Eating Weeds: Cleavers Coffee

One of the world’s most beloved beverages comes from a species of plant found in the fourth largest family of flowering plants. Rubiaceae, also known as the coffee or bedstraw family, consists of around 13,500 species, placing it behind just Asteraceae, Orchidaceae, and Fabaceae for the most number of species. Coffea arabica, and other species in the genus Coffea, are grown for their fruits which are used to make coffee. This makes Rubiaceae one of the most economically important plant families. A family this size is bound to be home to a weed or two, and in fact, one of the most widespread and obnoxious weeds is also a member of Rubiaceae.

Galium aparine, known commonly by a slew of names including cleavers, occurs naturally across large portions of Europe, Asia, North Africa, and possibly even parts of North America. It has been introduced as a weed in many locations across North America, South America, Australia, New Zealand, Japan, and parts of Africa. It is of particular concern in agricultural settings where its lengthy, sprawling branches and sticky leaves get tangled up in harvesting equipment, while its tiny, prickly fruits get mixed in with seeds of similar size like canola.

Galium aparine

Sticky willy, as it is also known, is an annual plant that, in some cases, can have two generations per year – one in the spring (having germinated the previous fall) and one in the summer. Its stems are square, though not as sharply square as plants in the mint family, and can grow to around six feet long. They are weak, brittle, and don’t stand upright on their own; instead they are found scrambling across the ground or, when given the opportunity, climbing up the lengths of other plants in order to reach the sunlight. Leaves occur in whorls of six to eight and are simple and slender with entire margins. Flowers are produced at leaf axils along the lengths of the branches and are tiny, four-petaled, star-shaped, and greenish white. Fruits are borne in pairs and are round, single-seeded, indehiscent nutlets. The stems, leaves, and fruits are covered in stiff, hooked hairs or trichomes, earning it other names like catchweed bedstraw, grip grass, stickyweed, and velcro plant.

flowers and immature fruit on Galium aparine

Galium aparine is a climbing plant, but unlike other climbing plants, it doesn’t twine up things or produce structures like tendrils to hold itself up. Instead, its ability to climb is made possible by its abundant bristly hairs. A paper published in Proceedings of the Royal Society B (2011) investigates the way G. aparine climbs up other plants using the hairs on its leaves. A close inspection of the leaves reveals that the trichomes on the top of the leaf (the adaxial leaf surface) differ significantly from those found on the bottom of the leaf (the abaxial leaf surface). Adaxial trichomes curve towards the tip of the leaf, are hardened mainly at the tip, and are evenly distributed across the leaf surface. Abaxial trichomes curve towards the leaf base, are hardened throughout, and are found only on the midrib and leaf margins.

Having different types of hairs on their upper and lower leaf surfaces gives cleavers an advantage when it comes to climbing up neighboring plants. The authors of the paper describe the technique as a “ratchet mechanism.” When the upper surface of their leaf makes contact with the lower surface of another plant’s leaf, the flexible, outwardly hooked trichomes inhibit it from slipping further below the leaf and allow it to easily slide out from underneath it. When the lower surface of their leaf makes contact with the upper surface of another plant’s leaf, the stiff, inwardly hooked trichomes keep it attached to the leaf even if the other leaf starts to slip away and allows it to advance further across the leaf for better attachment and coverage. Using this ratchet mechanism, cleavers climb up the leaves of other plants, keeping their leaves above the other plant’s leaves, which gives them better access to sunlight. The basal stems of cleavers are highly flexible, which keeps them from breaking as the plant sways in the wind, tightly attached to their “host” plant.

fruits of Galium aparine

The hooked trichomes on the tiny fruits of cleavers readily attach to the fur and clothing of passing animals. The nutlets easily break free from the plants and can be transported long distances. They can also be harvested and made into a lightly caffeinated tea. Harvesting the fruit takes time and patience. I spent at least 20 minutes trying to harvest enough fruits for one small cup of cleavers coffee. The fruits don’t ripen evenly, and while I tried to pick mostly ripe fruits, I ended up with a selection of fruits in various stages of ripeness.

To make cleavers coffee, first toast the seeds for a few minutes in a pan heated to medium high, stirring them frequently. Next, grind them with a mortar and pestle and place the grinds in a strainer. Proceed as you would if you were making tea from loose leaf tea.

