A ten-year woodland restoration trajectory

Leighton Reid describes a long-term ecological research project at Shaw Nature Reserve (Franklin County, Missouri, USA). To learn more, read the new research paper (email the author for a pdf copy – jlreid@vt.edu) or tune in for a webinar from the Natural Areas Association on April 21 (register here).

In 2000, the Dana Brown Woods were dark and dense. Brown oak leaves and juniper needles covered the sparsely vegetated ground, and invasive honeysuckle was creeping in around the edges. Biologically, the woodland was getting dormant.

In contrast, the woods today are lit by sunlight everywhere except the lowest-lying streambanks, and the ground is hardly visible beneath a green layer of diverse, ground-level foliage. These changes were most likely caused by two actions: burning the woods, and cutting out invasive trees and shrubs.

Many practitioners have seen woodlands recover to some extent when they are burned, but few have documented the recovery as thoroughly and over so long a period of time as Nels Holmberg and James Trager.

Nels Holmberg (left) discussing the finer points of Rubus identification with Quinn Long in the Dana Brown Woods.

Nels is an ecologist and sheep farmer in Washington, Missouri. He has inventoried the plants at several state parks and natural areas. In 2000, Nels teamed up with Shaw Nature Reserve’s resident natural historian, James Trager, and together they designed a study to describe how ecological restoration was changing the woodland flora at the reserve. They picked the Dana Brown Woods as their study area.

In a nutshell, Nels and James chose 30 random points on a map. They divided the points evenly across three ecological communities. They placed 10 points in mesic woodlands – the gently sloping parts of the property where white oak and shagbark hickory were most prevalent. Ten points were in areas dominated by eastern red cedar – mostly thin-soiled ridgetops that faced the south, and ten points were in forest – the lower, thicker-soiled toe slopes where northern red oak and Shumard oak were dominant in the canopy with paw paws and spicebush down below.

Three ecological communities in the Dana Brown Woods: (A) red cedar dominated areas which, after removing red cedar, looked more like dolomite glades in some parts; (B) mesic woodlands with lots of oak and hickory in the canopy; and (C) forest – which had a much darker understory.

At each point, Nels hammered in a t-post, then walked 50 m in the steepest direction and hammered in another t-post. This was his transect. Every year for more than a decade (2000-2012), Nels walked the transects and recorded every stem of every species that was inside of 10 0.5-m² study plots. Actually, he did this twice per year – once in the spring to capture the ephemeral plants, and once in early summer. Over the course of the study he spent more than 200 days in the field.

Dana Brown Woods before (left) and after (right) red cedar removal, with Nels’s 30 transects. The horizontal axis of the image is about 0.9 km. Imagery is from Google Earth.

During this time the stewards at Shaw Nature Reserve were busy restoring the woods. From 2001-2012, they burned the woods five times. This amounted to about one fire every three years. In 2005-2006, they brought in a logging crew to remove all of the eastern red cedars.

James Trager lights a fire in a woodland at Shaw Nature Reserve.

One of several thousand red cedar stumps from trees that were harvested from the Dana Brown Woods in 2005-2006.

One of Nels’s sampling quadrats in the Dana Brown Woods. Photo: Nels Holmberg.

I met Nels and James in 2014. I had just joined Missouri Botanical Garden’s Center for Conservation and Sustainable Development as a postdoc, and I was looking for a local research project. I heard that Nels Holmberg had a giant dataset about woodland restoration, so I called him and asked if I could look at it. Nels said “Sure!”. I imagined he would send me an Excel file. Instead he brought in a giant cardboard box full of yellow legal pads where he had recorded his data.

One of hundreds of datasheets where Nels recorded his detailed observations.

It took a long time to digitize all of the data. There were more than 50,000 data points. But once we had it all together, this is what we learned:

After eleven years of restoration, the number of native plant species in Dana Brown Woods increased by 35%, from 155 species in 2001 to 210 species in 2012. This increase was linear. That is, the number of native species was still increasing at the end of the study. If we repeated the study today, we expect the number of native species would be even greater than in 2012.

