Inside of a Seed: Two Monocots

“Seeds are travelers in space and time – small packages of DNA, protein, and starch that can move over long distances and remain viable for hundreds of years. These packages have everything they need not only to survive, but also to grow into a plant when they encounter the right conditions.”      The Book of Seeds by Paul Smith


As illustrated in last week’s post, the mature seeds of dicots – depending on the species – can be either with or without endosperm (a starchy food packet that feeds a growing seedling upon germination). Seeds without endosperm store these essential sugars in their cotyledons. Monocotyledons (or monocots, for short) are a group of flowering plants (i.e. angiosperms) whose seedlings are composed of a single cotyledon. With the exception of orchids, the seeds of monocots always contain endosperm.

The first of two examples of monocot seeds is the common onion (Allium cepa). The embryo in this seed sits curled up, surrounded by endosperm inside of a durable seed coat.

If you have ever sown onion seeds, you have watched as the single, grass-like cotyledon emerges from the soil. The seed coat often remains attached to the tip of the cotyledon like a little helmet as it stretches out towards the sky. Soon the first true leaf appears, pushing out from the base of the cotyledon. The source of this first leaf is the plumule hidden within the cotyledon.

The fruit of plants in the grass family – including cereal grains like wheat, oats, barley, rice, and corn – is called a caryopsis. In this type of fruit, the fruit wall (or pericarp) is fused to the seed coat, making the fruit indistinguishable from the seed. The embryos in these seeds are highly developed, with a few more discernible parts. A simplified diagram of a corn seed (Zea mays) is shown below. Each kernel of corn on a cob is a caryopsis. These relatively large seeds are great for demonstrating the anatomy of seeds in the grass family.

In these seeds there is an additional layer of endosperm called aleurone, which is rich in protein and composed of living cells. The cells of the adjacent endosperm are not alive and are composed of starch. The embryo consists of several parts, including the cotyledon (which, in the grass family, is also called a scuttelum), coleoptile, plumule, radicle, and coleorhiza. The coleoptile is a sheath that protects the emerging shoot as it pushes up through the soil. The plumule is the growing point for the first shoots and leaves, and the radicle is the beginning of the root system. The emerging root is protected by a root cap called a calyptra and a sheath called a coleorhiza.

Germination begins with the coleorhiza pushing through the pericarp. It is quickly followed by the radicle growing through the coleorhiza. As the embryo emerges, a signal is sent to the endosperm to start feeding the growing baby corn plant, giving it a head start until it can make its own food via photosynthesis.

corn seeds (Zea mays)

Up Next: We’ll take an inside look at the seeds of gymnosperms.


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Inside of a Seed: Two Dicots

“A seed is a living thing that embodies roots, stems, leaves, and fruit in an embryonic state and retains the ability to convert the sun’s energy into a source of food.” — Seedtime by Scott Chaskey


Few things are more miraculous than seeds. Within them is a living plant in embryonic form. Under the right conditions, these tiny objects expand, pushing out the beginnings of the most minuscule weed to the most humongous tree. Looking at these otherwise unassuming specks, you would hardly guess that they held such potential.

Housed in a seed is the genetic material necessary for growth and reproduction, along with some stored sugars to get the plant started. All of this is enclosed in a protective case. It is a rare moment in a plant’s life – a time when it isn’t rooted in place and can, for a brief period, move around. With the help of agents like wind, water, and animals it can travel anywhere in the world, venturing as far as inches or miles from its parent plant. As long as it finds a suitable place to grow, its voyage is not in vain.

Seeds are the result of sexual reproduction in plants (with rare exceptions, which we will cover in a future post). After pollination, a pollen grain sends three haploid cells into the ovule of a flower. These cells unite with the haploid cells found within. One germ cell from the pollen grain goes to the formation of an embryo, while the other two cells help form endosperm, the food source for the developing embryo. The wall of the ovule becomes the outer layer of the seed, known as the seed coat or testa. The seed matures as the fruit it is nested in ripens. Eventually, the fetal plant within the seed is ready to find a new home.

Seed heads of rubber rabbitbrush (Ericameria nauseosa) – the fuzzy pappus attached to the fruits allows seeds to float in the breeze and travel away from their parent plant.

