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Using Defoliation of Dominant Grasses to Increase Prairie Plant Diversity

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 In many prairies, the primary suppressors of plant diversity are dominant grasses – both native and non-native.  These grasses, left unchecked, can monopolize light, moisture, and nutrients to the point that few other plant species can coexist with them.  I’m not sure why some prairies suffer from this more than others.  There is some evidence that hemi-parasitic and allelopathic plants such as pussy toes, false toadflax, and wood betony can play a role in suppressing grasses and facilitating forb diversity, but I don’t think that’s the whole answer because I’ve seen very diverse prairie plant communities without those species – or with only a few scattered populations of them.  Regardless of the reasons, we are left with many prairies that have lost – or are losing – plant diversity through domination by grasses, and we have to decide what to do with them.  Some of those prairies are restored (reconstructed) prairies that started out with high plant diversity but have since lost much of that diversity.  Others are remnant prairies that have been degraded by overgrazing and/or broadcast herbicide application.  Still others are relatively diverse remnant prairies that are slowly losing diversity as individual forbs die without reproducing.

Big bluestem. A good native grass, but sometimes so dominant that overall plant diversity suffers.

I think one of the best tools we have for combating grass domination in prairies is defoliation – the removal of above-ground portions of plants.  Defoliation has always been a major component of prairie ecosystems through fire and herbivory, and mowing and (non-lethal “burn back”) herbicide applications are additional options at our disposal today.  Plants respond to defoliation in various ways, depending upon the severity of defoliation, the frequency and/or duration of the defoliating event, the stage of the plant’s growth at the time of defoliation, and each species’ genetic programming.  The way each plant, and its neighbors, respond to a defoliation event determines which plants will gain or lose territory.  In other words, defoliation influences the competition between plants – and manipulating competition between plants is really what most prairie management is all about. 

Some of the earliest research I’m aware of on the effects of prairie plant defoliation can be found in range management research from the 1950’s and 1960’s.  The earliest paper that is often cited from that era is by F.J. Crider, who documented the effects of defoliation on the root growth of grasses.  He (and others since) found that a severe defoliation of a grass plant resulted in an immediate cessation of root growth as plants reallocated resources from root growth to regrow leaves and stems.  More importantly, those grass plants actually abandoned sections of living roots as well – shrinking the total root mass of the plant fairly dramatically.  This makes sense, since the plant has to support those roots through photosynthesis, and a severe defoliation takes away most of the plant’s ability to photosynthesize. 

A simplified look at how dominant grasses can affect plant diversity. In the top example (A) grasses have monopolized both aboveground (light) and belowground resources (moisture and nutrients). After defoliation (B), both the above and belowground parts of the grasses have shrunk, freeing up resources and allowing other plants to establish within that lost territory. As the grasses recover from the stress of defoliation, their vigor and size increase, but the new plants have a fighting chance - at least for a while - to hold the new ground they've taken.

Those early range science research data provide some useful context for today’s prairie management, but those researchers were primarily trying to figure out the intensity of grazing they could employ while still maintaining a dominant stand of grass.  As prairie managers, by contrast, we want to reduce the dominance of grass to increase the diversity of other plants.  We can still learn from what those range scientists discovered; we just want to employ it in a different way.  Since plants primarily compete for light, moisture, and nutrients, we want to find ways to make those three kinds of resources more available to plants other than dominant grasses.  Defoliation can reduce shading aboveground (removal of leaves and stems), while simultaneously freeing up the availability of moisture and nutrients belowground (reduction of root masses).

In prairies where dominant grass species are suppressing plant diversity, we want to defoliate those grasses in a way that forces them to cede territory to other species.  In order for that to work, the first important factor is that the defoliation has to happen during the growing season.  Defoliating a dormant plant (e.g. with an early spring burn) doesn’t have any impact on its root system, which is a critically-important part of its competitive ability.  In order to force a plant to reallocate resources away from its roots, defoliation needs to take place after the plant has already invested significant resources in above-ground growth.  This is why a late-spring burn can have a significant (if temporary) impact on cool-season exotic grasses such as smooth brome.  Burning, grazing, or mowing grasses when they are just starting to flower has the biggest impact on most species because they have invested the maximum amount in their above-ground growth by that point.  Alternatively, repetitive mowing or grazing of grasses can also have a strong – and perhaps longer lasting – impact on their root systems because every time the grasses start to regrow, they get nipped off again, forcing them to regroup and reallocate resources time after time.

