Summer of Weeds: Common Mullein

The fuzzy, gray-green leaves of common mullein are familiar and friendly enough that it can be hard to think of this plant as a weed. Verbascum thapsus is a member of the figwort family and is known by dozens of common names, including great mullein, Aaron’s rod, candlewick, velvet dock, blanket leaf, feltwort, and flannel plant. Its woolly leaves are warm and inviting and have a history of being used as added padding and insulation, tucked inside of clothing and shoes. In Wild Edible and Useful Plants of Idaho, Ray Vizgirdas writes, “the dried stalks are ideal for use as hand-drills to start fires; the flowers and leaves produce yellow dye; as a toilet paper substitute, the large fresh leaves are choice.”

Common mullein is a biennial that was introduced to eastern North America from Eurasia in the 1700’s as a medicinal plant and fish poison. By the late 1800’s it had reached the other side of the continent. In its first year it forms a rosette of woolly, oblong and/or lance-shaped leaves. After overwintering it produces a single flower stalk up to 6 feet tall. The woolly leaves continue along the flower stalk, gradually getting smaller in size until they reach the inflorescence, which is a long, dense, cylindrical spike. Sometimes the stalk branches out to form multiple inflorescences.

First year seedlings of common mullein (Verbascum thapsus)

The inflorescence doesn’t flower all at once; instead, a handful of flowers open at a time starting at the bottom of the spike and moving up in an irregular pattern. The process takes several weeks to complete. The flowers are about an inch wide and sulfur yellow with five petals. They have both female and male sex parts but are protogynous, meaning the female organs mature before the male organs. This encourages cross-pollination by insects. However, if pollination isn’t successful by the end of the day, the flowers self-pollinate as the petals close. Each flower produces a capsule full of a few hundred seeds, and each plant can produce up to 180,000 seeds. The seeds can remain viable for over 100 years, sitting in the soil waiting for just the right moment to sprout.

Common mullein is a friend of bare, recently disturbed soil. It is rare to see this plant growing in thickly vegetated areas. As an early successional plant, its populations can be abundant immediately after a disturbance, but they do not persist once other plants have filled in the gaps. Instead they wait in seed form for the next disturbance that will give them the opportunity to rise again. They can be a pest in gardens and farm fields due to regular soil disturbance, and are often abundant in pastures and rangelands because livestock avoid eating their hairy leaves. Because of its ephemeral nature, it is generally not considered a major weed; however, it is on Colorado’s noxious weed list.

Several features make common mullein a great example of a drought-adapted plant. Its fleshy, branching taproot can reach deep into the soil to find moisture, the thick hairs on the leaves help reduce water loss via transpiration, and the way the leaves are arranged and angled on the stalk can help direct rain water down toward the roots.

Common mullein has an extensive history of ethnobotanical uses. Medicinally it has been used internally to treat coughs, colds, asthma, bronchitis, and kidney infections; and as a poultice to treat warts, slivers, and swelling. The dried flower stalks have been used to make torches, and the fuzzy leaves have been used as tinder for fire-making and wicks in lamps.

The hairy leafscape of common mullein (Verbascum thapsus)

More Resources:

Quote of the Week:

From Gaia’s Garden by Toby Hemenway

Here’s why opportunistic plants are so successful. When we clear land or carve a forest into fragments, we’re creating lots of open niches. All that sunny space and bare soil is just crying out to be colongized by light- and fertlity-absorbing green matter. Nature will quickly conjure up as much biomass as possible to capture the bounty, by seeding low-growing ‘weeds’ into a clearing or, better yet, sprouting a tall thicket stretching into all three dimensions to more effectively absorb light and develop deep roots. … When humans make a clearing, nature leaps in, working furiously to rebuild an intact humus and fungal layer, harvest energy, and reconstruct all the cycles and connections that have been severed. A thicket of fast-growing pioneer plants, packing a lot of biomass into a small space, is a very effective way to do this. … And [nature] doesn’t care if a nitrogen fixer or a soil-stabilizing plant arrived via continental drift or a bulldozer’s treads, as long as it can quickly stitch a functioning ecosystem together.