As with so many things in biology, there is no single type of seed. When it comes to seed anatomy, most seeds consist of the same basic components, but each species of plant has its own unique seed. In fact, a well-trained taxonomist can identify plants simply by observing their seeds. With such a wide variety of seeds, it is difficult to organize them into discrete categories, but we still try. What follows is an introduction to two types of seeds – endospermic and non-endospermic – using two basic examples.

The first thing you should know about these two examples is that both species are dicotyledons (or dicots, for short). This means that when the baby plant emerges, it has two cotyledons, which are also called embryonic leaves because they look like little leaves. All flowering plants have been divided into two groups based on the number of cotyledons they have, the second group being the monocotolydons (or monocots) which have only one cotyledon. This is an old-fashioned way to classify plants, but it is still useful in some instances.

Endospermic Seeds

The seeds of the castor bean plant (Ricinus communis) are endospermic seeds. This means that they retain the endosperm that was formed when two pollen grain cells joined up with the haploid cell in the ovule. The endosperm will help feed the growing embryo as it germinates. The two cotyledons are visible within the seed, but they are thin and broad, leaving plenty of space in the seed for the endosperm. The cotyledons are part of the embryo and are attached to the radicle, which is the embryonic root. The radicle is the first thing to emerge from the seed upon germination. The area between the radicle and the cotyledon is known as the hypocotyl. It becomes the stem of the germinating seedling.

An elaisome is attached to the outside of the seed coat of castor bean seeds. This fleshy, nutrient-rich appendage is particularly attractive to ants. They carry the seeds back to their colony and feed the elaisome to their young. The seeds, however, remain unconsumed. In this way, the ants aid in the seeds’ dispersal.

seeds of castor beans (Ricinus communis)

Non-endospermic Seeds

The seeds of plants in the bean family (Fabaceae) are non-endospermic seeds. This means that as the embryo develops, it uses up the majority of the endosperm within the seed. The food necessary for the seedling to get its start is all stored in its cotyledons. The common pea (Pisum sativum) is a good example of this. The embryo – which consists of the cotyledons, plumula (or plumule), hypocotyl, and radicle – takes up all available space inside of the seed coat. After germination, as the seedling develops, the plumule appears above the cotyledons and is the growing point for the first true leaves and stems.

seeds of the common pea (Pisum sativum)

In future posts, we will look at a few other types of seeds, as well as discuss various other seed-related topics. If you have a story to share about seeds, please do so in the comment section below.

Attempts to Avenge the Acts of Cirsium arvense – A Biocontrol Story

Some weeds are so noxious, their crimes so heinous, and their control so challenging that desperation leads us to introduce other non-native organisms to contain them. Alien vs. alien duking it out in a novel environment. It seems counterintuitive – if an introduced species has reached the status of invasive, is it worth the risk of bringing in yet another foreign species in attempt to defeat it? We all know what happened to the old lady who swallowed the spider to catch the fly, yet for decades now we have been doing just this. It’s something we call classical biological control – introducing pathogens, insects, or other organisms to help control the spread of problematic ones.

Such attempts mostly fail, but we keep trying. The attempts made on Cirsium arvense exemplify this. The trouble is that even when such efforts fail, they aren’t always benign, as we shall see.

Canada thistle, a misnomer for Cirsium arvense, is a European native that has been acting in the role of noxious weed for centuries, even in its native land. First introduced accidentally to eastern North America sometime in the 1600’s, it has made its way across the continent and has since become one of our worst weeds in both natural and agricultural settings, as well as in our yards and gardens. Its seeds get around, carried by wind and water, attached to animals or deposited in their dung, stowing away as contaminants in crop seed or passengers in the ballast water of ships. But casual dispersal by seed isn’t quite as troubling as what it does once it takes root.

Several related species of thistle are also pesky weeds, but unlike Cirsium arvense, they are mostly annuals or biennials, spreading only by seed. Cirsium arvense is a perennial plant with roots that spread deep and wide. New shoots form readily along the spreading roots, forming a veritable thicket of stems that can be dozens of feet wide and giving the plant a more appropriate common name, creeping thistle.