The immediate result of that kind of severe and/or repeated defoliation of dominant grasses is a release of opportunistic plants that thrive under low levels of competition.  This includes many annual and biennial plants, but also perennial plants that are built to move quickly into open space.  The quick flush of these plants often turns people off of defoliation because of a widely-held misperception that those “weedy” plants are outcompeting “good” plants.  In truth, the weedy plants are only able to grow because the competition that normally holds them in check has been suppressed.  When the defoliation event is over, the dominant grasses and other perennial plants will slowly recover their vigor – at which point the weedy plants will retreat and wait for another opportunity.  Rather than indicating a problem, I use the presence of weedy plants to tell me that my defoliation treatment has succeeded in weakening dominant grasses and has opened up space for other plants to take advantage of. 

As common as the overly-dominant grass problem is in prairie conservation, there is a frustrating scarcity of research that addresses it.  However, a recently-published research project by Kat McCain and others at Kansas State University provides some very nice insight into what can happen when dominant grass species are suppressed in a restored (reconstructed) prairie.  Kat and her colleagues studied plots of seeded prairie that had become heavily dominated by big bluestem and switchgrass over time.  They found that removing half or all of the big bluestem tillers (stems) – by clipping and herbicide application – from a plot significantly increased plant diversity.  Interestingly, removing switchgrass tillers in the same way had much less impact.  Following the removal of big bluestem tillers, the researchers saw increases in the vegetative cover of some forb species, including roundheaded bushclover (Lespedeza capitata),  pitcher sage (Salvia azurea), and blue wild indigo (Baptisia australis) within those plots, as well as new establishment of forb species including whorled milkweed (Asclepias verticillata), green antelopehorn milkweed (Asclepias viridis), leadplant (Amorpha canescens), roundheaded bushclover, and heath aster (Symphyotrichum ericoides).  In other words, suppression of big bluestem competition led to increased vigor among existing forbs and also allowed new plants to establish in the territory previously held by the dominant grass.  The study bolsters the theory that grass competition is suppressing forb diversity in many prairies, but also provides information on how plant communities might respond if that grass competition is reduced.

Of course there is a difference between simple defoliation and the kind of clipping/herbicide combination used by McCain and her colleagues.  In addition, most defoliation treatments (especially prescribed fire and haying) in prairie management are non-selective, meaning that all plants are simultaneously defoliated – not just the ones we want to suppress.  Uniform defoliation likely decreases some of the benefits of suppressing grass vigor because the vigor of the plants we hope will respond is suppressed as well.  However, there will be still be plants that can take advantage of the newly available light and soil resources following a uniform defoliation treatment, and by altering the timing of defoliations from year to year, we can ensure that a variety of species get the opportunity to respond.

Haying is a good example of uniform defoliation. Every plant gets cut at the same height and at the same time.

 

Ideally, though, we would like the ability to defoliate only those species that are suppressing plant diversity.  One way to do that is by using a selective herbicide such as Poast, which affects only grasses (not forbs, sedges, or other plants).  Poast is labeled for control of annual grasses, but at light rates can also provide short-term burn-back (defoliation) of perennial grasses as well, and some prairie managers have seen plant diversity increase following treatments.  Because it can kill annual grasses, and calibrating the appropriate application rate with the desired result can be tricky, it’s probably best to use this treatment on restored prairie rather than on remnant prairies for now – and to test it on small patches first. 