Book Review: Weeds Find a Way

At what age do we become aware that there are profound differences among the plants we see around us? That some are considered good and others evil. Or that one plant belongs here and another doesn’t. Most young children (unless an adult has taught them) are unaware that there is a difference between a weed and a desirable plant. If it has attractive features or something fun to interact with – like the seed heads of dandelions or the sticky leaves of bedstraw – they are all the same. At some point in our trajectory we learn that some plants must be rooted out, while others can stay. Some plants are uninvited guests – despite how pretty they might be – while others are welcome and encouraged.

But weeds are resilient, and so they remain. Weeds Find a Way, written by Cindy Jenson-Elliot and illustrated by Carolyn Fisher, is a celebration of weeds for their resiliency as well as for their beauty and usefulness. This book introduces the idea of weeds to children, focusing mainly on their tenacity, resourcefulness, and positive attributes rather than their darker side. “Weeds are here to stay,” so perhaps there is a place for them.

The book begins by listing some of the “wondrous ways” that weed seeds disperse themselves: “floating away on the wind,” attaching themselves to “socks and fur,” shot “like confetti from a popped balloon.” And then they wait – under snow and ice or on top of hot sidewalks – until they find themselves in a time and place where they can sprout. Eventually, “weeds find a way to grow.”

Weeds also “find a way to stay.” We can pull them up, but their roots are often left behind to “sprout again.” Pieces and parts break off and take root in the soil. Animals may swoop in to devour them, but weeds drive them away with their thorns, prickles, and toxic chemicals. In these ways they are a nuisance, but they can be beautiful and beneficial, too.

This illustrated story of weeds is followed by some additional information, as well as a list of common weeds with brief descriptions. Weeds are defined as plants “thought to be of no value that grow in places where people do not want them to grow,” adding that even “misunderstood and underappreciated plants that are native to a region and have multiple uses” can be labeled weeds.

The concept of weeds as invasive species is also addressed; some introduced plants move into natural areas and can “crowd out native vegetation, block streams, and drive away wild animals.” That being said, weeds also provide us with “endless opportunities to study one of nature’s most wonderful tools: adaptation.” Weeds are problematic as much as they are useful, it’s simply a matter of perspective.

A criticism of this book might be that it doesn’t focus enough on the negative aspects of weeds. There is plenty of that elsewhere. The aim of this book is to connect us with nature, and as Jensen-Elliot writes, “you don’t need a garden to know that nature is at work.” When there is a weed nearby, nature is nearby. Weeds “adapt and grow in tough times and desolate places,” and they make the world beautiful “one blossom at a time.”

Summer of Weeds: Salsify

Picking a favorite weed is challenging. If we dismiss entirely the idea that a person is not supposed to like weeds, the challenge is not that “favorite weeds” is an oxymoron; it is, instead, that it is impossibile to pick one weed among hundreds of weeds that is the most attractive, the most impressive, the most useful, the most forgiving, whatever. For me, salsify is a top contender.

Salsify and goatsbeard are two of several common names for plants in the genus Tragopogon. At least three species in this genus have been introduced to North America from Europe and Asia. All are now common weeds, widespread across the continent. All have, at some point, been cultivated intentionally for their edible roots and leaves, but Tragopogon porrifolius – commonly known as oyster plant or purple salsify – may be the only one that is intentionally grown in gardens today. Its purple flowers make it easy to determine between the other two species, which have yellow flowers.

As it turns out, I am not familiar with purple salsify. I don’t think it is as common in western North America as it is in other parts of the continent. In fact, the most common of the three in my corner of the world appears to be Tragopogon dubius, commonly known as western salsify. Tragopogon pratensis (meadow salsify) makes an appearance, but perhaps not as frequently. To complicate matters, hybridization is common in the genus, so it may be difficult to tell exactly what you are looking at.

western salsify (Tragopogon dubius)

Regardless, salsify is a fairly easy weed to identify. It is a biennial (sometimes annual, sometimes perennial) plant that starts out as a rosette of gray-green leaves that are grass-like in appearance. Eventually a flower stalk emerges, adorned with more grass-like leaves, branching out to form around a half dozen flower heads. Salsify is in the aster family, in which flower heads typically consist of a tight grouping of disc and ray florets. In this case, only ray florets are produced. The florets are yellow or lemon-yellow, and each flower head sits atop a series of pointed bracts which encase the flower (and the forming seed head) when closed. Examining the length of the bracts is one way to tell T. dubius (bracts extend beyond the petals) from T. pratensis (bracts and petals are equal in length).

