Purist or Pragmatist? Identifying and Addressing Non-Native vs. Invasive Species in Prairies.

Recently, there has been a lot of consternation and confusion among biologists and the public about invasive species.  Much of the confusion comes from misusing the term “invasive species”, and particularly the practice of using the terms “non-native (or exotic) species” and “invasive species” interchangeably.  This really needs to stop.

In North America, non-native species are generally defined as species that were not present in an ecosystem prior to European settlement.  There is plenty of discussion to be had about whether or not that is a useful definition, but there you go.  While definitions of invasive species vary, most ecologists use a definition similar to that used by the National Invasive Species Informational Center, which has two important parts.  First, the species must be non-native to the ecosystem in question.  Second, and most importantly, a non-native species must either cause – or be likely to cause – harm to the environment or human health in order to be considered invasive.  That ability of the species to cause harm when it’s been introduced into an ecosystem is the key characteristic that splits invasive species from all other non-native species.

A third term often thrown into invasive species discussions is “weed.”  The common definition of a weed is “a plant out of place” – meaning that any plant that shows up where someone doesn’t want it can be considered a weed.  I don’t have any problem with that definition, but there are at least two problems that arise from using the term “weed” interchangeably with “invasive species”.  First, of course, weeds can only be plants, and there are plenty of invasive species that aren’t plants.  Second, because of the definition, a plant that is called a weed by one person might be a wonderful plant to the next.  Lots of native perennial prairie wildflowers show up in weed books and websites, for example, because the authors of those sites see them as extraneous to pastures, yards, or other non-prairie habitats.  Many other “weedy species” (native and non-native) are important components of prairies – temporarily occupying space abandoned by stressed perennial plants until those perennials recover from fire, grazing, or other disturbances.

Common mullein (Verbascum thapsis) is a non-native plant considered to be invasive in some places, but not in Nebraska. In our Platte River Prairies it appears when dominant grasses are weakened, but doesn't persist when those grasses recover.

Several high profile articles and opinion pieces have recently questioned the value of fighting against species simply because they are “non-native”.  I think it’s a great topic of discussion, albeit one that makes many ecologists uncomfortable.  However, some of the authors – and MANY of those who have subsequently reported on the original pieces – make the very dangerous error of interchanging the terms “non-native” and “invasive” as they discuss the topic.  Even The Nature Conservancy’s Chief Scientist made this mistake in a blog post he wrote about a journal article by Mark Davis and his colleagues. 

Here’s the thing, it’s really easy for those of us working to conserve native ecosystems to become purists about the species we want in those ecosystems.  We are fortunate in North America to have ecosystems that consist largely of the same species that were present in those systems at the time of European settlement.  There are advantages to having ecological systems full of those “native” species because those species have had time to develop relationships and counterbalances with/to each other.  However, very few ecological communities (none that I work with) remain completely devoid of non-native species, and keeping them “pure” gets harder all the time.  Drawing a line in the sand and working to prevent any non-native species from getting a foothold in ecological communities is not only impossible, it’s foolhardy as well.

Don’t get me wrong, we spend a lot of time fighting invasive species in our Platte River Prairies.  It’s by far the most time-consuming and expensive part of our stewardship work – and extremely important.  However, we’re as careful as we can be to prioritize those species that we work against so we aren’t spending time and effort on species that aren’t causing any problems.  (If you’re interested, I wrote an earlier post on prioritizing invasive species.)

As I consider whether or not a particular species is one to attack or live with, I focus mainly on the question of whether that species adds to, or subtracts from, the species diversity, ecological resilience, or ecological function of my prairies.  Does the new species outcompete others to the point where small or large scale diversity declines?  Or does the species actually increase species diversity by adding itself (and its functions) to the prairie?

For example, I don’t worry about plant species like the biennial plant goatsbeard, aka salsify (Tragopogon dubious), even though it’s a non-native species.  My experience with goatsbeard is that it periodically appears here and there in prairies – mainly where defoliation has weakened dominant grasses and opened space for the establishment of new plants.  It certainly doesn’t appear to be eliminating other species or negatively altering habitat.  Rather, it’s inserted itself into the plant community in such a way that someone who didn’t know better would assume it was native. 

Goatsbeard, aka salsify, is a non-native plant species that doesn't appear to be invasive.