The stems of creeping thistle can grow more than four feet tall and are adorned with alternately arranged, prickly, lobed leaves. Groups of small, urn-shaped flowerheads are born at the tops of stems. Flowers are pink to purple, sweet smelling, and favored by pollinators. Individual plants either produce all male flowers or all female flowers, and since individual plants are actually large colonies, an adjacent colony of the opposite sex is necessary in order for the production of viable seeds. Like other plants in the aster family, the seeds come with a feathery pappus, suggesting wind dispersal. However, the pappus is often weakly attached, sloughing off without seeds in tow, leaving them to the fate of gravity.

flowers of creeping thistle (Cirsium arvense) via eol

It comes as no surprise that when plants readily spread by root, stolon, or rhizome, they are well suited to become some of our most bothersome weeds. Eliminating their seed heads does little to reduce their spread. Pulling them out of the ground is futile; you will never get all the roots. Tilling them under only aids in their dispersal since chopped up roots and stems now have the chance to produce new plants. Herbicide treatments can set them back, but they must be repeated on a long-term and exacting schedule in order to thoroughly kill the roots. Considering what we’re up against when it comes to plants like creeping thistle, it makes sense why we would introduce foreign fighters to do our bidding, especially if such fighters are enemies of the plant in their native land.

The list of insects that have been employed (or at least considered) in the fight against creeping thistle is extensive. It includes thistle tortoise beetle (Cassida rubiginosa), seedhead weevil (Rhinocyllus conicus), thistle crown weevil (Trichosirocalus horridus), thistle gall fly (Urophora cardui), thistle stem weevil (Ceutorhynchus litura), thistle bud weevil (Larinus planus), seedhead fly (Orellia ruficauda), thistle flea beetle (Altica carduorum), thistle leaf beetle (Lema cyanella), painted lady (Vanessa cardui), and sluggish weevil (Cleonus piger). Unfortunately, and perhaps not surprisingly, as Bugwood reports, “biocontrol currently provides little or no control of Canada thistle populations, although some agents weaken and kill individual plants.” Despite the fact that there are well over 100 known organisms that consume or attack Cirsium arvense, nothing manages to do long-term damage.

thistle tortoise beetle (Cassida rubiginosa) – a common biocontrol agent of invasive thistle species (via wikimedia commons)

The status of creeping thistle biocontrol efforts on two South Dakota wildlife refuges was reported on in a 2006 issue of Natural Areas Journal. Multiple introductions of at least half a dozen different insect species had occurred beginning in 1986. Nearly 20 years later, they were not found to have had a significant effect on creeping thistle populations. The authors concluded stating they “do not advocate further releases or distribution in the northern Great Plains of the agents” examined in their study. They also advised that “effectiveness be a primary consideration” of any new biocontrol agents and expressed concern that some introduced insects have the potential to attack native thistles.

North America is home to quite a few native thistles, several of which are rare or threatened. A USDA guide to managing creeping thistle in the Southwest highlights the importance of protecting native thistles – “especially rare or endangered species” – from biocontrol agents and gives two examples of endangered thistles in New Mexico that are at risk of such agents.

The federally threatened species, Pitcher’s thistle (Cirsium pitcheri), which is restricted to sand dune shorelines along the upper Great Lakes, has quite a bit working against it. An added blow came a few years ago when it was discovered that the flowerheads of Pitcher’s thistle were being damaged by the thistle bud weevil (Larinus planus), a biocontrol agent employed against creeping thistle in the area. A paper published in Biological Conservation in 2012 examining the extent of weevil damage on the rare thistle cautioned that, “although some biological control agents may benefit some rare plant taxa, the negative impacts of both native insects and introduced herbivores are well documented.”

Pitcher’s thistle (Cirsium pitcheri) via eol

Classical biological control, if and when it works, can be quite valuable, especially if it reduces the need for other management inputs like herbicides and cultivation. Unfortunately, it is rarely successful and can have unintended consequences. Goldson et al. report in a 2014 issue of Biological Conservation that the success rate is only around 10% and that even that 10% is at risk of failing at some point. In his book, Where Do Camels Belong?, ecologist Ken Thompson cites that “only about one in three species introduced as biological controls establish at all, and only half of those that do establish (i.e. about 16% of total attempts) control the intended enemy,” adding that “biological control is just another invasion, albeit one we are trying to encourage rather than prevent, and its frequent failure is another example of how poorly we understand the effects of adding new species to ecosystems.”

Still, while some warn against being too optimistic, others argue that it is an essential tool in the war against invasive species and, while acknowledging that a few introductions have gone awry, assert that “significant non-target impacts” are rare. Clearly, this is a rich topic ripe for healthy debate and one that I will continue to explore. If you have thoughts or resources you’d like to share, please do so in the comment section below.