The toasted fruits and resulting tea should smell similar to coffee. The smell must not be strong, because my poor sense of smell didn’t really pick up on it. The taste is coffee-like, but I thought it was more similar to black tea. Sierra tried it and called it “a tea version of coffee.” If the fruits were easier to collect, I could see myself making this more often, but who has the time?

The leaves and stems of Galium aparine are also edible, and the plant is said to be a particular favorite of geese and chickens, bringing about yet another common name, goosegrass. In the book Weeds, Gareth Richards discusses the plant’s edibility: “It’s edible for humans but not that pleasant to eat; most culinary and medicinal uses center around infusing the plant in liquids.” Cooking with the leaves or turning them into some sort of spring tonic is something I’ll consider for a future post about eating cleavers.

More Eating Weeds Posts on Awkward Botany

Apriums and Plumcots and Pluots, Oh My!

I was once a teenage paper carrier in small town Idaho. One of my stops was an apartment complex, and for much of the year, this was an uneventful stop. But for a few weeks in the summer, the purple-leaved plum trees out front had ripe fruit on them, and each time I was there, I would stop and take a few. In general, I don’t get that excited about fruit, but I enjoyed eating these plums. This variety of plum is typically planted for its looks rather than its fruit, and it may even be the tree that recently received a pitifully low score on an episode of Completely Arbortrary. Ornamental plum or not – and low cone score or not – I thought the fruit was good.

Many of the things we eat are a result of crosses between two related species, and plums are a great example of this. Species are species because they are reproductively isolated. A species does not typically mate with a member of another species and create viable offspring, except this happens all the time both naturally and artificially. In many cases, the offspring isn’t actually viable, but there is offspring nonetheless, and in the case of plants, that offspring can then reproduce asexually – by leaf, stem, or root cuttings or by some other means – and the resulting hybrid can exist indefinitely. One species mating with another species (specifically two species that are members of the same genus) is called interspecific hybridization, and there is a good chance that you’ve eaten something recently that is a result of this.

One of the most widely grown species of plum, Prunus domestica (commonly known as European plum), is a result of interspecific hybridization that occurred many centuries ago. A paper published in Horticulture Research (2019) confirmed that P. domestica originated as a cross between Prunus cerasifera and Prunus spinosa, the latter of which may have also been a result of interspecific hybridization. There are over 400 species in the genus Prunus that are distributed across temperate regions in the northern hemisphere. Within this genus is the subgenus Prunus (or Prunophora), a group that includes dozens of familiar species such as the plums, apricots, peaches, and almonds. Due to their close relationship, both natural and artificial hybridization among members of this subgenus is common, which explains the origin of Prunus domestica, as well as the majority of the plums we grow today.

Current commercial production of plums in North America is largely thanks to work done by Luther Burbank in the late 19th to early 20th centuries. Burbank was obsessed with plant breeding and released hundreds of new varieties of all kinds of different plants during his decades long career. He seemed particularly interested in plums, developing 113 different cultivars, which account for more than half of all his fruit releases. Probably his most well known plum variety is ‘Santa Rosa,’ which thanks to modern day genetics has been determined to be a cross between at least four different species of plum.

apriums

Early colonizers to the American continent were mainly growing varieties of the European plum they had brought over from Europe. North America is also home to several species of plums, which are used by indigenous populations. Shortly before Burbank began working with plums on his farm in California in 1881, Asian plum species were imported to the U.S., and breeders began using them in crosses with both European and North American plum species. Burbank became particularly engulfed in these efforts. In an article published in HortScience (2015), David Karp writes, “In the history of horticulture it is rare to find an individual who almost single-handedly created a new commercial industry based on a novel fruit type as Luther Burbank did for Asian-type plums in the United States.” Most Asian-type plums sold in stores today are hybrids of several different plum species due to the numerous complex crosses that Burbank made.

