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:]

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:]

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.

More Posts about Invasive Species:

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

As humans move around the globe, they are regularly accompanied by plants. Some plant species are intentional guests, while others are interlopers. This steady movement of plants from one region to another results in plants being introduced to areas where they are not native. In this regard, they are aliens. Some of these alien species will take up permanent residence and, as a result, can disrupt ecosystems, compete with native plant species, and cause economic damage. This earns them the title “invasive”. But not all introduced plant species achieve this. In fact, many will find themselves in a new region but will be unable to colonize. Others will colonize but not become fully established. Still others become established but will not spread. In all cases there are factors at play that either aid or limit an introduced plant species in becoming invasive.

In a review published in New Phytologist (2007), Kathleen Theoharides and Jeffrey Dukes examine four stages of invasion (transport, colonization, establishment, and landscape spread) and some of the “filters” that occur in each stage that help determine whether or not an alien plant species will become invasive. In their introduction they clarify, “these stages are not discrete, and filters will likely affect more than one stage,” but by analyzing each of the stages we can better determine how and why some introduced species are successful at becoming invasive while others are not. Generalities derived from this investigation can “be used to predict the outcome of invasion events, or to explore mechanisms responsible for deviations from these generalizations.”

Theoharides, K. A. and Dukes, J. S. (2007), Plant invasion across space and time: factors affecting nonindigenous species success during four stages of invasion. New Phytologist, 176: 256–273. doi:10.1111/j.1469-8137.2007.02207.x

In part one, we will look at the first two stages of invasion: transport and colonization.

Species have always moved around from region to region by various means. However, as Theoharides and Dukes write, “current species movements are happening faster than before and from more distant regions, primarily as a result of global commerce and travel.” When it comes to human-mediated dispersal, many plants may never be transported by humans, while others simply won’t survive the journey. Species that are widespread may have a better chance of being transported because they are more likely to make contact with humans. Transporting high numbers of propagules (i.e. seeds, spores, cuttings) generally increases the likelihood that a species will survive the journey.

The invasion of Phalaris arundinacea (reed canary grass) was facilitated by multiple introduction events from a variety of sources within the native European range.” — Theoharides, K. A. and Dukes, J. S. (2007) [photo credit:]

Plants are transported by humans for many reasons. Sometimes its accidental, but often it is purposeful for either utilitarian or aesthetic purposes. Plants provide us with food, fuel, forage, building materials, clothing, and medicine. Over millennia, we have selected suites of species that are ideal for such purposes, and we have carried them with us into new regions or brought them home from other parts of the world. Not all of these species are well-behaved in their new homes, and many have become invasive. These species are given an advantage because they have been selected for traits like cold hardiness, disease resistance, and high yield. When they are transported, they are brought to locations with similar climates. “Climate matching, combined with intentional cultivation, greatly increases the likelihood that [these] species will escape cultivation.”

Surviving transportation is not a guarantee that alien plants will successfully colonize a new area. Myriad environmental conditions and biological processes stand in their way. Much depends on propagule pressure – “the combined measure of the number of individuals reaching a new area in any one release event and the number of discrete release events.” Where propagule pressure is high, colonization is more likely. Repeated introductions across a large area offer the species a greater chance of finding itself in a suitable location as well as a greater level of genetic variation. Disturbed environments with less competitors and increased resources (i.e. light, moisture, soil nutrients) are often easier to colonize than locations with a high level of biodiversity and fewer available resources.

“Populations of Salix babylonica (weeping willow) in New Zealand may have invaded from a single cutting.” — Theoharides, K. A. and Dukes, J. S. (2007) [photo credit:]

Climate is one of the main filters of colonization, yet plant species have still managed to colonize regions with very different climates compared to what they’re used to, while other plant species have been unsuccessful in colonizing regions with similar climates. Plant species that originate from wide geographic ranges tend to have “broader climatic tolerances” – a trait that along with phenotypic plasticity and a high level of genetic variability can enable a species to adapt to new and challenging environments. Other advantageous traits include “fast growth, self-compatibility, a short juvenile period, and seeds that germinate without a pre-treatment.”

If and when colonization is achieved, establishment is no guarantee. “In order for a plant to establish itself it must continue to increase from low density over the long term.” Small numbers of plants may successfully reproduce, but environmental factors, genetic issues, and biological competition may still stand in their way. Species that invade disturbed sites where resource availability is temporarily high, may soon find themselves in a resource-limited situation. As a result, their populations may dwindle.

With transport and colonization accomplished, establishment is the next goal. Establishment and landscape spread will be covered in part two.