The number of native species increased at different speeds and to different degrees in different ecological communities. In the lower and wetter forest areas, the numbers didn’t really shift very much. They jumped around but not in one direction. In the woodland areas, the number of native species increased by about 23% in the first three years and then leveled out. But in the higher and drier areas where red cedars had been dominant, the number of plants increased linearly by 36%.

Changes in the number of native plant species recorded over time in the Dana Brown Woods. On the left are overall changes for the whole management unit. On the right are changes for different ecological communities within the management unit. The management interventions are shown in gray.

The plant species that benefited from the restoration were mostly forbs and grasses. A couple of the biggest “winners” were black snakeroot (Sanicula odorata) and nodding fescue (Festuca subverticillata). There were also some “losers”: Virginia creeper (Parthenocissus quenquefolia) and spring beauty (Claytonia virginica) both declined over time. Relatively few of the species that became more common were “conservative” – i.e., dependent on intact habitat. Mostly they were more widespread and tolerant species.

Co-author Olivia Hajek demonstrates a hog peanut (Amphicarpaea bracteata) – a good representative of the type of species that benefited most from the restoration. Hog peanut is an herbaceous legume that is common in many woodlands, including disturbed ones.

Our study did not include a control treatment, but counterfactuals exist at Shaw Nature Reserve (although they are becoming fewer and fewer with the excellent stewardship of Mike Saxton and many others). There are still thick patches of eastern red cedar covering remnant glades on parts of the property. Woodlands that have not been regularly burned are now filled with bush honeysuckle (Lonicera maackii), wintercreeper (Euonymus fortunei), and other invaders. And low-lying forest that has not been restored is very dark with fire-intolerant sugar maple (Acer saccharum) casting much of the shade. If we had included a control treatment in our experiment, these are probably the trends we would have found – definitely not a spontaneous resurgence of diverse native plants.

Fragrant sumac (Rhus aromatica) was present at the outset of restoration and remained relatively stable.

Why does this work matter? The biggest value of this study is that it shows a relatively long-term restoration trajectory, and it does so in fine botanical detail. Many managers and scientists already have data to show that fire and tree thinning increase woodland plant diversity. This study adds another dimension. It shows how quickly plant diversity recovered. It also shows how the speed and shape of the recovery varied across the landscape. We hope that other scientists and practitioners will compare the recovery trajectories in the Dana Brown Woods to their own natural areas. To facilitate that, we have made all of the underlying data freely available online.

Buffalo clover (Trifolium reflexum) is a conservative species that is present in Dana Brown Woods but was not detected in any of the survey plots.

One of the next steps for this research is to figure out how and when to re-introduce some more conservative plants. Although the Dana Brown Woods became much more diverse as it was being restored, most of the plants were early successional or generalist species. We found very few habitat specialists that cannot tolerate disturbance, which suggested to us that some of these species may have been lost from the site at some time in the past. To learn how conservative plants might be re-introduced, we have started a new experiment testing the effects of soil microbes, competition, and time since the start of restoration on the success of introduced seedlings from seven conservative plant species. In the next year or two, we hope to have new information and recommendations for restorationists looking to add more specialized biodiversity to their woodlands.

Freemont’s leather flower (Clematis fremontii) is a restricted species occurring on dolomite glades in southeastern Missouri. Although it is present at Shaw Nature Reserve less than one kilometer from Dana Brown Woods, it has not colonized the restored glade habitats there. This photo is from Valley View Glade near Hillsboro, Missouri.

To learn more about this research, you can read the original research paper in Natural Areas Journal. Email me for a pdf copy (jlreid@vt.edu). You can also tune in on April 21 for a webinar on this work. Register here.

How does fire affect ant-mediated seed dispersal?