In contrast, I worry a great deal about species such as crown vetch and smooth brome, because both can form large monocultures, and if left unchecked, can transform a prairie plant community into one dominated by only a few plant species.  Not only does that affect the plant species pushed out of the community, it severely alters habitat conditions for pollinators and other species that rely on those now missing plant species.  Further, it changes the habitat structure and food availability for many animals.  We try to eradicate crown vetch as soon as we find it, because it’s not (yet?) widespread on our sites.  We know we can’t eradicate smooth brome because it’s already all over the place, but we do design management strategies to suppress it when possible, and to encourage other plant species to fill space left by weakened brome plants. 

We also, by the way, suppress or kill native species that act aggressively and simplify either species composition or habitat structure.  For example, eastern red cedars are native species that encroach upon prairies when fire is absent.  We clear cedar trees that are too large to kill with fire, and use periodic prescribed fire to stop others before they grow very large.  In addition, we use herbicides to kill many other native tree and shrub species, including honey locust, smooth sumac, rough-leaved dogwood, and many others when their presence works against our ecological goals for a site.  Similarly, most of our fire and grazing management is designed to suppress the dominance of native warm-season grasses like big bluestem and indiangrass because those grasses can reduce plant diversity if given the chance.

This eastern red cedar had a short life prior to our prescribed fire that killed it. Cedars are native to Nebraska, but when left unchecked can drastically impact prairie habitats and species diversity.

There are, of course, invasive and other non-native animals too – not just plants – but for the most part, prairies in our areas have few animal species that we know to be causing problems.  There are a few invasive insect species that were introduced as biocontrol for invasive thistle species but subsequently began attacking native thistles – but I really don’t know much about them.  However, a couple of non-prairie birds provide good examples of non-native vs. invasive animals.  European starlings are non-native birds that are a substantial reason for the decline of native cavity nesting birds such as red-headed woodpeckers.  Starlings, therefore, are invasive species.  In contrast, Eurasian collared-doves are a new species of bird to Nebraska, but at least to my knowledge, no one has shown any reason to be concerned about that.  Collared-doves may just be a new species of bird that has successfully joined the community of species in our towns (where they are usually found).

There are, of course, some important differences between non-native invaders like crown vetch and smooth brome and native species like cedars and big bluestem.  Those differences include a long history of inter-species competition that has built plant communities that are unlikely to allow any one species to become overly dominant – or that at least have the capacity to prevent that dominance under certain fire, grazing, or other conditions.  Most successful invasive species are able to invade and dominate regardless of managment conditions, and require direct focused attack in order to control them.  An additional difference is that with native species, we know what we’re getting.  We have a pretty good idea of how they grow, how they interact with other species, and what factors give them an advantage or disadvantage within their communities.  With non-native species, we don’t have that history to draw upon as we try to decide whether or not to control them.  That uncertainty is a huge challenge for those of us who have decided not to be purists about native/non-native species, but to instead work against those species that decrease diversity and resilience.  How do we know which new species will be problems?

Sometimes we can use information from other land managers or published journal articles that have already evaluated species which are just showing up in our prairies.  However, even in those cases, it can be hard to know for sure how a new species will act when it comes in.  Varying soils and precipitation rates are just two of many factors that can lead to a species becoming invasive at one site and simply non-native at others.  Sweet clover is a great example of a species that is considered to be an awful invasive species in some places and simply a big ugly non-native plant in others.  I don’t like sweet clover, but I haven’t seen it decrease plant diversity over time at our sites.  In prairies we manage with periodic grazing, sweet clover is largely a non-issue because it is so palatable to cattle.  In our other prairies, it becomes visually abundant in some years and not in others, and the surrounding plant community seems to be just the same after a flush of sweet clover as it was before it.  I’ve spoken to other prairie managers around the country who see sweet clover as I do, but I also know managers who see it acting much more aggressively.  We’ve made the decision not to worry about sweet clover on our prairies, but I certainly don’t blame others who spend time and effort suppressing it.

Sweet clover is a species that is considered by some to be very invasive, but we tend to think of it as a big ugly plant that comes and goes without lasting impacts. This is a photo I took this summer in some of our research plots where we are tracking sweet clover and other non-native species in high and low diversity plant communities.