Burbank is also said to be the first to cross plums and apricots, creating the first of many cultivars of the plumcot. Plum and apricot crosses didn’t really catch on for a few more decades, and when they did, it was thanks to the work of Floyd Zaiger of Zaiger Genetics who developed and released numerous varieties. Apriums and Pluots are Zaiger Genetics trademarks, along with a few other unlikely crosses with plums and their related counterparts.

plumcots

A plumcot is the simplest cross. It is said to be 50% Asian plum (Prunus salicina) and 50% apricot (Prunus armeniaca). However, due to all the breeding of Asian plums carried out by Burbank and others, the Asian plum involved in the cross is typically a hybrid with other plum species, as discussed in a recent paper published in Plants (2022). An aprium is the result of a cross between a plumcot and an apricot, making it 75% apricot and 25% plum, while a pluot is a cross between a plumcot and a plum, making it 75% plum and 25% apricot. There is typically much more that goes into making these crosses, but that’s the general idea. If you’re lucky, you can find all three of these intraspecific crosses in a produce section near you, but it may not be clear what cultivar you’re purchasing. Myriad cultivars have been released of each of these hybrids – each one varying in color, size, flavor, disease resistance, etc. – and unfortunately most grocery stores don’t include cultivar names on their products, so it’s difficult to know what you’re getting.

At Awkward Botany Headquarters, there is a plum tree growing in our front yard. We didn’t plant it, so at this point I have no idea what species or cultivar it is. The plums are delicious though, and the leaves aren’t purple like the plums I used to eat on my paper route. Considering all of the intraspecific crossing that has gone on with plums, it’s quite likely that it is a combination of different species, which isn’t going to make it easy to figure out. But I’ll do my best.


Check out the linktree for various ways to follow and support Awkward Botany.

The Serotinous Cones of Lodgepole Pine

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

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

needles of lodgpole pine (Pinus contorta)

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

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

closed cone of lodgepole pine (Pinus contorta)

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

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

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

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

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

open cones of lodgepole pine (Pinus contorta)

Further Reading and Viewing:

The Flight of the Dandelion

The common dandelion (Taraxacum officinale) comes with a collection of traits that make it a very successful weed. Nearly everything about it screams success, from its asexually produced seeds to its ability to resprout from a root fragment. Evolution has been kind to this plant, and up until the recent chemical warfare we’ve subjected it to, humans have treated it pretty well too (both intentionally and unintentionally).

One feature that has served the dandelion particularly well is its wind-dispersed seeds. Dandelions have a highly-evolved pappus – a parachute-like bristle of hairs attached to its fruit by a thin stalk. The slightest breath or puff of wind will send this apparatus flying. Once airborne, a dandelion’s seed can travel up to a kilometer or more away from its mother plant, thereby expanding its territory with ease.

Such a low-growing plant achieving this kind of distance is impressive. Even more impressive is that it manages to do this with a pappus that is 90% empty space. Would you leap from a plane with only 10% of a parachute?

Dandelion flight was investigated by researchers at the University of Edinburgh, who used a wind tunnel along with long-exposure photography and high-speed imaging to observe the floating pappus. Their research was presented in a letter published in an issue of Nature in October 2018. Upon close examination, they observed a stable air bubble floating above the pappus as it flew. This ring-shaped air bubble – or vortex – which is unattached to the pappus is known as a separated vortex ring. While this type of vortex ring had been considered theoretically, this marked the first time one had been observed in nature.

Seeing this type of air bubble associated with the dandelion’s pappus intrigued the researchers. About a 100 filaments make up the parachute portion of the pappus. They are arranged around the stalk, leaving heaps of blank space in between. The air bubble observed was not what was expected for such a porous object. However, the researchers found that the filaments were interacting with each other in flight, reducing the porosity of the pappus. In their words, “Neighboring filaments interact strongly with one another because of the thick boundary layer around each filament, which causes a considerable reduction in air flow through the pappus.”

The pappus acts as a circular disk even though it is not one, and its limited porosity allows just enough air movement through the filaments that it maintains this unique vortex. “This suggests,” the researchers write, “that evolution has tuned the pappus porosity to eliminate vortex shedding as the seed flies.” Fine-tuned porosity and the resultant unattached air bubble stabilizes the floating fruit “into an equilibrium orientation that minimizes [its] terminal velocity, allowing [it] to make maximal use of updrafts.” The result is stable, long distance flight.

Wind-dispersed seeds come in two main forms: winged and plumed. Winged seeds are common in trees and large shrubs. They benefit from the height of the tree which allows them to attain stable flight. While such seeds have the ability to travel long distances, their success is limited on shorter plants. In this case, plumed seeds, like those of the dandelion, are the way to go. As the researchers demonstrated, successful flight can be achieved by bristles in place of wings. The tiny seeds of dandelions seen floating by on a summer breeze are not tumbling through the air haphazardly; rather, they are flying steadily, on their way to spoil the dreams of a perfect lawn.

