are shared between native plant communities and pollinator gardens, there is a risk that the pollen from one species will be transferred to another species. High quantities of this “heterospecific pollen” can result in reduced seed production. “Low-quality pollination in terms of heterospecific pollen from wildflower plantings may be especially detrimental for wild remnant species.”

The third interaction involves gene flow between pollinator gardens and native plant communities. Pollen that is transferred from closely related species (or even individuals of the same species but from a different location) can have undesired consequences. In some cases, it can increase genetic variation and help address problems associated with inbreeding depression. In other cases, it can introduce traits that are detrimental to native plant populations, particularly traits that disrupt adaptations that are beneficial to surviving in urban environments, like seed dispersal and flowering time. Whether gene flow between the two groups will be positive or negative is difficult to predict, and “the likelihood of genetic extinction versus genetic rescue will depend on remnant population size, genetic diversity, and degree of urban adaptation relative to the planted wildflowers.”

The fourth interaction involves pathogen transmission via shared pollinators. “Both bacterial and viral pathogens can be transmitted via pollen, and bacterial pathogens can be passed from one pollinator to another.” In this way, pollinators can act as “hubs for pathogen exchange,” which is especially concerning when the diseases being transmitted are ones for which the native plants have not adapted defenses.

photo credit: wikimedia commons

All of these interactions become more direct once wildflowers escape gardens and establish themselves among the native plants. And because the species in wildflower seed mixes are selected for their tolerance of urban conditions, “they may be particularly strong competitors with wild remnant populations,” outcompeting them for space and resources. On the other hand, the authors note that, depending on the species, they may also “provide biotic resistance to more noxious invaders.”

All of these interactions require further investigation. In their conclusion, the authors affirm, “While there is a clear potential for positive effects of urban wildflower plantings on remnant plant biodiversity, there is also a strong likelihood for unintended consequences.” They then suggest future research topics that will help us answer many of these questions. In the meantime, pollinator gardens should not be discouraged, but the plants (and their origins) should be carefully considered. One place to start is with wildflower seed mixes, which can be ‘fine-tuned’ so that they benefit our urban pollinators as well as our remnant native plants. Read more about plant selection for pollinators here.

Summer of Weeds: Willowherbs and Fireweed

Last week we discussed a plant that was introduced as an ornamental and has become a widespread weed. This week we discuss some native plants that have become weedy in places dominated by humans. Similar to pineapple weed, species in the genus Epilobium have moved from natural areas into agricultural fields, garden beds, and other sites that experience regular human disturbance. Some species in this genus have been deliberately introduced for their ornamental value, but others have come in on their own. In all cases the story is similar, humans make room and opportunistic plants take advantage of the space.

Epilobium species number in the dozens and are distributed across the globe. North America is rich with them. They are commonly known as willowherbs and are members of the evening primrose family (Onagraceae). They are herbaceous flowering plants with either annual or perennial life cycles and are commonly found in recently disturbed sites, making them early successional or pioneer species. Many are adapted to wet soils and are common in wetlands and along streambanks; others are adapted to dry, open sites. Hybridization occurs frequently among species in the Epilobium genus, and individual species can be highly variable, which may make identifying them difficult.

northern willowherb (Epilobium ciliatum)

At least two North American species are commonly weedy: E. ciliatum (northern willowherb) and E. brachycarpum (panicled willowherb). Regarding these two species, the IPM website of University of California states: “Willowherbs are native broadleaf plants but usually require a disturbance to establish. Although considered desirable members of natural habitats, they can be weedy in managed urban and agricultural sites.” The field guide, Weeds of the West, refers to E. brachycarpum as a “highly variable species found mostly on non-cultivated sites, and especially on dry soils and open areas.” E. ciliatum is notorious for being a troublesome weed in greenhouses and nurseries, as discussed on this Oregon State University page.