This kind of uncertainty about potential impacts creates an obvious conundrum.  We can’t fight every new species that comes into our prairies, but we don’t always know which to focus on and which not to.  Unfortunately, I don’t have any easy answers.  We all have to make our own decisions based on the best available information and the amount of time/effort we can spare.  For example, someone who owns a small prairie and has abundant resources might decide to prevent any new species from joining the prairie community to be completely safe.  On the other hand, most of us are forced to make difficult decisions (guesses?) because our resources are very limited. 

Maybe the key is simply to be thoughtful.  I think it’s a mistake to assume that a species is bad just because it’s new.  Not only are some non-native species fairly innocuous, many can provide benefits, including food sources for native animals, replacement for eradicated native species, etc.  However, it’s also a mistake to look at a few prominent examples of where non-native or invasive species turn out to provide unexpected benefits and assume that we should just accept all comers.  The successful nesting of endangered southwestern willow flycatchers in riparian habitats dominated by invasive salt cedar, for example, has led some to suggest that maybe salt cedar isn’t so bad after all.  This inaccurate, but frustratingly common, misreading of the situation ignores all of the damage to other aspects of the environment caused by salt cedar.  Clearly, there are some species that need to be controlled at all costs before they completely wreck important native habitats.

Salt cedar has been shown to be invasive, in that it has multiple negative impacts on the surrounding environment. Here it's growing in a patch of non-native Phragmites (common reed) - another invasive species.

All of us who manage prairies have to make daily decisions based on best available information, past experience, and gut feelings.  Decisions about invasive and other non-native species are really no different.  I certainly don’t plant seeds of non-native plant species in our prairie restorations, but I also don’t worry about most of the non-native plants that establish there.  Frequent conversations with other prairie managers help me keep track of which species are causing problems elsewhere, and that helps me make decisions about our sites.  I’m sure I’ll make (have made?) mistakes about which species to worry about, but I hope that careful monitoring of those species and their interactions with the surrounding community will help me catch those mistakes early. 

I’d love to have prairies with only native species in them.  I’d also love to have prairies with millions of contiguous acres, large herds of free-roaming bison, and staggering plant, insect, and animal diversity – along with a huge staff of well-trained and well-equipped land managers.  Unfortunately, the reality is that I manage relatively small and fragmented prairies with a small staff and old equipment – and our prairies contain quite a few non-native species.  I try to eradicate known invasive species as they appear, suppress the ones that are entrenched, and make smart decisions about species I’m unsure of.  Anyone have a better plan? 

 For those of you in Nebraska, there is a great invasive species site that contains current events, species lists, and other great information about invasive species.  Visit the Nebraska Invasive Species Project site here

Ecological Resilience in Prairies: Part 2

This is Part 2 of a two part series on ecological resilience in prairies.  In Part 1, I interviewed Dr. Craig Allen about the basic definition of ecological resilience and then wrote about the relevance and application or resilience to prairie ecosystems.  In Part 2, I explore how ecological resilience can influence the way we restore and manage prairies, and about how much we still have to learn about how to do that.

 

Influencing Resilience through Restoration and Management

Understanding ecological resilience should help us better design restoration and management strategies that build and maintain resilience in prairies.  Using the components of resilience discussed in Part 1, it seems apparent that when restoring (reconstructing) prairies, it’s important to maximize species diversity in seed mixtures.  More importantly, prairie restoration that adds to the size and connectivity of existing prairie remnants should make the entire complex of restored/remnant prairie more resilient (see earlier post on this subject).  Finally, selecting and altering restoration sites, when possible, to include topographic and other habitat type variation – and multiple examples of each type – can also help ensure the resilience of the resulting restored prairie. 

Designing management strategies for prairies that sustains ecological resilience is trickier because we still have much to learn.  We’re far from fully understanding the various stable states prairies may exist in, or flip to, let alone where the thresholds are between those states.  In addition, the level of plasticity, or range of adaptive capacity, of prairies is a subject of great debate right now among prairie ecologists – although the discussion is not usually framed in those terms.  The real question is – How much can prairies change their plant and animal species composition and still remain “in the bowl”? 