Further Reading (and Watching):

Investigating the Soil Seed Bank

Near the top of the world, deep inside a snow-covered mountain located on a Norwegian island, a vault houses nearly a million packets of seeds sent in from around the world. The purpose of the Svalbard Global Seed Vault is to maintain collections of crop seeds to ensure that these important species and varieties are not lost to neglect or catastrophe. In this way, our food supply is made more secure, buffered against the unpredictability of the future. Seed banks like this can be found around the world and are essential resources for plant conservation. While some, like Svalbard, are in the business of preserving crop species, others, like the Millennium Seed Bank, are focused on preserving seeds of plants found in the wild.

Svalbard Global Seed Vault via wikimedida commons

Outside of human-built seed banks, many plants maintain their own seed banks in the soil where they grow. This is the soil seed bank, a term that refers to either a collection of seeds from numerous plant species or, simply, the seeds of a single species. All seed bearing plants pass through a period as a seed waiting for the chance to germinate. Some do this quickly, as soon as the opportunity arises, while others wait, sometimes for many years, before germinating. Plants whose seeds germinate quickly, generally do not maintain a seed bank. However, seeds that don’t germinate right away and become incorporated in the soil make up what is known as a persistent soil seed bank.

A seed is a tiny plant encased in a protective layer. Germination is not the birth of a plant; rather, the plant was born when the seed was formed. The dispersal of seeds is both a spatial and temporal phenomenon. First the seed gets to where it’s going via wind, water, gravity, animal assistance, or some other means. Then it waits for a good opportunity to sprout. A seed lying in wait in the soil seed bank is an example of dispersal through time. Years can pass before the seed germinates, and when it does, the plant joins the above ground plant community.

Because seeds are living plants, seeds found in the soil seed bank are members of a plant community, even though they are virtually invisible and hard to account for. Often, the above ground plant community does not represent the population of seeds found in the soil below. Conversely, seeds in a seed bank may not be representative of the plants growing above them. This is because, as mentioned earlier, not all plant species maintain soil seed banks, and those that do have differences in how long their seeds remain viable. Depending on which stage of ecological succession the plant community is in, the collection of seeds below and the plants growing above can look quite different.

Soil seed banks are difficult to study. The only way to know what is truly there is to dig up the soil and either extract all the seeds or encourage them to germinate. Thanks to ecologists like Ken Thompson, who have studied seed banks extensively for many years, there is still a lot we can say about them. First, for the seeds of a plant to persist in the soil, they must become incorporated. Few seeds can bury themselves, so those with traits that make it easy for them to slip down through the soil will have a greater chance of being buried. Thompson’s studies have shown that “persistent seeds tend to be small and compact, while short-lived seeds are normally larger and either flattened or elongate.” Persistent seeds generally weigh less than 3 milligrams and tend to lack appendages like awns that can prevent them from working their way into the soil.

The seeds of moth mullein (Verbascum blattaria) are tiny and compact and known to persist in the soil for decades as revealed in Dr. Beal’s seed viability experiment. (photo credit: wikimedia commons)

Slipping into cracks in the soil is a major way seeds move through the soil profile, but it isn’t the only way. In a study published in New Phytologist, Thompson suggests that “the association between small seeds and possession of a seed bank owes much to the activities of earthworms,” who ingest seeds at the surface and deposit them underground. Later, they may even bring them back up the same way. Ants also play a role in seed burial, as well as humans and their various activities. Some seeds, like those of Avena fatua and Erodium spp., have specialized appendages that actually help work the seeds into the soil.

Not remaining on the soil surface keeps seeds from either germinating, being eaten, or being transported away to another site. Avoiding these things, they become part of the soil seed bank. But burial is only part of the story. In an article published in Functional Ecology, Thompson et al. state that burial is “an essential prelude to persistence,” but other factors like “germination requirements, dormancy mechanisms, and resistance to pathogens also contribute to persistence.” If a buried seed rots away or germinates too early, its days as a member of the soil seed bank are cut short.