E. ciliatum is a perennial that reproduces via both rhizomes and seeds. It reaches up to five feet tall and has oppositely arranged, lance-shaped leaves with toothed margins that are often directly attached to the stems. Its flowers are tiny – around a quarter of an inch wide – and white, pink, or purple with four petals that are notched at the tip. They sit atop a skinny stalk that is a few centimeters long, which later becomes the fruit. When dry, the fruit (or capsule) splits open at the top to reveal several tiny seeds with tufts of fine hairs.

northern willowherb (Epilobium ciliatum)

E. brachycarpum is an annual that reaches up to three feet tall and is highly branched. Its leaves are short and narrow and mostly alternately arranged. Its flowers and seed pods are similar to E. ciliatum. At first glance it can appear as one of many weeds in the mustard family; however, the tuft of hairs on its seeds distinguishes it as a willowherb.

Seeds and seed pods of panicled willowherb (Epilobium brachycarpum)

Weeds of North America by Richard Dickinson and France Royer describes one weedy species of willowherb that was introduced to North America from Europe – E. hirsutum. It is commonly referred to as great hairy willowherb, but some of its colloquial names are worth mentioning: fiddle grass, codlins and cream, apple-pie, cherry-pie, blood vine, and purple rocket. Introduced as an ornamental in the mid 1800’s, it is a semiaquatic perennial that can reach as tall as eight feet. It has small, rose-purple flowers and is frequently found growing in wetlands along with purple loosestrife (Lythrum salicaria).

Chamerion angustifolium – which is synonymously known as Epilobium angustifolium and commonly called fireweed – is distributed throughout temperate regions of the Northern Hemisphere. It is a rhizomatously spreading perennial that grows to nine feet tall; has lance-shaped, stalkless leaves; and spikes of eye-catching, rose to purple flowers. It is a true pioneer species, found in disturbed sites like clear-cuts, abandoned agricultural fields, avalanche scars, and along roadsides. It gets its common name for its reputation of being one of the first plants to appear after a fire, as John Eastman describes in The Book of Field and Roadside: “A spring fire may result in a profusion of growth as soon as 3 months afterward, testifying to fireweed’s ample seed bank in many wilderness areas.” Eastman goes on to write, “fireweed’s flush of abundance following fire may rapidly diminish after only a year or two of postburn plant growth.” This “flush of abundance” is what gives it its weedy reputation in gardens. With that in mind, it is otherwise a welcome guest thanks to its beauty and its benefit to pollinators.

fireweed (Chamerion angustifolium)

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Quote of the Week:

From the book Food Not Lawns by H.C Flores

Sometimes [weeding] feels like playing God – deciding who lives and who dies is no small matter – and sometimes it feels like war. … Take a moment to ponder the relationship of these plants to other living things around, now and in the future. Your weeds provide forage and habitat for insects, birds, and animals, as well as shelter for the seedlings of other plants. They cover the bare soil and bring moisture and soil life closer to the surface, where they can do their good work. Weeds should be respected for their tenacity, persistence, and versatility and looked upon more as volunteers than as invaders.


Summer of Weeds: Lambsquarters

Since we seem to be on the topic of edible weeds we may as well discuss lambsquarters, another frequently present and commonly eaten, nutritious and versitile weed. Botanically known as Chenopodium album, it is a member of the family Amaranthaceae and therefore related to several common (and uncommon) agricultural crops, including spinach (Spinacia oleracea), beets (Beta vulgaris), Swiss chard (also Beta vulgaris), amaranth (Amaranthus spp.), and red orach (Atriplex hortensis). Chenopodium, a genus consisting of 100 plus species, is also cultivated in various parts of the world for its edible leaves, stems, and seeds. Chenopodium quinoa, commonly known as quinoa, is now a popular “grain” in North America after being grown for millenia by Andean cultures.