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

The flowers of salsify open early in the morning and face the rising sun. By noon, they have usually closed. This phenomenon is the reason behind other common names like noonflower and Jack-go-to-bed-at-noon. Salsify’s timely flowering makes an appearance in Elizabeth Gilbert’s novel, The Signature of All Things: “Alma learned to tell time by the opening and closing of flowers. At five 0’clock in the morning, she noticed, the goatsbeard petals always unfolded. … At noon, the goatsbeard closed.”

The seed heads of salsify look like over-sized dandelions. Each seed (a.k.a. achene) is equipped with a formidable pappus, and with the help of a gust of wind, seeds can be dispersed hundreds of feet from the parent plant. The seeds don’t remain viable for very long, but with each plant producing a few hundred seeds and dispersing them far and wide, it is not hard to see why salsify has staying power.

Open, sunny areas are preferred by salsify, but it can grow in a variety of conditions. In The Book of Field and Roadside, John Eastman writes, “goatsbeards can establish themselves in bare soil, amid grasses and old-field vegetation, and in heavy ground litter; such adaptability permits them to thrive across a range of early plant successional stages.” Wild Urban Plants of the Northeast lists the following sites as “habitat prefrences” of meadow salsify: “abandoned grasslands, urban meadows, vacant lots, rubble dumps, and at the base of rock outcrops and stone walls.” While generally not considered a noxious weed, Tragopogon species are commonly encountered and widely naturalized. Last summer on a field trip to Mud Springs Ridge near Hells Canyon, salsify was one of only a small handful of introduced plants I observed looking right at home with the native flora.

Seed heads of western salsify (Tragopogon dubius) before opening

All that being said, why is salsify one of my favorite weeds? Its simple and elegant form appeals to me. Its gray- or blue-green stems and leaves together with its unique, yellow flowers are particularly attractive to me. And its giant, globe-shaped seed heads, which seem to glisten in the sun, captivate me. Its not a difficult weed to get rid off. It generally pulls out pretty easily, and it’s a satisfying feeling when you can get it by the root. It’s a sneaky weed, popping up full grown inside of another plant and towering above it, making you wonder how you could have missed such an intrusion. The roots are said to be the most palatable before the plant flowers, so if you can spot the young rosette – disguised as grass and also edible – consider yourself lucky. I haven’t tried them yet, but I will. [Editor’s note: Sierra tells me that I have eaten them in a salad she made, but at the time I didn’t know they were in there so I don’t remember what they tasted like.] If they are any good, that will be one more reason why salsify is one of my favorite weeds.

Bonus excerpt from Emma Cooper’s book, Jade Pearls and Alien Eyeballs, regarding Tragopogon porrifolius:

Salsify is often called the vegetable oyster, because its roots are supposed to have an oyster-like flavor although I suspect nobody would be fooled. The long roots are pale and a bit like carrots – they are mild and sweet and when young can be eaten raw. Mature roots are better cooked. Traditionally a winter food, any roots left in the ground in spring will produce a flush of edible foliage.

Introducing the Summer of Weeds

I spent the first five months of this year posting almost exclusively about invasive species. There is still plenty more to say on the topic, and I’m sure I will get back to that. However, it is time now to dive into the topic that I really want to explore. Weeds.

There is definitely crossover between the two topics – many weeds are invasive species – but there are clear distinctions, too. Oftentimes, weeds as a category of plants are unfairly and unjustly lumped under the title “invasive,” but any plant can be a weed at any moment in time if a human says so. That’s the difference. A plant does not have to prove that it is causing any sort of ecological or economic damage to be called a “weed;” it just has to be growing where a human doesn’t want it to. Yet, too quickly a plant “out of place” is cursed at using words like “invasive” or “noxious” regardless of its origin or behavior. I know I’m being overly semantic about this, but it seems unfair (and incorrect) to lump any and all plants that are bothering us for whatever reason into categories that have legal definitions.