This post was inspired in part by episode six of The Shape of the World podcast. I highly recommend listening to the entire series.

Botany in Popular Culture: The Tan Hua Flowers in Crazy Rich Asians

When a flower blooms, a celebration is in order. Flowers abound for much of the year, which means parties are called for pretty much non-stop (something Andrew W.K. would surely endorse). Since we can’t possibly celebrate every bloom, there are certain plants we have decided to pay more attention to – plants whose flowers aren’t so prolific, predictable, or long-lived; or plants whose flowers come infrequently or at odd times of the day (or night).

This is the case with the flowers of the night blooming cactus, Epiphyllum oxypetalum, which goes by many names including Dutchman’s pipe cactus, queen of the night, orchid cactus, night blooming cereus, and tan hua. Tan hua is the Chinese name for the plant, and this is how it is referred to in the book, Crazy Rich Asians by Kevin Kwan.

In the book, Nick Young brings his American girlfriend, Rachel Chu, to meet his ridiculously wealthy family in Singapore. Before the trip, Rachel was in the dark about the Young’s wealth. She first meets the family and their gargantuan mansion when Nick’s grandma, seeing that her tan hua flowers are about to bloom, throws an impromptu (and lavish) party. Nick refers to the flowers as “very rare,” blooming “extremely infrequently,” and “quite something to witness.”

In a seperate conversation, Nick’s cousin, Astrid, tries to convince her husband to attend the party by claiming, “it’s awfully good luck to see the flowers bloom.” Later, another one of Nick’s cousins tells Rachel, “it’s considered to be very auspicious to witness tan huas blooming.”

Tan hua (Epiphyllum oxypetalum) via wikimedia commons

Native to Mexico and Guatemala, E. oxypetalum was first brought to China in the 1600’s. Its beauty and intrigue along with its relative ease of cultivation helped it become popular and widespread across Asia and other parts of the world. Watching it bloom is considered a sacred experience by many, including in India, where it is said to bring luck and prosperity to households who are fortunate enough to see theirs bloom.

Epiphyllums are epiphytic, meaning they grow non-parasitically on the surfaces of other plants, such as in the crevices of bark or the crotches of branches. Like other cacti, they are essentially leafless, but their stems are broad, flat, and leaf-like in appearance. Showy, fragrant flowers are born along the margins of stems. The flowers of tan hua, as described in Crazy Rich Asians, appear as “pale reddish petals curled tightly like delicate fingers grasping a silken white peach.” A report (accompanied by photos) published by Sacred Heart University describes watching tan hua flowers progess from bud formation to full bloom, a process that took more than two weeks.

Tan huas are certainly not rare, as Nick described them. A number of Epiphyllum species and their hybrids are commonly cultivated; there is even an Epiphyllum Trail at San Diego Zoo’s Safari Park. Listed as “least concern” on the IUCN Red List, their popularity as ornamentals is noted but is not seen as affecting wild populations. Night blooming plants, while fascinating, aren’t all that rare either. Such plants have adapted relationships with creatures, like bats and moths, that are active during the night, employing their assistance with pollination. A paper published in Plant Systematics and Evolution describes the floral characteristics of Epiphyllum and similar genera: “The hawkmoth-flower syndrome, consisting of strongly-scented, night-blooming flowers with white or whitish perianths and long slender nectar-containing floral tubes is present in Cereus, Trichocereus, Selenicereus, Discocactus, Epiphyllum, and a number of other cactus genera.”

That being said, the specialness of a short-lived, infrequent, night blooming flower should not be understated, and really, parties being thrown in honor of any plant are something I can certainly get behind. Sitting in the courtyard late at night, the Young family and their guests watched as “the tightly rolled petals of the tan huas unfurled like a slow-motion movie to reveal a profusion of feathery white petals that kept expanding into an explosive sunburst pattern.” The look of it reminds Astrid of “a swan ruffling its wings, about to take flight.”

Later, “the tan huas began to wilt just as swiftly and mysteriously as they had bloomed, filling the night air with an intoxicating scent as they shriveled into spent lifeless petals.”


Additional Resources:


*Thank you Kathy for letting me borrow your Kindle so that I could write this post.