The seeds of redstem filare (Erodium circutarium) have long awns that start out straight, then coil up, straighten out, and coil up again with changes in humidity. This action helps drill the seeds into the soil. (photo credit: wikimedia commons)

Soil seed banks can be found wherever plants are found – from natural areas to agricultural fields, and even in our own backyards. Thompson and others carried out a study of the soil seed banks of backyard gardens in Sheffield, UK. They collected 6 soil cores each (down to 10 centimeters deep) from 56 different gardens, and grew out the seeds found in each core to identify them. Most of the seeds recovered were from species known to have persistent seed banks, and to no surprise, the seed banks were dominated by short-lived, weedy species. The seeds were also found to be fairly evenly distributed throughout the soil cores. On this note, Thompson et al. remarked that due to “the highly disturbed nature of most gardens, regular cultivation probably ensures that seeds rapidly become distributed throughout the top 10 centimeters of soil.”

Like the seed banks we build to preserve plant species for the future, soil seed banks are an essential long-term survival strategy for many plant species. They are also an important consideration when it comes to managing weeds, which is something we will get into in a future post.

Dr. Beal’s Seed Viability Experiment

In 1879, Dr. William J. Beal buried 20 jars full of sand and seeds on the grounds of Michigan State University. He was hoping to answer questions about seed dormancy and long-term seed viability. Farmers and gardeners have often wondered: “How many years would one have to spend weeding until there are no more weeds left to pull?” Seeds only remain viable for so long, so if weeds were removed before having a chance to make more seeds, the seed bank could, theoretically, be depleted over time. This ignores, of course, the consistent and persistent introduction of weed seeds from elsewhere, but that’s beside the point. The question is still worth asking, and the study still worth doing.

When Dr. Beal set up the experiment, he expected it would last about 100 years, as one jar would be tested every 5 years. However, things changed, and Dr. Beal’s study is now in its 140th year, making it the longest-running scientific experiment to date. If things go as planned, the study will continue until at least 2100. That’s because 40 years into the study, a jar had to be extracted in the spring instead of the fall, as had been done previously, and at that point it was decided to test the remaining jars at 10 year intervals. In 1990, things changed again when the period was extended to 20 years between jars. The 15th jar was tested in 2000, which means the next test will occur in the spring of next year.

In preparing the study, Dr. Beal filled each of the 20 narrow-necked pint jars with a mixture of moist sand and 50 seeds each of 21 plant species. All but one of the species (Thuja occidentalis) were common weeds. He buried the jars upside down – “so that water would not accumulate about the seeds” – about 20 inches below ground. Near each bottle he also buried seeds of red oak and black walnut, but they all rotted away early in the study.

After the retrieval of each bottle, the sand and seed mixture is dumped into trays and exposed to conditions suitable for germination. The number of germinates are then counted and recorded. Over the years, the majority of the seeds have lost their viability. In 2000, only three species germinated  – Verbascum blattaria, a Verbascum hybrid, and Malva rotundifolia. There were only two individuals of the Verbascum hybrid, and only one Malva rotundifolia. The seeds of Verbascum blattaria, however, produced 23 individuals, suggesting that even after 120 years, the seeds of this species could potentially remain viable long into the future.

moth mullein (Verbascum blattaria)

In the 2000 test, the single seedling of Malva rotundifolia germinated after a cold treatment. Had the cold treatment not been tried, germination may not have occurred, which begs the question, how many seeds in previous studies would have germinated if subjected to additional treatments? Dr. Beal himself had wondered this, expressing that the results he had seen were “indefinite and far from satisfactory.” He admitted that he had “never felt certain that [he] had induced all sound seeds to germinate.”

There are also some questions about the seeds themselves. For example, the authors of the 2000 report speculate that poor germination seen in Malva rotundifolia over most of the study period could be “the result of poor seed set rather than loss of long-term viability.” The presence of a Verbascum hybrid also calls into question the original source of those particular seeds. A report published in 1922 questions whether or not the seeds of Thuja occidentalis were ever actually added to the jars, and also expresses uncertainty about the identify of a couple other species in the study.

Despite these minor issues, Dr. Beal’s study has shed a great deal of light on questions of seed dormancy and long-term seed viability and has inspired numerous related studies. While questions about weeds were the inspiration for the study, the things we have been able to learn about seed banks has implications beyond agriculture. Seed bank dynamics are particularly important in conservation and restoration. If plants that have disappeared due to human activity have maintained a seed bank in the soil, there is potential for the original population to be restored.

In future posts we will dive deeper into seed banks, seed dormancy, and germination. In the meantime, you can read more about Dr. Beal’s seed viability study by visiting the following links:

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.