If you can’t already tell, I am obsessed with weeds. It’s a topic I have been thinking about fairly consistently for much of my adult life. For one thing, as part of my career I spend a huge portion of my time killing and controlling weeds. I comprehend fully the visceral reaction of seeing a garden overcome by weeds – the vile thoughts one can have towards a group of plants that are soiling what could otherwise be a beautiful landscape – and I know very well the backbreaking work and countless hours that go into removing uninvited plants (cursing the intruders along the way). I get why weeds are a problem, and I understand why they are a subject of so much vitriol. Yet, over the years I have developed a respect – even a love – for weeds (despite the fact that I still must remove them and that removing them continues to be an overwhelming task).

Unwanted plants have been following us around and getting in our way for millenia. Essentially, we are partners in crime. We intentionally and unintentionally bring plants from various parts of the world on our travels, and through disturbance we create conditions where introduced plants can settle in and thrive. Over time, some once beloved plants grow out of favor and transition from desirable to weedy. As our cycles of disturbance continue, we give early successional, opportunistic plant species a chance to perpetuate themselves, guaranteeing that we will keep such “weeds” with us forever. We reap what we sow; even though we generally don’t plant weeds on purpose, other actions ensure that they will be our constant companions.

The importance of weed control goes beyond the aesthetic. In horticulture and agriculture, weeds compete with crops for light, space, water, and nutrients. They also harbor pests and diseases, and their seeds can contaminate crops. In pastures and rangelands, some weeds poison livestock. Certain weeds are harmful to people, too. Other weeds are simply disruptive – getting tangled up in machinery, damaging infrastructure, blocking our vision along roadways, and even giving cyclists flat tires. Apart from all that, even if all weeds did was make our gardens look unsightly, I imagine we would still be pretty angry with them.

I am interested in weeds wherever they are, but the weeds that fascinate me the most are those that thrive in urban environments. Not necessarily the weeds in our yards, but the weeds that have escaped our fences and property lines; the ones in the margins. We see them in abandoned lots, along roadways, near irrigation channels, and in other neglected spaces. They pop up in the cracks of sidewalks, on rooftops, in the middle of decaying buildings, and anywhere else that people haven’t paid attention to in a while. Urban areas have, for the most part, been scraped of their native flora. Introduced species move in to fill that void. As Richard Mabey writes in his book about weeds, these plants “insinuate the idea of wild nature into places otherwise quite shorn of it;” they are “the very essence of wildness.” Novel ecosystems, like those created by urbanization and human development, are with us whether we like it or not. There is a “wildness” to them that is unlike other cultivated and manicured areas maintained by humans. These urban wild places are worth a closer look.

So, what is the Summer of Weeds?

Put simply, it’s an exploration of weeds. Throughout the summer I will be profiling some of the weeds I come across in my daily life. I will include photos, a brief description, and some interesting facts about each species. I will also include quotes about weeds from various sources, as well as videos, links, resources, and whatever else I come across that seems worth sharing. I hope you enjoy it. If you have anything to add along the way – specifically any personal thoughts or stories to share about weeds – please do. You can contact me via the usual ways: in the comment section below, through the Contact page, or on Twitter, Tumblr, Facebook, or Instagram. Happy Summer!

Love and Hate – The Story of Purple Loosestrife

In the early 1800’s, seeds of purple loosestrife found their way to North America. They arrived from Europe several times by various means – accidentally embedded in the ballast of ships, inadvertently tucked in sheep’s wool, and purposely carried in the hands of humans. Native to much of Europe and parts of Asia and commonly found growing in wetlands and other riparian areas, purple loosestrife’s appealing spikes of magenta flowers, sturdy, upright growth habit, and ease of propagation made it a prized ornamental; its abundant nectar made it a favorite of beekeepers.

During its first 150 years or so in North America, purple loosestrife became naturalized in ditches, wet meadows, and the banks of streams, rivers, lakes, and ponds while also enjoying a place in our gardens. Concern about its spread was raised in the first half of the twentieth century, but it wasn’t until the 1980’s after an extensive survey was done and a special report was issued by the U.S. Fish and Wildlife Service that attitudes about purple loosestrife shifted dramatically. At that point, it was no longer a benign invader and welcome garden companion. It was, instead, a biological menace that needed to be destroyed.