Chenopodium album is a summer annual that reaches up to 6 feet tall with sturdy, angular stems and triangular, diamond-shaped, or lance-shaped leaves with irregularly toothed margins. The leaves are green on top and mealy gray-white on bottom. The flowers are tiny, petal-less, and organized in tight clusters at the ends of branches. In Botany In a Day, Thomas Elpel describes the flowers as “little green ‘globs’ forming along an upright stalk, sometimes colored with specks of yellow.” They are generally wind-pollinated, but are occassionally visited by pollinating insects. Each plant can produce tens of thousands of seeds, which are potentially viable for up to 40 years.

Inflorescence of lambsquarters (Chenopodium album)

Lambsquartes is one of many common names for C. album (others include goosefoot, fat hen, baconweed, mealweed, frostblite, and wild spinach), and is a name with several proposed origins. Is it because the plant is commonly found growing in the manure-rich soils of barnyards? Or is it because the fuzzy undersides of the leaves are reminiscent of sheep’s wool? Perhaps it is because per weight, the harvested plants and a quarter of lamb contain roughly the same amount of protein? Who knows? Despite all this talk of sheep, however, large quantities of lambsquarters are reported to be poisonous to both sheep and pigs.

Though lambsquarters prefers nutrient-rich soils, it tolerates a wide variety of soil types, including dry, compacted, urban soil. It is drawn to all sorts of disturbed sites and is particularly abundant in gardens, agricultuaral fields, and roadsides. It readily hybridizes with other Chenopodium species, including the North American native C. berlandieri. In The Book of Field and Roadside, John Eastman calls it “one of the wold’s most abundant and noxious weeds,” because “it competes with some 40 crops [and] is especially invasive in tomato, potato, sugar beet, soybean, and corn fields.”

Eastman goes on to hint at lambsquarters’ potential for phytoremediation: “The plant accumulates high levels of nitrates and pesticides in addition to its oxalic acid content.” It also takes up heavy metals, including zinc, copper, and lead. This phenomenon is worth a future post, so stay tuned.

Leaf of lambsquarters (Chenopodium album)

That being said, when harvested from a non-polluted site, lambsquarters is a very nutritious spinach-like green both raw and cooked. Younger leaves and plants are preferred because older ones tend to be higher in oxalic acid. The seeds are also edible and, like quinoa, can be used in a similar manner as common grain and cereal crops. Harvester ants and various bird species also collect and consume the seeds. The roots of lambsquarters are high in saponin and can be used to make soap.

There are many reasons to be impressed with Chenopodium album, including its ability to tolerate drougt and frost, its adaptability to all types of soil, its highly nutritious plant parts (but also potentially toxic due to accumalation of pollutants and oxalic acid), and its competitive and persistent nature. Ehrenfried Pfeiffer, author of Weeds and What They Tell, was in awe of this “most enduring annual weed” and its goosefoot family relatives, writing: “We have the feeling that the goosefoot was destined to play a better role than to become an obnoxious weed. They follow closely man’s steps, showing their inclination to be domesticated. Probably future plant breeders may develop new cultivated varieties out of this family long after our present cultivated plants have degenerated, for it is their extreme vitality and preserverence to grow that makes the goosefoot family so interesting.”

Pfeiffer’s predictions haven’t quite come to pass, but time will tell.

More lambsquarters flowers


According to an article posted on LiveScience, lambsquarters is one of “The Five Healthiest Backyard Weeds.” The list includes two other weeds we have covered during the Summer of Weeds: Purslane and Plantain.


Summer of Weeds: Plantains

This is a revised version of two ethnobotany posts that appeared previously on Awkward Botany: White Man’s Foot, part one and part two. Plantains have a long history of ethnobotonical uses, as well as a bad reputation of being pesky, hard-to-eliminate weeds. The two most common introduced plantain species in North America are broadleaf plantain (Plantago major) and lanceleaf plantain (Plantago lanceolata). Wherever our daily travels take us, chances are there is a plantain growing nearby.


