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

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

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

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

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

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

flowers of creeping thistle (Cirsium arvense) via eol

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

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

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

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

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

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

Pitcher’s thistle (Cirsium pitcheri) via eol

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

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

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

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When Urban Pollinator Gardens Meet Native Plant Communities

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

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

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

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

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

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

photo credit: wikimedia commons

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

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

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

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

photo credit: wikimedia commons

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

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

Rare and Endangered Plants: Texas Wild Rice

Some plants have native ranges that are so condensed that a single major disturbance has the potential to wipe them out of existence completely. They are significantly more vulnerable to change than neighboring plant species, and for this reason they often find themselves on endangered species lists. Zizania texana is one of those plants. Its range was never large to begin with, and due to increased human activity it now finds itself on the brink of extinction.

Zizania texana is one of three species of wild rice found in North America. The other two, Z. palustris and Z. aquatica, enjoy much broader ranges. Both of these species were once commonly harvested and eaten by humans. Today, Z. palustris is the most commercially available of the two. Commonly known as Texas wild rice, Z. texana, was not recognized as distinct from the other two Zizania species until 1932.

Herbarium voucher of Texas wild rice (Zizania texana) - photo credit: University of Texas Herbarium

Herbarium voucher of Texas wild rice (Zizania texana) – photo credit: University of Texas Herbarium

Texas wild rice is restricted to the headwaters of the San Marcos River in Central Texas. The river originates from a spring that rises from the Edwards Aquifer. It is a mere 75 miles long, but is home to copious amounts of wildlife, including several rare and endangered species. Before the 1960’s, Texas wild rice was an abundant species found along several miles of the San Marcos River. Its population and range has since been greatly reduced, and the native population is now limited to about 1200 square meters within the first two miles of the river.

Texas wild rice is an aquatic grass with long, broad leaves that remains submerged in the clear, flowing, spring-fed water of the river until it is ready to flower. Flower heads rise above the water, and each flower spike consists of either male or female flowers. The flowers are wind pollinated, but research has revealed that the pollen does not travel far and does not remain viable for very long. If a male flower is further than about 30 inches away from a female flower, the pollen generally fails to reach the stigma. The plants also reproduce asexually by tillering, but plants produced this way are genetically identical to the parent plant.

As people settled in the area around San Marcos Springs and began altering the river for their own use, Texas wild rice had to put up with a series of assaults and dramatic changes, including increased sediment and nutrient loads, variations in water depth and speed, trampling, and mechanical and chemical removal of the plant itself. Sexual reproduction became more difficult. In his book, Enduring Seeds, Gary Paul Nabhan describes one scenario: “streamflow had been increased to the extent that the seedheads, which were formerly raised a yard above the water, [were] now constantly being pummeled by the current so that they [remained] submerged, incapable of sexual reproduction.”

San Marcos, Texas – where the headwaters of the San Marcos River is located and where Texas wild rice has long called its home – is the location of Texas State University and is part of the Greater Austin metropolitan area. Thus, Zizania texana now finds itself confined to a highly urbanized location. The San Marcos Springs and River are regularly used for recreation, which leads to increased sediments, pollution, and trampling. Introduced plant species compete with Texas wild rice, and introduced waterfowl and aquatic rodents consume it. In this new reality, sexual reproduction will remain a major challenge, and a return to its original population size seems veritably impossible.

Texas wild rice (Zizania texana) and its urbanized habitat - photo credit: The Edwards Aquifer

Texas wild rice (Zizania texana) and its urbanized habitat – photo credit: The Edwards Aquifer

Attempts have and are being made to maintain the species in cultivation and to reintroduce it to its original locations, but its habitat has been so drastically altered that it will need constant management and attention for such efforts to be successful. As Nabham puts it, it is a species that has “little left of [its] former self in the wild – it is a surviving species in name more than in behavior…The wildness has been squeezed out of Texas rice.”