Lytrhrum salicaria – commonly known as purple loosestrife, spiked willow-herb, long purples, rainbow weed, etc. – is an herbaceous perennial in the family Lythraceae. It reaches up to two meters tall; has square or angular stems with lance-shaped, stalkless leaves up to ten centimeters long; and ends in dense, towering spikes of pink-purple, 5-7 petaled flowers. The flowers attract a wide variety of pollinating insects – mostly bees – and afterwards produce small capsules full of tiny, red-brown seeds. Charles Darwin thoroughly studied the flowers of purple loosestrife; he was intrigued by the plant for many reasons, including its heterostyly (a topic for another post).

Lythrum salicaria (purple loosestrife) – image credit: wikimedia commons

Purple loosestrife seeds remain viable for up to 20 years and are transported by wind, water, and in mud stuck to the feet of birds. Apart from seeds, populations expand clonally as root crowns grow larger each year and produce increasingly more stems. Broken stem pieces also take root in mud, creating new plants. Purple loosestrife’s ability to form expansive populations in a quick manner, pushing other plants aside and forming what appears to be a dense monoculture, is part of the reason it has earned itself a place among the International Union for Conservation of Nature’s list of 100 World’s Worst Invasive Alien Species.

But is this ranking justified? In a paper published in Biological Invasions in 2010, Claude Lavoie compares news reports about purple loosestrife around the turn of the century with data presented in scientific papers and finds that the reports largely exaggerate the evidence. Purple loosestrife was being accused of all manner of crimes against nature and was being condemned before there was sound evidence to justify such actions.

It began with the U.S. Fish and Wildlife Service’s special report published in 1987. According to Lavoie, “a long list of the impacts of the species on wetland flora and fauna [was] presented,” but the claims were not supported by observational or experimental data – “the impacts [were] only suspected.” Regardless, wetland managers began campaigns against purple loosestrife in order to convince the public that it was a Beautiful Killer. News outlets were quick to spread the word about this “killer” plant. When biological control programs began in the 1990’s, news outlets reported on their success. Little empirical evidence had been published on either topic, and debates about purple loosestrife’s impacts remained unsettled in the scientific community.

The flowers of purple loosestrife (Lythrum salicaria) – photo credit: wikimedia commons

Around this time, five reviews were published examining the evidence against purple loosestrife. Lavoie reports that all but one of them “rely on a relatively high number of sources that have not been published in peer-reviewed journals.” After examining the reviews, Lavoie concludes: “although each review provided valuable information on purple loosestrife, most were somewhat biased and relied on a substantial amount of information that was anecdotal or not screened by reviewers during a formal evaluation process. Only one review was impartial, and this one painted an inconclusive picture of the species.”

Research has continued regarding the impacts of purple loosestrife, and so Lavoie examined 34 studies that were published during the 2000’s in search of conclusive evidence that the plant is as destructive to wetlands and wildlife as has been claimed. Upon examination he concludes that “stating that this plant has ‘large negative impacts’ on wetlands is probably exaggerated.” The most common accusation – that purple loosestrife crowds out native plants and forms a monoculture – “is controversial and has not been observed in nature (with maybe one exception).” Lavoie finds that there is “certainly no evidence that purple loosestrife ‘kills wetlands’ or ‘creates biological deserts,'” and “there are no published studies [in peer-reviewed journals] demonstrating that purple loosestrife has an impact on waterfowl or fishes.” All other negative claims against purple loosestrife “have not been the object of a study,” except for its impact on amphibians, which had at that time only been tested on two species, one “reacting negatively.” Certain claims – such as purple loosestrife’s impact on wetland hydrology – should be studied more in depth “considering the apparent public consensus on the detrimental effects of purple loosestrife” on wetland ecosystems.

Lavoie agrees that it is reasonable to control purple loosestrife when the intention is to reduce additional pressures on an ecosystem that is already highly threatened. However, he warns that “focusing on purple loosestrife instead of on other invasive species or on wetland losses to agriculture or urban sprawl could divert the attention of environmental managers from more urgent protection needs.” There is mounting evidence that purple loosestrife invasions are disturbance-dependent and are “an indicator of anthropogenic disturbances.” In order to protect our wetlands, we must first protect them against undue disturbance. Lavoie supports using the Precautionary Principle when dealing with introduced species; however, he finds the approach “much more valuable for newcomers than for invaders coexisting with native species for more than a century.”