Plantago major is in the plantain family (Plantaginaceae) a family that consists of at least 90 genera, including common ornamental plants like Veronica (speedwells), Digitalis (foxgloves), and Antirrhinum (snapdragons). The genus Plantago, commonly known as plantains, consists of around 200 species distributed throughout the world in diverse habitats. Most of them are herbaceous perennials with similar growth habits.

Originating in Eurasia, P. major now has a cosmopolitan distribution. It has joined humans as they have traveled and migrated from continent to continent and is now considered naturalized throughout most temperate and some tropical regions. P. major has a plethora of common names – common plantain being the one that the USDA prefers. Other names include broadleaf plantain, greater plantain, thickleaf plantain, ribgrass, ribwort, ripplegrass, and waybread. Depending on the source, there are various versions of the name white man’s foot. Along the same line, a common name for P. major in South Africa is cart-track plant.

common plantain (Plantago major)

Common plantain starts by forming a rosette of broad leaves usually oriented flat against the ground. The leaves are egg-shaped with parallel veins; occasionally, leaf margins are faintly toothed. The inflorescence is a leafless spike up to 20 centimeters tall or taller with several tiny flowers that are a dull yellow-green-brown color. The flowers are wind pollinated and highly prone to self-pollination. The fruits are capsules that can contain as many as 30 seeds; an entire plant can produce as many as 15,000 seeds. The seeds are small, brown, sticky, and easily transported by wind or by adhering to shoes, clothing, animals, and machinery. They require light to germinate and can remain viable for up to 60 years.

Common plantain prefers sunny sites but can also thrive in part shade. It adapts to a variety of soil types but performs best in moist, clay-loam soils. It is often found in compacted soils and is very tolerant of trampling. This trait, along with its low-growing leaves that easily evade mower blades, explains why it is so common in turf grass. It is highly adaptable to a variety of habitats and is particularly common on recently disturbed sites (both natural and human caused). It is an abundant urban and agricultural weed.

Illustration of three Plantago species from Selected Weeds of the United States (Agriculture Handbook No. 366) circa 1970

Even though it is wind pollinated, its flowers are visited by syrphid flies and various bee species which feed on its pollen. Several other insects feed on its foliage, along with a number of mammalian herbivores. Cardinals and other bird species feed on its seeds.

Humans also eat plantain leaves, which contain vitamins A, C, and K. Young, tender leaves can be eaten raw, while older leaves need to be cooked as they become tough and stringy with age. The medicinal properties of  P. major have been known and appreciated at least as far back as the Anglo-Saxons, who likely used a poultice made from the leaves externally to treat wounds, burns, sores, bites, stings, and other irritations. It has also been used to stop cuts from bleeding and to treat rattlesnake bites. Apart from external uses, the plant was used internally as a pain killer and to treat ulcers, diarrhea, and other gastrointestinal issues.

P. major has been shown to have antibacterial, anti-inflammatory, antioxidant, and other biological properties; several chemical compounds have been isolated from the plant and deemed responsible for these properties. It is for this reason that P. major and other Plantago species have been used to treat such a diverse number of ailments. The claims are extensive and worth exploring. You can start by visiting the following sites:

Excerpts about plantains from The Book of Field and Roadside by John Eastman

Concerning their cosmopolitan nature: “Although both plantains [P. major and P. lanceolata] are Eurasian natives, they have long been thoroughly naturalized global residents; the designation ‘alien’ applies to them in the same sense that all white and black Americans are alien residents.”

In which I learned a new term: “Both species are anthropophilic (associate with humans); they frequent roadsides, parking areas, driveways, and vacant lots, occurring almost everywhere in disturbed ground. Where one species grows, the other can often be found nearby.”