Death by Crab Spider, part two

Crab spiders that hunt in flowers prey on pollinating insects. Thus, pollinating insects tend to avoid flowers that harbor crab spiders. We established this in part one. Now we ask, what effect, if any, does this interaction have on a crab spider infested plant’s ability to reproduce? More importantly, what are the evolutionary implications of this relationship?

In a study published in Ecological Entomology earlier this year, Gavini, et al. found that pollinating insects avoided the flowers of Peruvian lily (Alstroemeria aurea) when artificial spiders of various colors and sizes were placed in them. Bumblebees and other bees were the most frequent visitors to the flowers and were also the group “most affected by the presence of artificial spiders, decreasing the number of flowers visited and time spent in the inflorescences.” This avoidance had a notable effect on plant reproduction, namely a 25% reduction in seed set and a 15% reduction in fruit weight. The most abundant and effective pollinator, the buff-tailed bumblebee, was deterred by the spiders, leading the researchers to conclude that, “changes in pollinator behavior may translate into changes in plant fitness when ambush predators alter the behavior of the most effective pollinators.”

Peruvian lily (Alstroemeria aurea) via wikimedia commons

But missing from this discussion is the fact that crab spiders don’t only eat pollinators. Any flower visiting insect may become a crab spider’s prey, and that includes florivores. In which case, crab spiders can benefit a plant, saving it from reproduction losses by eating insects that eat flowers.

In April of this year, Nature Communications published a study by Knauer, et al. that examined the trade-off that occurs when crab spiders are preying on both pollinators and florivores. Four populations of buckler-mustard (Biscutella laevigata ssp. laevigata) were selected for this study. Bees are buckler-mustard’s main pollinator, and in concurrence with other studies, they significantly avoided flowers when crab spiders were present.  Knauer, et al. also determined that bees and crab spiders are attracted to the same floral scent compound, β-ocimene. This compound not only attracts pollinators, but is also emitted when plants experience herbivory, possibly to attract predators to come and prey on whatever is eating them.

buckler-mustard (Biscutella laevigata) via wikimedia commons

In this study, the predators called upon were crab spiders. Florivores had a notable impact on plants in this study, and the researchers found that when crab spiders were present, florivores were significantly reduced, thereby reducing their negative impact. They also noted that “crab spiders showed a significant preference for [florivore-infested] plants over control plants.”

And so it is, a plant’s floral scent compound attracts pollinators while simultaneously attracting the pollinator’s enemy, who is also called in to protect the flower from being eaten. Luckily, in this case, buckler-mustard is easily pollinated, so the loss of a few pollinators isn’t likely to have a strong negative effect on reproduction. As the authors write, “pollinators are usually abundant and the low number of ovules per flower makes a few pollen grains sufficient for a full seed set.”

crab spider on zinnia

But none of these studies are one size fits all. Predator-pollinator-plant interactions are still not well understood, and there is much to learn through future research. A meta-analysis published in the Journal of Animal Ecology in 2011 looked at the research that had been done up to that point. Included were a range of studies involving sit-and-wait predators (like crab spiders and lizards) as well as active hunters (like birds and ants) and the effects of predation on both pollinators and plant-eating insects. They concluded that where carnivores “disrupted plant-pollinator interactions, plant fitness was reduced by 17%,” but thanks to predation of herbivores, carnivores helped increase plant fitness by 51%. This suggests that carnivores, overall, have a net positive effect on plant fitness.

Many pollinating insects have an advantage over plant-eating insects because they move quickly from flower to flower and plant to plant, unlike many herbivores which move more slowly. This protects pollinators from predation and helps explain why plant-pollinator interactions are not disrupted as easily by carnivores. Additionally, as the authors note, “plants may be buffered against loss of pollination by attracting different types of pollinators, some of which are inaccessible to carnivores.”

But again, there is still so much to discover about these complex interactions. One way to gain a better understanding is to investigate the effects of predators on both pollinators and herbivores in the same study, since many of the papers included in the meta-analysis focused on only one or the other. As far as crab spiders go, Knauer, et al. highlight their importance in such studies. There are so many different species of crab spiders, and they are commonly found on flowers around the globe, so “their impact on plant evolution may be widespread among angiosperms.”

In other words, while we still have a lot to learn, the impact these tiny but skillful hunters have should not be underestimated.