A field of purple loosestrife in Massachusetts – photo credit: wikimedia commons

Purple loosestrife has found its way to nearly every state in America and most of the Canadian provinces. Peter Del Tredici writes in Wild Urban Plants of the Northeast, “Conservationists despise purple loosestrife, despite its beauty, and it is listed as an invasive species in most of the states where it grows.” By listing a plant as a noxious weed, landowners are obligated to remove it. Care must be taken though, as removal of purple loosestrife can result in a secondary invasion by noxious weeds with an even worse track record, such as common reed or reed canary grass. “Hardly a gain from the biodiversity point of view,” quips Lavoie.

Claude Lavoie’s paper and the papers he references are definitely worth reading. It is important that we continue to study purple loosestrife and species like it in order to fully understand the impact that introduced species are having on natural areas, especially since it is unlikely that we will ever completely eliminate them. On that note, I’ll leave you with this passage from The Book of Swamp and Bog by John Eastman:

The situation is easy for environmentalists to deplore. This plant, like few others, stirs our alien prejudice. Our native cattails, for example, are almost as rudely aggressive and competitive in many wetland areas as purple loosestrife. Yet, because cattails obvioulsy ‘belong here,’ they seldom evoke the same outraged feelings against their existence. … With the spread of purple loosestrife, we have new opportunities to witness the phases of an ever-recurring ecological process. We can watch it affect, change, adapt, and refit both its own elements and those of invaded communities into new arrangements of energy efficiency. The point is that we might as well study this process rather than simply deplore it; we have few alternatives.

Horticulture’s Role in the Spread of Invasive Plants

I live in the city of Boise – a bustling metropolis by Idaho’s standards. It is located in the high desert of the Intermountain Northwest in a region called the sagebrush steppe. Our summers are hot and dry, and our native flora reflects this.

When I leave my apartment I am greeted by a flowering quince (Chaenomeles sp.). At this time of year it is in full bloom and looking amazing. It originated in East Asia. To my left I see a tree of heaven (Ailanthus altissima), a common urban tree that came to America from China via Europe. To my right there is a row of Norway maples (Acer platanoides), another popular urban tree. As its common name suggests, it is a European species that is distributed across large portions of eastern and central Europe. None of these plants are native to the sagebrush steppe, nor would they survive the harsh conditions without supplemental irrigation. All are horticultural introductions.

Tree of Heaven (Ailanthus altissima) – photo credit: wikimedia commons

But there is another thing that at least two of these species have in common. Tree of heaven and Norway maple are considered invasive species in North America due to their propensity to spread into natural areas and disrupt native ecosystems. They also have a reputation of being pesky urban weeds.

My experience isn’t unique. Yards across North America are planted largely with species that are not native to this continent, and while most species stay where we plant them, a significant portion of them have leaped out of our tidy landscapes and disseminated themselves across natural areas, earning them the title invasive species.

In a paper published in BioScience (2001), Sarah Hayden Reichard and Peter White discuss the role that horticulture has played in introducing invasive species to the United States. Humans have a long history of moving plants from one part of the world to another for food, fuel, and fiber. However, collecting plants from around the world and organizing them into gardens for aesthetic purposes is, by comparison, a more recent thing. Species used for ornamental horticulture are what Reichard and White are concerned about.

As an introduction, Reichard and White offer a quick history of the beginnings of ornamental horticulture in the United States. This period is summed up well in an article by Richard Mack and Mark Lonsdale in the same issue of BioScience:

As colonists became more secure in their new environments, they began to import ornamental species from their homelands and elsewhere, in simultaneous quests for both familiar and unfamiliar plants. These plant importations sprang from deep-seated or primal aspects of human behavior shared by people in former colonies and homelands alike. … Many needed to be reassured with familiar plants from home, and they also had seemingly antithetical desires to experience novel, exotic ornamental plants.

Today, plant explorations continue throughout the world, often with the goal of introducing new plant species to the horticulture trade, and avid gardeners remain eager to find something new and interesting to add to their yards. There is nothing inherently wrong with this. Nor is there anything inherently wrong with filling our yards with exotic plants. The trouble comes when these plants escape cultivation and cause problems in neighboring ecosystems. Bringing awareness to this darker side of ornamental horticulture is what Reichard and White endeavor to do.