Illustration by Amelia Hansen from The Book of Field and Roadside by John Eastman

Medicinal and culinary uses according to Eastman: “Plantains have versatile curative as well as culinary properties; nobody need go hungry or untreated for sores where plantains grow. These plants contain an abundance of beta carotene, calcium, potassium, and ascorbic acid. Cure-all claims for common plantain’s beneficial medical uses include a leaf tea for coughs, diarrhea, dysentery, lung and stomach disorders, and the root tea as a mouthwash for toothache. … Their most frequent and demonstrably effective use as a modern herb remedy, however, is as a leaf poultice for insect bites and stings plus other skin irritations. The leaf’s antimicrobial properties reduce inflammation, and its astringent chemistry relieves itching, swelling, and soreness.”

Even the seeds are “therapeutic”: “The gelatinous mucilage surrounding seeds can be readily separated, has been used as a substitute for linseed oil. Its widest usage is in laxative products for providing bulk and soluble fiber called psyllium, mainly derived from the plantain species P. ovata and leafy-stemmed plantain (P. psyllium), both Mediterranean natives.”

An excerpt from Weeds: In Defense of Nature’s Most Unloved Plants by Richard Mabey

“Plantain, ‘the mother of worts,’ is present in almost all the early prescriptions of magical herbs, back as far as the earliest Celtic fire ceremonies. It isn’t clear why such a drab plant – a plain rosette of grey-green leaves topped by a flower spike like a rat’s-tail – should have had pre-eminent status. But its weediness, in the sense of its willingness to tolerate human company, may have had a lot to do with it. The Anglo-Saxon names ‘Waybroad’ or ‘Waybread’ simply mean ‘a broad-leaved herb which grows by the wayside.’ This is plantain’s defining habit and habitat. It thrives on roadways, field-paths, church steps. In the most literal sense it dogs human footsteps. Its tough, elastic leaves, growing flush with the ground, are resilient to treading. You can walk on them, scuff them, even drive over them, and they go on living. They seem to actively prosper from stamping, as more delicate plants around them are crushed. The principles of sympathetic magic, therefore, indicated that plantain would be effective for crushing and tearing injuries. (And so it is, to a certain extent. The leaves contain a high proportion of tannins, which help to close wounds and halt bleeding.)”


Seed Dispersal via Caching – The Story of Antelope Bitterbrush

Generally speaking, individual plants produce an enormous amount of seeds. This may seem like a huge waste of resources, but the reality is that while each seed has the potential to grow into an adult plant that will one day produce seeds of its own, relatively few may achieve this. Some seeds will be eaten before they get a chance to germinate. Others germinate and soon die from lack of water, disease, or herbivory. Those that make it past the seedling stage continue to face similar pressures. Reaching adulthood, then, is a remarkable achievement.

Antelope bitterbrush is a shrub that produces hundreds of seeds per individual. Each seed is about the size of an apple seed. Some seeds may be eaten right away. Others fall to the ground and are ignored. But a large number are collected by rodents and either stored in burrows (larder hoarding) or in shallow depressions in the soil (scatter hoarding). It is through caching that antelope bitterbrush seeds are best dispersed. When rodents fail to return to caches during the winter, the seeds are free to sprout in the spring. Some of the seedlings will dry out and others will be eaten, but a few will survive, making the effort to produce all those seeds worth it in the end.

Fruits forming on antelope bitterbrush (Purshia tridentata)

Antelope bitterbrush (Purshia tridentata) is in the rose family and is often simply referred to as bitterbrush. It occurs in grasslands, shrub steppes, and dry woodlands throughout large sections of western North America. It is a deciduous shrub that generally reaches between three and nine feet tall but can grow up to twelve feet. It has wedge-shaped leaves that are green on top, grayish on bottom, and three-lobed. Flowers are yellow, strongly fragrant, and similar in appearance to others in the rose family. Flowering occurs mid-spring to early summer. Fruits are achenes – single seeds surrounded by papery or leathery coverings. The covering must rot away or be removed by animals before the seed can germinate.