Highlights from the Western Society of Weed Science Annual Meeting

Earlier this month, I went to Garden Grove, California to attend the 71st annual meeting of the Western Society of Weed Science. My trip was funded by an Education and Enrichment Award presented by the Pahove Chapter of the Idaho Native Plant Society. It was a great opportunity for a weeds-obsessed plant geek like myself to hang out with a bunch of weed scientists and learn about their latest research. What follows are a few highlights and takeaways from the meeting.

General Session

Apart from opening remarks and news/business-y stuff, the general session featured two invited speakers: soil ecologist Lydia Jennings and historian David Marley. Lydia’s talk was titled “Land Acknowledgement and Indigenous Knowledge in Science.” She started by sharing a website called Native Land, which features an image of the Earth overlayed with known “borders” of indigenous territories. By entering your address, you can see a list of the tribes that historically used the land you now inhabit. It is important for us to consider the history of the land we currently live and work on. Lydia then compared aspects of western science and indigenous science, pointing out ways they differ as well as ways they can be used in tandem. By collaborating with tribal nations, weed scientists can benefit from traditional ecological knowledge. Such knowledge, which has historically gone largely unrecognized in the scientific community, should receive more attention and acknowledgement.

David Marley was the comic relief. Well-versed in the history of Disneyland, he humorously presented a series of stories involving its creation. Little of what he had to say related to weed science, which he openly admitted along the way; however, one weeds related story stood out. Due to a lack of funds, the early years of Tomorrowland featured few landscape plants. To make up for that, Walt Disney had signs with fake Latin names created for some of the weeds.

Weeds of Range and Natural Areas

I spent the last half of the first day in the “Weeds of Range and Natural Areas” session where I learned about herbicide ballistic technology (i.e. killing plants from a helicopter with a paintball gun loaded with herbicide). This is one of the ways that Miconia calvescens invasions in Hawaii are being addressed. I also learned about research involving plant debris left over after logging. When heavy amounts of debris are left in place, scotch broom (Cytisus scoparius) infestations are thwarted. There was also a talk about controlling escaped garden loosestrife (Lysimachia punctata) populations in the Seattle area, as well as a few talks about efforts to control annual grasses like cheatgrass (Bromus tectorum) in sagebrush steppes. Clearly there are lots of weed issues in natural areas, as that only covers about half the talks.

Basic Biology and Ecology

On the morning of the second day, the “Basic Biology and Ecology” session held a discussion about weeds and climate change. As climate changes, weeds will adapt and find new locations to invade. Perhaps some weeds won’t be as problematic in certain areas, but other species are sure to take their place. Understanding the changes that are afoot and the ways that weeds will respond to them is paramount to successful weed management. This means documenting the traits of every weed species, including variations between and among populations of each species, so that predictions can be made about their behavior. It also means anticipating new weed species and determining ways in which weeds might exploit new conditions.

No doubt there is much to learn in order to adequately manage weeds in a changing climate. An idea brought up during the discussion that I was particularly intrigued by was using citizen scientists to help gather data about weeds. Similar to other organizations that collect phenological data from the public on a variety of species, a website could be set up for citizen scientists to report information about weeds in their area, perhaps something like this project in New Zealand. Of course, there are already a series of apps available in North America for citizen scientists to report invasive species sightings, so it seems this is already happening to some degree.

Teaching and Technology Transfer

A highlight of the afternoon’s “Teaching and Technology Transfer” session was learning about the Wyoming Restoration Challenge hosted by University of Wyoming Extension. This was a three year long contest in which thirteen teams were given a quarter-acre plot dominated by cheatgrass with the challenge to restore the plant community to a more productive and diverse state. Each team developed and carried out their own strategy and in the end were judged on a series of criteria including cheatgrass and other weed control, plant diversity, forage production, education and outreach, and scalability. Preliminary results can be seen here; read more about the challenge here and here.

And so much more…

Because multiple sessions were held simultaneously, I was unable to attend every talk. I also had to leave early on the third day, so I missed those talks as well. However, I did get a chance to sit in on a discussion about an increasingly troubling topic, herbicide-resistant weeds, which included a summary of regional listening sessions that have been taking place in order to bring more attention to the subject and establish a dialog with those most affected by it.