Another way to get selective defoliation is by the use of grazing.  In an earlier post, I described our use of patch-burn grazing in our Platte River Prairies as a way to increase and maintain plant diversity.  Patch-burn grazing is essentially a technique that uses patches of burned prairie within a larger prairie to attract grazing animals, concentrating grazing activity in those burned areas while allowing other areas to recover.  Under a light stocking rate, we find that cattle – even in the burned patches – are very selective about the plants they choose to eat.  Their top choice of grasses in the spring is smooth brome, and their summer favorite is big bluestem.  These happen to be two of the top three grasses that appear to stifle plant diversity in our prairies (the third is Kentucky bluegrass, which cattle like less well).  In our application of patch-burn grazing, a burned patch of prairie is normally grazed intensively for an entire season before the next patch is burned and cattle shift their attention to that.  That length of intense defoliation has significant impacts on the plants that are grazed – and again, under light stocking rates, the primary plants that are defoliated are smooth brome and big bluestem.  Interestingly, switchgrass is much less attractive to cattle and is often left ungrazed – or lightly grazed – in our prairies.  I found it intriguing (and encouraging!) that McCain and her colleagues found that switchgrass appeared to have much less impact on plant diversity than big bluestem did.

The effects of selective grazing in a restored prairie. This photo shows the burned patch with a patch-burn grazing system where a light stocking rate allows cattle to be selective about their eating preferences. Big bluestem is cropped very short, while other grasses and forbs are ungrazed or lightly grazed. Species such as hoary vervain aren't typically grazed even under high stocking rates, but many species such as purple prairie clover (front left), illinois bundleflower (middle left) and stiff sunflower (blooming) are commonly considered to be favorites of cattle - but only at higher stocking rates.

It would stand to reason that selective grazing of big bluestem and smooth brome would favor the expansion of the ungrazed plant species growing with those grasses.  While I’ve not had the time or resources to conduct much full-scale research (help wanted!), I do have data that supports that idea.  Through annual data collection of plant species frequency, I’ve found that the density of species (the number of plant species per 1m2 plot) increases by 20 to 30 percent in the year following the burn/graze treatment in a patch of prairie – in both restored and remnant prairies. 

Data from two prairies under patch-burn grazing. In both cases, the graphs show the number of plant species per square meter over time from the year of fire and intense grazing through the two subsequent years. The East Dahms site is a degraded remnant prairie and the Dahms 95 site is a restored prairie that was seeded in 1995 with over 150 plant species. The error bars represent 95% confidence intervals.

Of course, the increase I see in species density following grazing includes many plants such as ragweed and other opportunistic species, but I also see species like purple prairie clover (Dalea purpurea), Illinois bundleflower (Desmanthus illinoiensis), and stiff sunflower (Helianthus laetiflorus) respond as well.   In addition to seeing this in my plot data, I can walk out into the prairies and see seedlings of these species around the adult ungrazed plants.  Not all of those young plants survive their first year or two, but some permanent plot data I’ve looked at shows that at least some of them do.  By the way, I see similar post-grazing increases in plant species density and establishment of species like prairie clover under patch-burn grazing with higher stocking rates (less selective grazing) as well. 

As I said in my previous post on grazing, I’m not advocating that all prairies need to be grazed.  I’m not even advocating grazing as the solution to all prairies that suffer from overly-dominant grasses.  However, as we search for answers to address grass suppression of plant diversity, grazing certainly appears to be one viable alternative that is worth more investigation.  I’m continuing to experiment with variations in the way we employ fire and grazing treatments, and will keep learning as I go.  I’m also combining seed additions with those grazing treatments to see if I can take advantage of the open space created by defoliation to help establish new plants from seed – something that appears to happen rarely without some kind of suppression of surrounding vegetation.  I’m seeing some positive results, but it’s too early to know how well it will work long-term, and I’m still tweaking seeding rates and other factors.

Whether it’s grazing, prescribed fire, haying, or herbicide application, defoliation may be the most powerful tool available to help us suppress dominant grasses and increase plant diversity in prairies – where that is an issue.  The biggest obstacle to its application is probably the fear of causing damage to prairie by burning, cutting, or grazing plants during the growing season, but I think that fear ignores the resilience of prairie communities.  We still have a lot to learn about the most effective ways to apply defoliation to achieve our objectives, but the only way we’ll learn is by experimentation.  I hope you’ll join me in testing these methods and tracking the results.  If you do, please share what you learn with the rest of us so we can all work to figure this out.

Why Every Prairie Really is Unique – and Why it Matters.

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Confirmation that every prairie has its own unique composition of plant and insect species, and discussion about why that’s important for conservation.