“Thomas Jefferson, an avid horticulturist, also introduced several species. He may have been the first person to introduce Cytisus scoparius (Scotch broom) as an ornamental species; that plant is now an invasive species in many parts of North America.” — Reichard and White (2001) [photo credit: www.eol.org]

Major players involved in the global movement of horticultural specimens include botanical gardens and arboreta, nurseries, garden clubs and horticultural societies, and the seed trade industry. The motives for transporting species vary among the groups, as do their roles in addressing the invasive species issue. Many botanical gardens have extensive plant exploration programs, which today are often more conservation focused than they were in the past; however, some of the species acquired during these explorations are released to the public, often without certainty that they won’t spread.

Even though most nurseries don’t have active plant exploration programs, they may acquire plants from nurseries or other institutions that do. For business reasons, plants may be sold before they have been properly screened for invasive-ness. Some retail nurseries make an effort to not sell plants that are known invasives in their regions. However, there are plenty of mail order nurseries that may not be aware of or may simply ignore the fact that they are shipping plants to regions where they are invasive. Seed exchanges between garden clubs and botanical societies, as well as the seed trade industry, are also responsible for shipping species to areas where they are currently or may become invasive.

“Uninformed people sometimes dump their aquarium water and plants into local water sources, and many of the aquarium plants survive and multiply. Hydrilla verticillata, a very aggressive aquatic weed in the South, was probably introduced to provide a domestic source of this plant for the aquarium trade.” — Reichard and White (2001) [photo credit: wikimedia commons]

Plant exploration will continue, and many new plants will be introduced to the public through the horticulture trade. Rules and regulations help restrict some plant movement, but in a capitalist society such restrictions will ultimately be, as Reichard and White write, “a compromise between ideal invasive plant exclusion and trade facilitation.” Plants can be screened for invasivibility, but it is difficult to know if, when, and where a species may become invasive. Furthermore, given enough time, a species that appeared to stay put can suddenly start to spread (or could have been spreading all along unnoticed).

Reichard and White acknowledge that “the burden of finding a solution to the problems posed by invasive plants does not necessarily fall on the shoulders of [the horticulture] industry.” Various groups from broad disciplines will have to come to together to work towards a solution. Reichard and White offer some suggestions for working together. For example, invasive species biologists can share their research with the horticulture industry which can, in turn, communicate this information to the public through garden writers and speakers. Botanical gardens can take a leadership role by vowing to “first do no harm to plant diversity and natural areas” and by providing public education about the issue.

Efforts can be made to ban the sale of problematic plants and to encourage proper screening of new introductions, but public demand for certain plants may remain. So, “better communication from ecologists to the public about which species are causing problems will discourage people from buying them.” Involving the public in eradication efforts can also help raise awareness, as people can see first hand that plants in their yards have invaded the wild.

When Alien Plants Invade – The Four Stages of Invasion, part two

In a review published in New Phytologist (2007), Kathleen Theoharides and Jeffrey Dukes examine four stages of invasion as they relate to alien (i.e. introduced or non-indigenous) plant species. In part one we discussed transport and colonization, in which species must survive being transported long distances and then take root and reach maturity in an unfamiliar location. Alien plant species don’t become invasive until they have reached the last two stages: establishment and landscape spread. Removal of the species upon reaching these stages is no easy task. Luckily, introduced species have a few barriers to overcome before this point.

An established population is one that is self-sustaining and expanding. Environmental conditions may be a limiting factor, as they were during colonization, but the main constraints at this stage are “biotic filters.” Theoharides and Dukes define these as “barriers to invasion created by the actions or presence of living organisms.” Through competition for various resources, as well as herbivory and disease, neighboring organisms affect the survival, growth, and reproduction of introduced plant species. Thus, “traits that enhance competitive performance, reduce niche overlap between [introduced species] and natives, or increase enemy resistance may be most important during establishment… Other advantageous traits include secondary chemical compounds that deter herbivores, ‘novel weapons’ such as root exudates that negatively impact other plants, fast growth, and high fecundity.”