One final highlight was getting to meet up with Heather Olsen and talk to her briefly about her work in updating the Noxious Weed Field Guide for Utah. This work was aided by the Invasive Plant Inventory and Early Detection Prioritization Tool, which is something I hope to explore further.

If you are at all interested in weeds of the western states, the Western Society of Weed Science is a group you should meet. They are fun and friendly people who really know their weeds.

See Also: Highlights from the Alaska Invasive Species Workshop 

Lettuce Gone Wild, part two

The lettuce we eat is a close relative to the lettuce we weed out of our gardens. Last week we discussed the potential that wild relatives may have for improving cultivated lettuce. But if wild lettuce can be crossed with cultivated lettuce to create new cultivars, can cultivated lettuce cross with wild lettuce to make it more weedy?

Because so many of our crops are closely related to some of the weeds found along with them or the plants growing in nearby natural areas, the creation of crop-wild hybrids has long been a concern. This concern is heightened in the age of transgenic crops (also known as GMOs), for fear that hybrids between weeds and such crops could create super weeds – fast spreading or highly adapted weeds resistant to traditional control methods such as certain herbicides. To reduce this risk, extensive research is necessary before such crops are released for commercial use.

flowers of prickly lettuce (Lactuca serriola)

There are no commercially available, genetically modified varieties of cultivated lettuce, so this is not a concern when it comes to crop-wild hybrids; however, due to how prevalent weedy species like prickly lettuce (Lactuca serriola) are, hybridization with cultivated lettuce is still a concern. So, it is important to understand what the consequences might be when hybridization occurs.

In a paper published in Journal of Applied Ecology in 2005, Hooftman et al. examined a group of second-generation hybrids (L. sativa x L. serriola), and found that the hybrids behaved and appeared very similarly to non-hybrid prickly lettuce. They also found that the seeds produced by the hybrids had a significantly higher germination rate than non-hybrid plants. This is an example of hybrid vigor. Thus, “if hybridization does occur, this could lead to better performing and thus potentially more invasive (hybrid) genotypes.” However, the authors cautioned that “better performing genotypes do not automatically result in higher invasiveness,” and that much depends on the conditions they are found in, the level of human disturbance, etc.

Another thing to consider is that hybrids are not stable. In an article published in Nature Reviews Genetics in 2003, Stewart et al. adress the “misunderstanding that can arise through the confusion of hybridization and … introgression.” It is wrong to assume that hybrids between crops and wild relatives will automatically lead to super weeds. For this to occur, repeated crosses with parental lines (also known as backcrossing) must occur, and “backcross generations to the wild relative must progress to the point at which the transgene [or other gene(s) in question] is incorporated into the genome of the wild relative.” That is what is meant by “introgression.” This may happen quickly or over many generations or it may never happen at all. Each case is different.

prickly leaf of prickly lettuce (Lactuca serriola)

In a paper published in Journal of Applied Ecology in 2007, Hooftman et al. observe the breakdown of crop-wild lettuce hybrids. They note that “fitness surplus through [hybrid vigor] will often be reduced over few generations,” which is what was seen in the hybrids they observed. One possible reason why this occurs is that lettuce is predominantly a self-crossing species; outcrossing is rare, occurring 1 – 5% of the time thanks to pollinating insects. But that doesn’t mean that a stable, aggressive genotype could never develop. Again, much depends on environmental conditions, as well as rates of outcrossing and other factors relating to population dynamics.

A significant expansion of prickly lettuce across parts of Europe led some to hypothesize that crop-wild hybrids were partly to blame. In a paper published in Molecular Ecology in 2012 Uwimana et al. ran population genetic analyses on extensive data sets to determine the role that hybridization had in the expansion. They concluded that, at a level of only 7% in wild habitats, crop-wild hybrids were not having a significant impact. They observed greater fitness in the hybrids, as has been observed in other studies (including the one above), but they acknowledged the instability of hybrids, especially in self-pollinating annuals like lettuce.

seed head of prickly lettuce (Lactuca serriola)

It is more likely that the expansion of prickly lettuce in Europe is due to “the expansion of favorable habitat as a result of climate warming and anthropogenic habitat disturbance and to seed dispersal because of transportation of goods.” Uwimana et al. did warn, however, that “the occurrence of 7% crop-wild hybrids among natural L. serriola populations is relatively high [for a predominantly self-pollinating species] and reveals a potential [for] transgene movement from crop to wild relatives [in] self-pollinating crops.”