What if humans had stayed out of it? Would a plant with such a limited range and such difficulty reproducing sexually persist for any great length of time? It’s hard to say. If it disappears completely, what consequences will there be? It is known to provide habitat for the fountain darter, an endangered species of fish, as well as several other organisms; however, the full extent of its ecological role remains unclear. It will be nursed along by humans for the foreseeable future, but it may never regain its full glory. It is a species teetering on the edge of extinction, simultaneously threatened and cared for by humans – a story shared by so many other species around the world.

Additional Resources:

Field Trip: Mud Springs Ridge and Cow Creek Saddle

Last weekend I went on two all day field trips that were part of Idaho Native Plant Society‘s annual meeting. The second field trip was in a location with a climate considerably warmer and drier than the first field trip. The flora was much more familiar to me since it was similar to what I generally see in southern Idaho. We visited two sites: Mud Springs Ridge and Cow Creek Saddle. Both are high on a mountain ridge (around 5300 feet in elevation) flanked by the Salmon River canyon on the east and the Snake River canyon on the west. The tiny town of Lucile, Idaho was just below us to the east, and if we would have continued down the other side of the mountain, we would have arrived at Hells Canyon National Recreation Area. These sites are high elevation grasslands, and there was a huge diversity of grasses and forbs to explore.

Taking decent photos of the plants was a challenge as the sun was shining brightly and there was a constant breeze. Photographs don’t quite cut it anyway. The views were incredible. Standing on a ridge top peering across a meadow full of wildflowers with more mountains in the distance. Mass amounts of lupines and paintbrushes mixed with grasses and other plants being tossed about in the breeze. Little rock gardens randomly dispersed across the hillsides. You kind of had to be there.

A view across the meadow at Mud Springs Ridge

A view across the meadow at Mud Springs Ridge

Searching for Silene spaldingii - an Idaho endemic - on the mountainside

Fellow botany geeks searching for Silene spaldingii (Spalding’s catchfly) – a rare, imperiled plant species

Gnarly, old curl-leaf mountain mahogany (Cercocarpus ledifolius) growing out of a rock outcrop

Gnarly, old Cercocapus ledifolius (curl-leaf mountain mahogany) growing out of a rock outcrop

Close up of Cercocarpus ledifolius

Cercocarpus ledifolius (curl-leaf mountain mahogany)

Orthocarpus tenuifolius (owl's clover)

Orthocarpus tenuifolius (thin-leaved owl’s clover)

Castilleja (indian paintbrush)

Castilleja hispida (harsh paintbrush)

Castilleja cusickii (Cusick's paintbrush)

Castilleja cusickii (Cusick’s paintbrush)

Lewissia columbiana v. wallowaensis

Lewisia columbiana var. wallowensis (Wallowa lewisia)

Lewissia columbiana v. wallowaensis

Lewisia columbiana var. wallowensis (Wallowa lewisia)

Erigeron

Erigeron davisii (Davis’ fleabane)

On cow creek saddle looking towards Salmon River canyon

On Cow Creek Saddle looking towards Salmon River canyon

On cow creek saddle looking towards Snake River canyon

On Cow Creek Saddle looking towards Snake River canyon

The field trips were incredible, and the annual meeting in general was a lot of fun. If you have a native plant society in your neck of the woods and you are not already a member, I highly recommend checking it out. Now, where to next?

A Rare Hawaiian Plant – Newly Discovered and Critically Endangered

Hawaii is home to scores of plant species that are found nowhere else in the world. But how did those plants get there? In geological time, Hawaii is a relatively young cluster of islands. Formed by volcanic activity occurring deep within the ocean, they only just began to emerge above water around 10 million years ago. At that point the islands would have been nearly devoid of life, and considering that they had never been connected to any other body of land and are about 2,500 miles away from the nearest continent, becoming populated with flora and fauna took patience and luck.

As far as plant life is concerned, seeds and spores had to either be brought in by the wind, carried across the ocean by its currents, or flown in attached to the feathers of birds. When humans colonized the islands, they brought seeds with them too; however, its estimated that humans didn’t begin arriving on the islands until about 1,700 years ago. The islands they encountered were no longer barren landscapes, but instead were filled with a great diversity of plant and animal life. A chance seed arriving on the islands once in a blue moon does not fully explain such diversity.