As an example, I manage a sand prairie that was hayed annually in the mid-summer for about 20 years before The Nature Conservancy purchased it in 2000.  Over that 20 year period, the plant community in that prairie adjusted to that annual haying regime.  Species such as stiff sunflower, leadplant, and sand cherry became restricted to a few steep slopes where hay equipment couldn’t go.  Early summer grass and forb species became very abundant, but later season flowering plants were less so because they were mowed off around their flowering time each year.  The prairie was a nice quality mixed-grass prairie, with good plant diversity, but definitely had the “look” of an annually-hayed prairie. 

Stiff sunflower has increased in abundance since annual haying ceased.

When we took over the management in 2000, we let the site rest for about 5 years and burned portions of it each year during that time.  Then, we began introducing some combined fire and grazing treatments at different intensities and at varying times of the season.  As a result, the prairie looks fairly different now.  Stiff sunflower and leadplant have spread considerably through the site, re-taking lower slopes where hay equipment had earlier eliminated them.  Cool-season grasses (native and non-native) change in abundance from year to year, but warm-season native grasses are certainly more dominant than they previously were.  Overall forb diversity is about the same as it was, but the relative abundance of many forbs has changed, and those abundances now vary from year to year, rather than remaining fairly stable.

In the context of plasticity, or adaptive capacity, this prairie has demonstrated that the 20 years of haying was not enough to move the plant community into a new stable state from which recovery, if that’s the right word, was not possible.  The community was altered by that haying regime, but upon alteration of that regime, the community composition morphed to match changing conditions – without losing plant diversity.  In other words, assuming that the prairie hasn’t lost anything critical during the 11 years of our management, both the haying regime and our current management have kept the prairie “in the bowl”, though it changed appearance fairly dramatically.  Its adaptive capacity is broad enough to include the “hayed” look and the “crazy Nature Conservancy management” look.  The real test of this, of course, would be to reintroduce haying for another 20 years and see if the plant community reverted back to something very similar to the condition it was in when we purchased it. 

This sand prairie has changed species composition substantially since we switched management from annual haying to a mixture of prescribed fire and grazing.

Here’s another example from my own experience.  We have a 45 acre restored prairie (prairie reconstruction) that was seeded in 1995 by Prairie Plains Resource Institute adjacent to a degraded remnant prairie.  The seed mixture included approximately 150 species of mesic prairie plants, most of which established successfully.  We managed the prairie with periodic spring fire for its first seven years, and then incorporated it into our experimental patch-burn grazing system (light stocking rate).  During nine years of patch-burn grazing management, (6 of which were during a severe drought) the plant species composition in any one place has bounced around quite a bit due to the fire/intense grazing/rest cycles imposed by the patch-burn grazing management system.  Overall, however, the prairie has maintained its mean floristic quality within 95% statistical confidence intervals (I collect annual data which entails calculating floristic quality within 100 1m2 plots and averaging the values across those plots).   Read more about our patch-burn grazing work and results at this restored prairie and others here.

That’s not to say the prairie hasn’t changed – it has.  Some plant species have increased in frequency among my annual data collection plots, some vary in frequency from year to year, and others stay fairly stable.  However, no species has dramatically declined over the time period.  (I wish I had data on other species, particularly insects, but I don’t.)  Perhaps the most interesting, and somewhat concerning, phenomenon has been an increase in the frequency of Kentucky bluegrass in my plot data.  The increase has been fairly steady over the nine years of patch-burn grazing and data collection, and bluegrass is now in about 75% of my 100 1m2 plots, though it rarely looks dominant where it occurs.  To this point, that increasing frequency doesn’t seem to be impacting the overall diversity or floristic quality of the plant community, but that doesn’t mean it won’t at some point.  Two possibilities are: 1) Our management is allowing bluegrass to enter the plant community but remain a minor component, or 2) Kentucky bluegrass is on a steady march of increasing dominance and will eventually turn my restored prairie into the same kind of low-diversity degraded prairie that exists in the adjacent remnant prairie.  I won’t be completely shocked by either scenario, but I have hope that #2 won’t happen because of the way bluegrass is acting in the community to this point.  It’s way too early to know for sure.

Interestingly, I have some large exclosures within this restored prairie that have never had grazing, only prescribed fire at a similar frequency to the grazed portion.  Those exclosures have very little Kentucky bluegrass in them – probably because of both the differing management and the fact that the exclosures are on the far side of the restoration from the neighboring bluegrass-dominated remnant prairie.  However, the exclosures also have much lower plant diversity and mean floristic quality than the grazed portion of the restored prairie.  Visually, they are dominated by warm-season grasses and a few large forbs (e.g., perennial sunflowers).  At this point, I prefer the grazed portion of the prairie because it seems to line up better with my management objectives of maintaining diverse and resilient plant communities (assuming it’s not slowly becoming a bluegrass wasteland). 