Death by Crab Spider, part one

When a bee approaches a flower, it is essentially approaching the watering hole. It comes in search of food in the form of pollen and nectar. As is this case with other animals who come to feed at the watering hole, a flower-visiting bee makes itself vulnerable to a variety of predators. Carnivores, like the crab spider, lie in wait to attack.

The flowers of many plants rely on visits from bees and other organisms to assist in transferring pollen from stamens to stigmas, which initiates reproduction; and bees and other flower visitors need floral resources to survive. Crab spiders exploit this otherwise friendly relationship and, in doing so, can leave lasting impacts on both the bees and the flowers they visit.

Species in the family Thomisidae are commonly referred to as crab spiders, a name that comes from their resemblance to crabs. Crab spiders don’t build webs to catch prey; instead they either actively hunt for prey or sit and wait for potential prey to happen by, earning them the name ambush predators. Of the hundreds of species in this family, not all of them hunt for prey in flowers; those that do – species in the genera Misumena and Thomisus, for example – are often called flower crab spiders.

white crab spider (Thomisus spectabilis) on Iris sanguinea — via wikimedia commons

Most crab spiders are tiny – mere millimeters in size – and they have a number of strategies (depending on the species) to obscure their presence from potential prey. They can camouflage themselves by choosing to hunt in a flower that is the same color as they are or, in the case of some species, they can change their color to match the flower they are on. Some species of crab spiders reflect UV light, which bees can see. In doing so, they make themselves look like part of the flower.

Using an Australian species of crab spider, researchers found that honey bees preferred marguerite daisies (Chrysanthemum frutescens) on which UV-reflecting crab spiders were present, even when the scent of the flowers was masked. The spiders’ presence was seen as nectar guides, which “bees have a pre-existing bias towards.” Members of this same research team also determined that both crab spiders and honey bees choose fragrant flowers over non-fragrant flowers, and that, ultimately, “honey bees suffer apparently from responding to the same floral characteristics as crab spiders do.”

Needless to say, crab spiders are crafty. So the question is, when killing machines like crab spiders are picking off a plant’s pollinators, does this affect its ability to reproduce? First let’s consider how pollinators react to finding crab spiders hiding in the flowers they hope to visit.

goldenrod crab spider (Misumena vatia) preying on a pollinator — via wikimedia commons

A study published in Oikos in 2003 observed patches of common milkweed (Asclepias syriaca) – one set was free of crab spiders, the other set was not – and tracked the visitations of four species of bees – the common honey bee and three species of bumble bees. They compared visitation rates between both sets of milkweed patches and found that the smallest of the three bumble bee species decreased its frequency of visitation to the crab spider infested milkweeds. Honey bees also appeared to visit the infested milkweeds less, but the results were not statistically significant. The two larger species of bumble bees continued to forage at the same rate despite the presence of crab spiders.

During the study, crab spiders were seen attacking bees numerous times. Six attacks resulted in successful kills, and of the bees that escaped, 80% left the flower and either moved to a different flower on the same plant, moved to a different plant, or left the patch altogether. These results indicate a potential for the presence of crab spiders to effect plant-pollinator interactions, whether its directly (predation) or indirectly (bees avoiding flowers with crab spiders).

Another study published in Behavioral Ecology in 2006 looked at two species of bees – the honey bee and a species of long-horned bee – and their reactions to the presence of crab spiders on the flowers of three different plant species – lavender (Lavandula stoechas), crimson spot rockrose (Cistus ladanifer), and sage-leaf rockrose (Cistus salvifolius). Honey bees were about half as likely to select inflorescences of lavender when crab spiders were present, and they avoided the crab spider infested flowers of crimson spot rockrose with a similar frequency. On the other hand, the long-horned bee visited the flowers of crimson spot rockrose to the same degree whether or not a crab spider was present.

bee visiting sage-leaf rock rose (Cistus salvifolius) — via wikimedia commons

The researchers then exposed honey bees to the flowers of sage-leaf rockrose that were at the time spider-free but showed signs that crab spiders had recently visited. Some of the flowers featured the scent of crab spiders, others had spider silk attached to them, and others had the corpses of dead bees on them. They found that even when crab spiders were no longer present, the bees could still detect them. Honey bees were particularly deterred by the presence of corpses. The long-horned bees were also exposed to the flowers with corpses on them but didn’t show a significant avoidance of them.