Bitterbrush is an important species for wildlife. It is browsed by mule deer, pronghorn antelope, bighorn sheep, and other ungulates, including livestock. It provides cover for birds, rodents, reptiles, and ungulates. Its seeds are collected by harvester ants and rodents, its foliage is consumed by tent caterpillars and other insects, and its flowers are visited by a suite of pollinators. For all that it offers to the animal kingdom, it also relies on it for pollination and seed dispersal. The flowers of bitterbrush are self-incompatible, and if it wasn’t for ants and rodents, the heavy seeds – left to rely on wind and gravity – would have trouble getting any further than just a few feet from the parent plant.

Antelope bitterbrush (Purshia tridentata) in full bloom – photo credit: wikimedia commons

In a study published in The American Naturalist (February 1993), Stephen Vander Wall reported that yellow pine chipmunks were the primary dispersal agents of bitterbrush seeds in his Sierra Nevada study area. The optimal depth for seedling establishment was between 10-30 millimeters. Seeds that are cached too near the surface risk being pushed out of the ground during freeze and thaw cycles where they can desiccate upon germination. Cached bitterbrush seeds benefit when there are several seeds per cache because, as Vander Wall notes, “clumps of seedlings are better able to push through the soil and can establish from greater depths than single seedlings.”

Another study by Vander Wall, published in Ecology (October 1994), reiterated the importance of seed caching by yellow pine chipmunks in the establishment of bitterbrush seedlings. Seed caches, which consisted of anywhere from two to over a hundred seeds, were located as far as 25 meters from the parent plant. Cached seeds are occasionally moved to another location, but Vander Wall found that even these secondary caches produce seedlings. Of course, not all of the seedlings that sprout grow to maturity. Vander Wall states, “attrition over the years gradually reduces the number of seedlings within clumps.” Yet, more than half of the mature shrubs he observed in his study consisted of two or more individuals, leading him to conclude that “they arose from rodent caches.”

A study published in the Journal of Range Management (January 1996) looked at the herbivory of bitterbrush seedlings by rodents. In the introduction the authors discuss how “rodents [may] not only benefit from antelope bitterbrush seed caches as a future seed source, but also benefit from the sprouting of their caches as they return to graze the cotyledons of germinating seeds.”  In this study, Ord’s kangaroo rats, deer mice, and Great Basin pocket mice were all observed consuming bitterbrush seedlings, preferring them even when millet was offered as an alternative. The two species of mice also dug up seedlings, possibly searching for ungerminated seeds. Despite seed dispersal via caching, an overabundance of rodents can result in few bitterbrush seedlings reaching maturity.

A cluster of antelope bitterbrush seedlings that has been browsed. “Succulent, young seedlings are thought to be important in the diets of rodents during early spring because of the nutrients and water they contain.” — Vander Wall (1994)


Photos of antelope bitterbrush seedling clusters were taken at Idaho Botanical Garden, where numerous clusters are presently on display along the pathways of the native plant gardens and the adjoining natural areas. 


Beavers and Water Lilies – An Introduction to Zoochory

Beavers are classic examples of ecosystem engineers. It is difficult to think of an animal – apart from humans – whose day-to-day activities have more impact on the landscape than beavers. Their dam building activities create wetlands that are used by numerous other species, and their selective harvesting of preferred trees affects species composition in riparian areas. And that’s just the start. Their extensive evolutionary history and once widespread distribution has made them major players in the landscape for millions of years.

Today, the beaver family (Castoridae) consists of just two extant species: Castor fiber (native to Eurasia) and Castor canadensis (native to North America). Both species were hunted by humans to the brink of extinction but, thanks to conservation efforts, enjoy stable populations despite having been eliminated from much of their historical ranges. Before the arrival of Europeans, North American beavers are estimated to have been anywhere from 60 million to 400 million strong. Extensive trapping reduced the population to less than half a million. Today, 10 million or more make their homes in rivers, streams, and wetlands across the continent.