Eva Colberg describes her ongoing research at Shaw Nature Reserve. She is a Ph.D. student in the Biology Department at the University of Missouri St. Louis.

In the late 1940s, Ohio-born entomologist Mary Talbot spent her days crouched in the woods of St. Charles, MO, tracking ant activity in painstaking detail through the seasons. Similarly, last summer I tried my hand at watching ants in the woodlands of Shaw Nature Reserve, with the addition of crumbled pecan shortbread cookies and the help of my field assistant, Dayane Reis. Foraging ants flocked to the buttery feast, the contrast of the crumbs’ sandy color against dark soil and leaf litter allowing us to easily follow the cookie thieves back to their nests.

A plot of flagged ant nests (found by following cookie-bearing ants) in the Dana Brown Woods, one of the management units at Shaw Nature Reserve.

We watched at least seven different species of ants run off with the cookie crumbs, but I was most interested in the winnow ant (Aphaenogaster rudis). Reddish-brown, long-legged, and narrow-waisted due to a double-segmented petiole (the connection between the abdomen and thorax), the winnow ant worker is an elegant lady. She is also remarkably swift-footed and strong, adept at carrying chunks of pecan cookie or naturally occurring analogs.

A winnow ant (Aphaenogaster rudis) worker, with the petiole and post-petiole that give the species its svelte waist. From her head to the end of her abdomen, this ant is about 4.5 mm long.

To an ant, a cookie more or less resembles an insect carcass, a staple of many ant diets. Chemically and nutritionally, the seeds of many of Missouri’s spring-flowering herbs also resemble a delicious dead insect (or cookie). From an ant’s point of view, this means food for larvae. From a seed’s point of view, this means dispersal. Hitchhiking to an ant’s nest gives the seed a new location to germinate and grow away from the parent plant, and potentially a multitude of other benefits such as escape from predation or better soil conditions. In any case, this is ant-mediated seed dispersal, or myrmecochory.

A field ant (Formica subsericea) grabs a bloodroot (Sanguinaria canadensis) seed by its elaiosome, the oily, nutritious appendage that most resembles a dead insect and attracts ants.

In other parts of the world, benefits of myrmecochory include enhanced survival and germination after fire. In arid, fire-prone areas of both Australia and South Africa, ants bury seeds deep enough to buffer the intense heat of fire, but shallow enough that the heat weakens the seed coat and increases the odds of germination. Thus, the ants protect the seed from the flames while still providing exposure to a Goldilocks level of heat.

Just as in Australia and South Africa, fire is (or was, and with the help of land managers is once again becoming) also a frequent occurrence in Missouri. At Shaw Nature Reserve, managers use prescribed burns to restore an open structure to the reserve’s oak-hickory woodlands. But, is ant-mediated seed dispersal interacting with fire the same way here as in those other fire-adapted ecosystems?

This is a key question of my dissertation research at University of Missouri St. Louis. Using cookies to find winnow ant nests last summer helped me test methods and plan out my experiments for this coming year. Specifically, I will be tracking where the ants take their seeds, whether ants disperse seeds more or less in the year after a fire, and whether the presence and timing of surface fire affects the germination of the seeds after dispersal. Stay tuned!

You can keep up with Eva Colberg on Twitter (@ColbergEva) or by checking out her science communication initiative Science Distilled STL.

Seed Banking for Conservation and Restoration

Meg Engelhardt is Missouri Botanical Garden’s Seed Bank Manager. She describes her ex situ conservation program and its applications for ecological restoration.

Seeds have been stored for food since the dawn of agriculture, but in recent decades seed banks have become an increasingly relied upon tool for plant conservation. Ideally we would conserve all plants in their natural habitats, or in situ. After all, when plant populations remain intact so do the relationships with other organisms in their ecosystems. Intact plant populations also maintain gene flow within the species, helping populations continually adapt to their surrounding environments. Unfortunately, it is not always possible to maintain wild plant populations. Even when resources are available, maintaining natural plant populations may be impossible due to habitat fragmentation or destruction, range shifts due to climate change, pollinator loss, or any list of known or unknown factors resulting in population decline. This is where seed banking, or ex situ conservation, may play a supporting role.