An interesting side note about the presence of silk on flowers. As stated earlier, crab spiders do not spin webs; however, they do spin silk for other reasons, including to tether themselves to flowers while hunting. The authors recount, “on several occasions when an attempted attack was observed during this study, it was only the presence of a silk tether that prevented spiders being carried away from flowers by their much larger prey.”

So, again, if bees are avoiding flowers due to the presence of predators like crab spiders, what effect, if any, is this having on the plants? We will address this question in part two.

Eating Weeds: Clovers

If you ever spent time hunting for four-leaf clovers in the lawn as a kid, in all likelihood you were seeking out the leaves of Trifolium repens or one of its close relatives. Commonly known as white clover, the seeds of T. repens once came standard in turfgrass seed mixes and was a welcome component of a healthy lawn thanks to its ability to fix atmospheric nitrogen and provide free fertilizer. But around the middle of the 20th century, when synthetic fertilizers and herbicides became all the rage, clover’s reputation shifted from acceptable to disreputable. Elizabeth Kolbert, in an article in The New Yorker about American lawns, recounts the introduction of the broadleaf herbicide 2,4-D: “Regrettably, 2,4-D killed not only dandelions but also plants that were beneficial to lawns, like nitrogen-fixing clover. To cover up this loss, any plant that the chemical eradicated was redefined as an enemy.”

white clover (Trifolium repens) in turf grass

This particular enemy originated in Europe but can now be found around the globe. It has been introduced both intentionally and accidentally. Commonly cultivated as a forage crop for livestock, its seeds can be found hitchhiking to new locations in hay and manure. Clover honey is highly favored, and so clover fields are maintained for honey production as well. Its usefulness, however, doesn’t protect it from being designated as a weed. In Weeds of North America, white clover is accused of being “a serious weed in lawns, waste areas, and abandoned fields.”

White clover is a low-growing, perennial plant that spreads vegetatively as well as by seed. It sends out horizontal shoots called stolons that form roots at various points along their length, creating a dense groundcover. Its compound leaves are made up of three, oval leaflets, and its flower heads are globe-shaped and composed of up to 100 white to (sometimes) pink florets. Rich in nectar, the flowers of white clover draw in throngs of bees which assist in pollination. Closely related and similar looking strawberry clover, Trifolium fragiferum, is distinguished by its pink flowers and its fuzzy, rounded seed heads that resemble strawberries or raspberries. Red clover, T. pratense, grows more upright and taller than white and strawberry clovers and has red to purple flowers.

leaves and seed heads of strawberry clover (Trifolium fragiferum)

Clovers are tough plants, tolerating heat, cold, drought, and trampling. Lawns deprived of water go brown fairly quickly, revealing green islands of interlopers, like clover, able to hang in there throughout dry spells. These days, many of us are reconsidering our need for a lawn. Lawns are water hogs that require a fair amount of inputs to keep them green and free of weeds, pests, and diseases. The excessive amounts of fertilizers and pesticides dumped on them from year to year is particularly troubling.

Along with our reconsideration of the lawn has come clover’s return to popularity, and turfgrass seed mixes featuring clover are making a comeback. To keep clover around, herbicde use must be curbed, and so lawns may become havens for weeds once more. Luckily, many of those weeds, including clover, are edible, so urban foragers need only to step out their front door to find ingredients for their next meal.

The leaves and flowers of clover can be eaten cooked or raw. Fresh, new leaves are better raw than older leaves. That being said, clover is not likely to be anyone’s favorite green. Green Deane refers to it as a “survival or famine food” adding that “only the blossoms are truly pleasant to human tastes,” while “the leaves are an acquired or tolerated taste.” In The Book of Field and Roadside, John Eastman remarks: “As humanly edible herbs, clovers do not rank as choice. Yet they are high in protein and vitamins and can be eaten as a salad or cooked greens and in flower head teas. Flower heads and leaves are much more easily digested after boiling.”

I tried strawberry clover leaves and flower heads in a soup made from a recipe found in the The Front Yard Forager by Melany Vorass Herrara. Around two cups of clover chopped, cooked, and blended with potatoes, scallions, and garlic in vegetable or chicken broth is a fine way to enjoy this plant. I don’t anticipate eating clover with great frequency, partly because it is included in a list of wild edible plants with toxic compounds in The North American Guide to Common Poisonous Plants and Mushrooms and also because I have to agree with the opinions of the authors quoted above – there are better tasting green things. Either way, it’s worth trying at least once.

clover soup

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