North American beaver (Castor canadensis) - photo credit: wikimedia commons

North American beaver (Castor canadensis) – photo credit: wikimedia commons

Beavers are herbivores, and they harvest trees and shrubs to build dams and lodges. Their interactions with plants are legion, and so what better way to introduce the concept of animal-mediated seed dispersal than beavers. Plants have several strategies for moving their seeds around. Wind and gravity are popular approaches, and water is commonly used by plants both aquatic and terrestrial. Partnering with animals, however, is by far the most compelling method. This strategy is called zoochory.

Zoochory has many facets. Two major distinctions are epizoochory and endozoochory. In epizoochory, seeds become attached in some form or fashion to the outside of an animal. The animal unwittingly picks up, transports, and deposits the seeds. The fruits of such seeds are equipped with hooks, spines, barbs, or stiff hairs that help facilitate attachment to an animal’s fur, feathers, or skin. A well known example of this is the genus Arctium. Commonly known as burdock, the fruits in this genus are called burs – essentially small, round balls covered in a series of hooks. Anyone who has walked through – or has had a pet walk through – a patch of burdocks with mature seed heads knows what a nuisance these plants can be. But their strategy is effective.

The burs of Arctium - photo credit: wikimedia commons

The burs of Arctium – photo credit: wikimedia commons

Endozoochory is less passive. Seeds that are dispersed this way are usually surrounded by fleshy, nutritious fruits desired by animals. The fruits are consumed, and the undigested seeds exit out the other end of the animal with a bit of fertilizer. Certain seeds require passage through an animal’s gut in order to germinate, relying on chemicals produced during the digestion process to help break dormancy. Other seeds contain mild laxatives in their seed coats, resulting in an unscathed passage through the animal and a quick deposit. Some plants have developed mutualistic relationships with specific groups of animals regarding seed dispersal by frugivory. When these animal species disappear, the plants are left without the means to disperse their seeds, which threatens their future survival.

Beavers rely on woody vegetation to get them through the winter, but in warmer months, when herbaceous aquatic vegetation is abundant, such plants become their preferred food source. Water lilies are one of their favorite foods, and through both consumption of the water lilies and construction of wetland habitats, beavers help support water lily populations. This is how John Eastman puts it in The Book of Swamp and Bog: “Beavers relish [water lilies], sometimes storing the rhizomes. Their damming activities create water lily habitat, and they widely disperse the plants by dropping rhizome fragments hither and yon.”

Fragrant water lily (Nympaea odorata) - photo credit: wikimedia commons

Fragrant water lily (Nymphaea odorata) – photo credit: wikimedia commons

The seeds of water lilies (plants in the family Nymphaceae) are generally dispersed by water. Most species (except those in the genera Nuphar and Barclaya) have a fleshy growth around their seeds called an aril that helps them float. Over time the aril becomes waterlogged and begins to disintegrate. At that point, the seed sinks to the bottom of the lake or pond where it germinates in the sediment. The seeds are also eaten by birds and aquatic animals, including beavers. The aril is digestible, but the seed is not.

In her book, Once They Were Hats, Frances Backhouse writes about the relationship between beavers and water lilies. She visits a lake where beavers had long been absent, but were later reintroduced. She noted changes in the vegetation due to beaver activity – water lilies being only one of many plant species impacted.

Every year in late summer, the beavers devoured the seed capsules [of water lilies], digested their soft outer rinds and excreted the ripe undamaged seeds into the lake. Meanwhile, as they dredged mud from the botom of the lake for their construction projects, they were unintentionally preparing the seed bed. Seeing the lilies reminded me that beavers also inadvertantly propagate willows and certain other woody plants. When beavers imbed uneaten sticks into dams or lodges or leave them lying on moist soil, the cuttings sometimes sprout roots and grow.

Other facets of zoochory include animals hoarding fruits and seeds to be eaten later and then not getting back to them, or seeds producing fleshy growths that ants love called elaiosomes, resulting in seed dispersal by ants. Animals and plants are constantly interacting in so many ways. Zoochory is just one way plants use animals and animals use plants, passively or otherwise. These relationships have a long history, and each one of them is worth exploring and celebrating.