This is where an evolutionary process called adaptive radiation comes in. A single species has the potential to diverge rapidly into many new species. This typically happens in new habitats where little or no competition exists and there are few environmental stresses. Over time, as genetic diversity builds up in the population, individuals begin to adapt to specific physical factors in the environment, such as soil type, soil moisture, sun exposure, and climate. As individuals separate out into these ecological niches, they can become reproductively isolated from other individuals in their species and eventually become entirely new species.

This is the primary process that led to the great floral diversity we now see on the Hawaiian Islands. Adaptive radiations resulted in more than 1000 plant species diverging from around 300 seed introductions. Before western colonization, there were more than 1,700 documented native plant species. Much of this diversity is explained by the rich diversity of habitats present on the volcanic islands, which lead to many species becoming adapted to very specific sites and having very limited distributions.

A small population size and a narrow endemic range is precisely the reason why Cyanea konahuanuiensis escaped detection until recently. In September 2012, researchers on the island Oahu arrived at a drainage below the summit of Konahua-nui (the tallest of the Ko’olau Mountains). They were surveying for Cyanea humboldtiana, a federally listed endangered species that is endemic to the Ko’olau Mountains. In the drainage they encountered several plants with traits that differed from C. humboldtiana, including hairy leaves, smooth stems, and long, hairy calyx lobes. They took pictures and collected a fallen leaf  for further investigation.

Ko'olau Mountains of O'ahu (photo credit: wikimedia commons)

Ko’olau Mountains of O’ahu (photo credit: wikimedia commons)

Initial research suggested that this was a species unknown to science. More information was required, so additional trips were made, a few more individuals were found, and in June 2013, a game camera was installed in the area. The camera sent back three photos a day via cellular phone service and allowed the team of researchers to plan their next trip when they were sure that the flowers would be fully mature. Collections were kept to a minimum due to the small population size; however, using the material they could collect, further analyses and comparisons with other species in the Cyanea genus and related genera demonstrated that it was in fact a unique species, and so they gave it the specific epithet konahuanuiensis after the mountain on which it was found. It was also given a common Hawaiian name, Haha mili’ohu, which means “the Cyanea that is caressed by the mist.”

The hairy flowers and leaves of Cyanea konahuanuiensis. Purple flowers appear June-August. (photo credit: www.eol.org)

The hairy flowers and leaves of Cyanea konahuanuiensis. Purple flowers appear June-August. (photo credit: www.eol.org)

The total population of Cyanea konahuanuiensis consists of around 20 mature plants and a couple dozen younger plants. It is considered “critically imperiled” and must overcome some steep conservation challenges in order to persist. To start with, the native birds that pollinate its flowers and disperse its seeds may no longer be present. Also, it is likely being eaten by rats, slugs, and feral pigs. Add to that, several invasive plant species are found in the area and are becoming increasingly more common. While the researchers did find some seedlings in the area, all of the fruits that they observed aborted before they had reached maturity. Lastly, the population size is so small that the researchers say a landslide, hurricane, or flash flood “could obliterate the majority or all of the currently known plants with a single event.”

Seeds collected from immature fruits from two plants were sown on an agar medium at the University of Hawaii Harold L. Lyon Arboretum. The seeds germinated, and so the researchers plan to continue to collect seeds “in order to secure genetic representations from all reproductively mature individuals in ex situ collections.”

Single stem of Cyanea konahuanuiensis (photo credit: www.eol.org)

Single stem of Cyanea konahuanuiensis (photo credit: www.eol.org)

C. konahuanuiensis is not only part of the largest botanical radiation event in Hawaii, but also the largest on any group of islands. At some point in the distant past, a single plant species arrived on a Hawaiian island and has since diverged into at least 128 taxa represented in six genera, Brighamia, Clermontia, Cyanea, Delissea, Lobelia, and Trematolobelia, all of which are in the family Campanulaceae – the bellflower family. Collectively these plants are referred to as the Hawaiian Lobelioids. Cyanea is by far the most abundant genus in this group consisting of at least 79 species. Many of the lobelioids have narrow distributions and most are restricted to a single island.

Sources