Plants compete for light, moisture, and soil nutrients, as well as for pollinators and seed dispersers. Competition inhibits the establishment of invaders when neighboring plants consume available resources more efficiently. Introduced plants risk being outcompeted by plants that are of the same functional type (plants that are “morphologically, phenologically, and physiologically similar”). They also risk competition by a single dominant species (or group of similar species) or by “an assemblage of species with different traits.” As a general rule, plant communities with greater diversity are more resistant to invasion.

“In forests of the northeastern USA, Alliaria petiolata, an herbaceous mustard species, contains a type of phytotoxic glucosinolate that appears to disrupt the mutualism between arbuscular mycorrhizal fungi and hardwood canopy trees.” – – Theoharides and Dukes (2007) [photo credit: eol.org]

Two hypotheses postulate the success of some plant invaders in establishing themselves: the enemy release hypothesis and the evolution of increased competitive ability hypothesis. In the first hypothesis, plant species – having been removed from their native habitat – are freed from their natural enemies and are thus able to allocate more resources to growth and reproduction. The second hypothesis states that, in light of “reduced enemy pressure,” introduced species quickly evolve to allocate resources “from enemy defense to faster growth.” Escape from herbivory and diseases, however, is likely not the only factor in the success of invaders, and much still depends on the competitiveness of the plant and the availability of key resources.

After introduced plants become established, a lag phase generally occurs before landscape spread. This can be a result of a lack of genetic variation, a dearth of suitable habitat, unfavorable environmental conditions, or some combination of the three. New introductions may occur, and the population may continue to adapt and expand. Suitable habitat may be made available, and environmental conditions may shift. In time, landscape spread becomes a possibility.

Landscape spread occurs when multiple populations of a species are connected via long-distance dispersal. At this “metacommunity scale,” populations of an introduced plant species interact across a large area, with each population in a different stage of colonization and establishment. This means that transport, colonization, and establishment are all at play during the landscape spread stage.

Abutilon theophrasti (velvetleaf) was originally introduced before 1700 in the USA. This species has only recently become an aggressive invader as a result of the evolution of different life-history strategies based on the nature of competition in its new environment.” — Theoharides and Dukes (2007) [photo credit: wikimedia commons]

Dispersal ability and habitat connectivity are key factors in determining the success of an introduced plant species during landscape spread. Long-distance dispersal can occur via wind, water, or animals. Species that depend on animals to spread their seeds rely on specific animals to be present. The seeds of Prunus serotina (black cherry), for example, are dispersed by birds. So, landscape spread is reliant on birds and “roosting trees” where the birds can perch and defacate the seeds. In many cases, “humans also play a large role in intraregional dispersal.”

Habitats vary across the landscape due to a combination of numerous geological and biological processes. The disturbance regime – “the frequency, spatial extent, severity, and intensity of killing events over time” – also helps determine landscape patterns. Natural disturbances, such as fire, weather, and natural disasters, are differentiated from disturbances caused by human activity. Large scale development and disturbance of natural areas by humans disrupts the natural disturbance regime and alters historical landscape patterns. As the authors write, “alterations of the disturbance regime that increase resource availability or change landscape patterns can promote non-indigenous plant species spread by creating favorable patches for colonization and establishment.”

Fragmented landscapes consisting of small patches of natural areas dispersed among large areas of human development are particularly prone to invasion by introduced plant species for many reasons, including increased influx of propagules and a high degree of edge effects (habitat edges have environmental conditions that are generally more prone to invasion than habitat interiors).

Habitat patches can be connected via corridors. It is through these corridors that dispersal can occur between populations in a metacommunity. Corridors connect populations of both introduced and native plant species. However, “native plants often require wide undisturbed corridors of intact habitat, while [introduced plant species] may disperse best through strips of human-disturbed habitat or ‘disturbance corridors.'” The environmental conditions in disturbance corridors and the presence of dispersal agents (including humans and domesticated animals) help facilitate the connectivity of populations of introduced plant species and promote the colonization and establishment of new populations.

In their abstract, Theoharides and Dukes write, “both research and management programs may benefit from employing multiscale and stage approaches to studying and controlling invasion.” With their conclusion they provide a list of potential management strategies for each stage, and they advise employing “natural filters in order to prevent invasion succees.” Examples include reducing habitat fragmentation and edge effects, promoting intact native communities, reducing human disturbances, promoting natural disturbance regimes, and minimizing disturbance corridors.

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