Several years ago, I helped assemble a group of partners to begin some pilot research on what kinds of impacts habitat fragmentation may be having on the tallgrass prairies in southeastern Nebraska.  While those prairies are greatly fragmented compared to the extensive Nebraska Sandhills or Kansas Flint Hills prairies, there are still many blocks of hundreds to thousands of acres of prairie embedded within a matrix of cropland.  Those grassland complexes include a mixture of hayed prairies, grazed prairies, and Conservation Reserve Program (CRP) fields.  As conservation organizations consider the best ways to ensure continued survival of those important grasslands, many of the most pressing questions revolve around whether or not the prairie landscape is intact enough to support a functional ecosystem.

We still have a long way to go before we know much about the “functioning” of the southeast prairies, but we have collected enough data to know one thing:  every one of those prairies is unique.  While that not exactly earth shattering news, especially to biologists who have inventoried numerous prairies, it’s really very important.  To be more specific about what we’ve found, each prairie appears to have its own individual “signature” of plant and insect species compositions.  In other words, each has a mixture of species that is similar to, but also very different from, other prairies – even among prairies that are managed much the same way and that appear very similar from a distance.

For example, over the last couple of years we have done vegetation surveys in 24 hayed prairies as part of a couple of insect research projects.  We selected the 24 sites because we believed they were very similar to each other in plant community but varied in size and degree of isolation.  In other words, we tried to pick sites with identical plant communities.  Boy did we fail.  To be fair, the graduate student that selected the sites did so by driving gravel roads in April and May, and one hayed prairie looks a lot like another from the road at that time of year.  But we did try…

Staff from the Illinois Natural History Survey assisting with vegetation inventories in hayed prairies. Southeast Nebraska - Pawnee County.

There were some strong similarities among the prairies.  Most of them had had abundant cool-season grasses, but also strong representation from warm-season native grasses.  In addition, all of them had a good diversity of plant species – most averaged between 20 and 25 plant species per square meter.  Many of the more common wildflowers were found in all, or nearly all, of the prairies, and in similar amounts – including species like stiff sunflower (Helianthus pauciflorus), heath aster (Aster ericoides), black-eyed Susan (Rudbeckia hirta), pussytoes (Antennaria neglecta), and wild alfalfa (Psoralidium tenuiflorum).  Finally, the majority of the prairies had apparently been inter-seeded at some time in the past with non-native legumes such as red clover, sweet clover, and others, and those species were moderately to very abundant in those prairies – although not to the extent that they seemed to have an impact on overall plant diversity.

However, the differences among the prairies were as obvious as the similarities.  For example, of the 24 prairies, prairie phlox (Phlox pilosa) only appeared within our sampling plots (50 1m2 plots at each site) in three of them – but it was very abundant at those three sites.  Why is it abundant at a few sites and rare or missing from others?  Prairie blazing star (Liatris pycnostachya) showed a similar pattern.  Other important wildflower species – those considered to be among the core species of tallgrass prairie plant communities – only appeared in the data from about half of the sites.  Those species included compass plant (Silphium laciniatum), purple prairie clover (Dalea purpurea), hoary puccoon (Lithospermum canescens), and wild strawberry (Fragaria virginiana), among others.  To be clear, these data don’t indicate whether or not species were absent from an entire site – only that they were absent from the 50 1m2 plots we sampled from at each. Regardless, we saw significant differences in the relative abundance of those species from site to site.  Other important species that varied wildly in abundance among the prairies included prairie dropseed (Sporobolus heterolepis) and eastern gamagrass (Tripsacum dactyloides).  Those two species were in most of the prairies, but appeared in nearly 100% of the plots in some prairies and in only one or two plots in others.