This photo was taken when the 1995 prairie seeding was nearing the end of its establishment period (fifth growing season). The prairie has maintained its mean floristic quality (and all of its plant species) through years of patch-burn grazing and drought.

To relate this example back to adaptive capacity, it appears likely that the grazed portion of the restored prairie has the adaptive capacity to retain its integrity as a prairie community through fairly wild fluctuations in species composition as a result of stresses from fire, grazing, and drought.  This is, again, remembering that I’m only evaluating the plant community and that the experiment is far from over.  On the other hand, it appears the exclosed portions of the prairie have lost plant diversity over time.  Whether the communities in those exclosures are still “in the bowl” or in a new stable state of lower diversity is a big question.  To address it, I’m going to open one of them to fire/grazing this coming year and exclude a portion of the currently-grazed prairie and see what happens.  If the two plant communities trade identities to match their new management regimes, I’ll know that both were still “in the bowl” and understand more about the adaptive capacity of our prairies.  If they don’t, that will be equally instructive!

The Upshot

Building and sustaining ecological resilience in prairies may be the most important component of prairie conservation in the coming decades.  Threats from invasive species, habitat fragmentation and detrimental land management practices, compounded by climate change, will make conservation extremely difficult.  Armoring prairies with ecological resilience gives us the best chance of success. 

In order to build that resilience, we first have to understand it better.  It is certainly more complex than the few simple examples I’ve provided.  There are numerous belowground processes and systems we still know relatively little about.  Even aboveground, there are many more questions than answers regarding the way species interact with each other and their environment – and what is required to maintain those interactions.  To gain a better understanding of these natural systems, we have to rely on experimentation and observation.  I think there are essentially two broad questions:

  1. What is the adaptive capacity of prairies, and where are the thresholds between the desired state and other, less desired, stable states?  This will certainly vary between tallgrass and mixed-grass prairies, and between sand prairies and black soil prairies, etc., but there will almost surely be some consistent themes. 
  2. How important is it to keep the ball moving within the bowl?  In other words, are prairies really like our human bodies, in that the more we stress and rest them, the better prepared they are for future stresses?  (Does the bowl shrink if we don’t keep pushing at the edges?)  Or do we just have to keep prairies from being stressed too far in any particular direction? 

 

We can work toward answering these questions with direct experimentation on prairies we manage (similar to my simple experiment with grazing and exclosures in the restored prairie example presented earlier).  In addition, though, we can learn much from prairies that “flip” to less desirable stable states (hopefully not the ones we’re managing!) by documenting as much as we can about what happens to them and why. 

Most importantly, I hope that thinking about ecological resilience with regard to prairies will make you look at the prairies you’re familiar with in a new way.  Seeing prairies as balls rolling around in a bowl makes watching and managing prairies a much different experience than seeing them as a stable “climax community.”  When we expect change, it’s easier for us to perceive change, and the more observant we are, the more we’ll learn.  And goodness knows we have plenty to learn.

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If you’re interested in learning more about ecological resilience, here are some relevant references that Craig Allen recommends (I’m pretty sure it’s just a coincidence that he’s a co-author on all of them).  The Gunderson et al. book reprints a lot of the classic / foundational papers on the subject.    

Sundstrom, S., C. R. Allen and C. Barichievy.  Biodiversity, resilience, and tipping points in ecosystems.   Conservation Biology: in review.

Gunderson, L., C. R. Allen, and C. S. Holling. 2010.  Foundations of Ecological Resilience.  Island Press, New York, NY.  466pp.  

Allen, C. R., L. Gunderson, and A. R. Johnson.  2005.  The use of discontinuities and functional groups to assess relative resilience in complex systems.  Ecosystems 8:958-966.

Forys, E. A., and C. R. Allen.  2002.  Functional group change within and across scales following invasions and extinctions in the Everglades ecosystem.  Ecosystems 5:339-347.

Peterson, G., C. R. Allen, and C. S. Holling.  1998.  Ecological resilience, biodiversity and scale. Ecosystems 1:6-18.