Seed Dispersal via Caching – The Story of Antelope Bitterbrush

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

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

Fruits forming on antelope bitterbrush (Purshia tridentata)

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

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

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

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

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

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

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


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

Learning Lessons from Invaded Forests

In 1946, North American beavers were introduced to the archipelago of Tierra del Fuego at the tip of South America in an attempt to start an industry based on beaver fur. Although this industry has not thrived, beavers have multiplied enormously. By cutting trees and building dams, they have transformed forests into meadows and also fostered the spread of introduced ground cover plants. Now numbering in the tens of thousands in both Chilean and Argentinian parts of the archipelago, beavers are the target of a binational campaign to prevent them from spreading to the mainland of these two nations. — Invasive Species: What Everyone Should Know by Daniel Simberloff

Beavers in South America are just one example of the series of effects a species can have when it is placed in a new environment. Prior to the arrival of beavers, there were no species in the area that were functionally equivalent. Thus, through their felling of trees and damning of streams, the beavers introduced novel disturbances that have, among other things, aided the spread of non-native plant species. Ecologists call this an invasional meltdown, wherein invasion by one organism aids the invasion of another, making restoration that much more difficult.

Complicated interactions like this are explored by David Wardle and Duane Peltzer in a paper published last month in Biological Invasions. Organisms from all walks of life have been introduced to forests around the world, and while many introductions have had no discernable impact, others have had significant effects both above and below ground.

The authors selected forest ecosystems for their investigation because “the imprint of different invaders on long-lived tree species can often be observed directly,” even when the invading organisms are doing their work below ground. Moreover, a greater understanding of “the causes and consequences of invasions is essential for reliably predicting large-scale and long-term changes” in forest ecosystems. Forests do not regenerate quickly, so protecting them from major disturbances is important. Learning how forests respond to invasions can teach us how best to address the situation when it occurs.

The authors begin by introducing the various groups of organisms that invade forests and the potential impacts they can have. This is summarized in the graphic below. One main takeaway is that the effects of introduced species vary dramatically depending on their specific attributes or traits and where they fall within the food chain. If, like the beaver, a novel trait is introduced, “interactions between the various aboveground and belowground components, and ultimately the functioning of the ecosystem” can be significantly altered.

Wardle, D.A., and D.A. Peltzer. Impacts of invasive biota in forest ecosystems in an aboveground-belowground context. Biological Invasions (2017).  doi:10.1007/s10530-017-1372-x

After highlighting some of the impacts that invasive species can have above and below ground, the authors discuss basic tenets of invasion biology as they relate to forest ecosystems. Certain ecosystems are more vulnerable to invasions than others, and it is important to understand why. One hypothesis is that ecosystems with a high level of species diversity are more resistant to invasion than those with low species diversity. This is called biotic resistance.  When it comes to introduced plants, soil properties and other environmental factors come in to play. One species of plant may be highly invasive in one forested ecosystem, but completely unsuccessful in another. The combination of factors that help determine this are worth further exploration.

When it comes to restoring invaded forests, simply eliminating invasive species is not always enough. Because of the ecological impacts they can have above and below ground, “invader legacy effects” may persist. As the authors write, this requires “additional interventions to reduce or remove [an invader’s] legacy.” Care also has to be taken to avoid secondary invasions, because as one invasive species is removed another can take its place.

Nitrogen-fixing plants (which, as the authors explain, “feature disproportionately in invasive floras”) offer a prime example of invader legacy effects. Introducing them to forest ecosystems that lack plants with nitrogen fixing capabilities “leads to substantially greater inputs of nitrogen … and enhanced soil fertility.” Native organisms – decomposer and producer alike – are affected. Simply removing the nitrogen fixing plants does not at once remove the legacy they have left. Examples include Morella faya invasions of forest understories in Hawaii and invasions by Acacia species in South Africa and beyond.

“It has been shown that co-invasion by earthworms enhances the effects that the invasive nitrogen fixing shrub Morella faya has on nitrogen accretion and cycling in a Hawaiian forest, by enhancing burial of nitrogen-rich litter.” – D.A. Wardle and D.A. Peltzer (2017) – photo credit: wikimedia commons

The authors conclude with a list of “unresolved issues” for future research. A common theme among at least a couple of their issues is the need for observing invasive species and invaded environments over a long period. Impacts of invasive species tend to “vary across both time and space,” and it is important to be able to predict “whether impacts are likely to amplify or dampen over time.” In short, “focus should shift from resolving the effects of individual invasive species to a broader consideration of their longer term ecosystem effects.”

This paper does not introduce new findings, but it is a decent overview of invasion biology and is worth reading if you are interested in familiarizing yourself with some of the general concepts and hypotheses. It’s also open access, which is a plus. One thing that is clear after reading this is that despite our growing awareness of the impacts of invasive species, there is still much to be learned, particularly regarding how best to respond to them.


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