Prior to initiating our research project that used those 24 hayed prairies, we did some broader surveys of prairies that included both hayed and grazed prairies to assess the degree of variety within plant and insect species at those sites.  (The grazed prairies, by the way, reflected many different kinds and intensities of grazing, but are lumped together here for simplicity.)  Because it was pilot data, we did some quick inventories of plants at each site, trying to list all the species we could find.  Not surprisingly, there were some stark differences between grazed and hayed prairies in their plant compositions.  In a few cases that variation could be explained by site conditions (such as rocky or steep terrain) because those conditions clearly made grazing more feasible than haying – in those cases plant species may have been responding to the site rather than the management.  In most cases, however, it was clear that the plants were responding to management.  Plant species like false gromwell (Onosmodium molle), tall boneset (Eupatorium altissimum), partridge pea (Chamaecrista fasciculata), and hoary vervain (Verbena stricta) have life strategies that allow them to thrive under grazing because they are less palatable to grazers than their neighbors and/or can respond more quickly to the space opened up by grazing.  Those and other similar species were found almost exclusively at grazed sites.  On the other hand, there were species like New Jersey tea (Ceanothus herbaceus) and golden alexander (Zizea aurea) that were found much more often at hayed sites than grazed.  While the majority of plant species were found across the entire range of site types, their relative abundances showed that most responded much better to either the hayed or grazed conditions.

Hoary vervain is a wildflower found mostly in grazed prairies in southeastern Nebraska, and one that is extremely important to pollinators, including regal fritillary butterflies.

Within those same grazed and hayed prairies, we also did some sweep netting of insects.  We targeted our sweeping so that we collected insects off of the same plant species across all of the sites (using a couple different plant species).  We wanted to see whether the insect communities using a particular plant species were essentially the same at all of the prairies.  They’re not.  What we found was that the insect data were even more chaotic looking than the plant data.  Very few insect species were found at all, or even the majority of sites.  That’s likely due to the relatively small amount of sampling we did.  But even with our small samples, it was clear that a large number of species were abundant at one site but absent from many others.  The other strong pattern was that the total abundance of insects was often quite a bit higher – among many species – at the sites we’d considered to be “lower quality” based on our plant data.

There are a number of reasonable – even obvious – explanations for these variations among prairies, including soil type, topography, past management, random chance, etc.  But more important are the lessons that come from recognizing those differences.  First, I think the differences demonstrate the importance of management.  Surely some of the differences between the vegetation composition in those 24 hayed prairies stem from variations between sites relative to the timing and frequency of annual haying, the consistency of timing and frequency over many years, and whether or not the prairies are fertilized.  Later season haying, for example, allows more (and different) plant species to complete their life cycle prior to cutting than early season haying (or twice-annual haying).  Completing their life cycle now and then is critical for the long-term survival of plants because it allows them to reproduce and store energy for the coming winter.

The existence of a combination of hayed and grazed prairies within the landscape is also important.  Because there are plant species (and likely insect species) that do better under grazing management than haying management, and vice versa, having both types of management within a neighborhood can maintain more biological diversity than either of the management types alone.  Of course, managing each of those prairies with a combination of fire, grazing, and perhaps haying, might allow even more plant and insect species to persist on each prairie – although the local abundances of some species might go down because they would lose the consistent management that had allowed them to become dominant.

From a conservation standpoint, one big lesson is that it’s not sufficient to simply rank prairies by floristic quality, size, or other measurements, and act to “protect” the best of those.  In order to conserve the complete diversity of life in the southeastern Nebraska prairies, we need to keep as many of the remaining prairies as we possibly can.  While the grazed sites might look less attractive to a botanist driving by compared to hayed prairies full of compass plant and blazing star flowers, the aesthetic preferences of botanists don’t necessarily reflect the biological needs of ecosystems.  While there was surprising variation between the hayed prairies we looked at, a very significant degree of biological diversity would be lost if the grazed prairies were plowed up.

Even if it were the desired strategy, there’s no way conservation organizations would or could buy up a significant number of the grasslands in southeastern Nebraska, so working collaboratively with the landowners of those prairies is of the utmost importance.  Relying on private landowners to conserve prairies has been a tough pill for many conservation organizations to swallow – including the one that employs me – but during the last several decades most have recognized that need.    The next trick is to refine our skill at working with those landowners without coming across as know-it-all biologists.  The most successful private lands liaisons from the conservation community have been those who listen well, recognize the importance and value of individual landowners’ priorities (remembering that it is THEIR land), and build relationships on mutual trust.  Through those relationships, we can alert landowners to aspects of their prairies and management options they might not be aware of, but the decisions about how to manage their sites still have to be left to the landowner.  Clearly, that will result in some prairies being managed in ways that make some of us uncomfortable, but we don’t want all the prairies to look the same anyway, right?