Collecting seeds from a dolomite glade in Franklin County, Missouri

Seed banks are long term storage facilities designed to keep seed viable for years and even decades. Those seeds can then be used for research, restoration, reintroduction, or education.

Maybe you have heard of the Svalbard “doomsday” Seed Vault, where seeds of more than 4000 plant species are stored deep below the permafrost near the north pole. Or perhaps the Millennium Seed Bank, which holds 13% of the world’s wild plant species and continues to collect the world’s threatened flora. Here in the United States we have the Native Plant Germplasm System, a network of 20 storage facilities across the country that store over 15,000 species, with a focus on agriculturally important species. On a more local scale, many small seed banks are used to conserve regionally important, threatened species.

Inside a very large freezer at the National Center for Genetic Resources Preservation in Ft. Collins, Colorado

The Missouri Botanical Garden has been seed banking for over thirty years. In 1984 the Center for Plant Conservation (CPC) was founded with Missouri Botanical Garden as a founding member. CPC is a network of 40 botanical institutions focused on ex situ conservation of rare plant material while also ensuring material is available for restoration and recovery efforts. Our CPC collection is currently maintained by staff who are actively seed banking, researching, and restoring populations of extremely rare native plants throughout southeastern US (Solidago ouachitensis, for example).

Additionally, seed collecting and short term seed storage has been going on for at least 25 years at Missouri Botanical Garden’s Shaw Nature Reserve. Horticulture staff at the Reserve are focused on local ecotype native plant horticulture and have been collecting seed from regional wild sources for use in small scale greenhouse propagation, use in the Whitmire Wildflower Garden, restoration projects throughout the Reserve, and various other partnerships that encourage native plant horticulture.

The Shaw Nature Reserve “seed closet” currently houses almost 500 different taxa ‒ most of which are local wild source (i.e., seed collected from plants growing in the wild) or second generation seeds (i.e., the first descendants of plants growing in the wild).

In 2013 the Missouri Botanical Garden Seed Bank was created with two main goals. First to advance seed banking at an institutional level by providing support and facilities. A new seed lab space was created at Shaw Nature Reserve which includes lab benches and space for processing collected seed and cleaning for storage as well as a refrigerator and freezer storage space. The second goal is to collect and store samples of Missouri’s entire flora, which includes roughly 2,055 taxa. Continually collecting and storing samples of all local species will ensure long term genetic conservation that can be made available for research, restoration, and recovery should the need arise.

Missouri Evening Primrose, Oenothera macrocarpa

Can the Ozark chinquapin successfully re-colonize interior highland forests?

Jenn Rosen is an undergraduate at the University of Missouri – Saint Louis. This summer, she participated in the Missouri Botanical Garden’s Research Experience for Undergraduates (REU) program. She spent eight weeks working with scientists in the Center for Conservation and Sustainable Development. Here, Jenn writes about her field experiment at Shaw Nature Reserve.

Conservationists and ecologists have recently sought to restore an ecologically important tree, the Ozark chinquapin (Castanea ozarkensis), that became threatened not by the effects of clear-cutting, but instead due to an incurable parasitic fungus called chestnut blight (Chyphonectria parasitica). After devastating its cousin (American chestnut) in eastern forests, the fungal disease moved west and began infecting chinquapin trees in the Ozark and Ouachita Mountains. The blight is presumed to have been brought into the United States by accident in the early 1900s when Chinese chestnuts (C. mollissima) were imported into the country. What makes the Ozark chinquapin and the American chestnut so susceptible to the disease is that the fungus’ wind-borne ascospores can easily enter through small cavities in their bark where they then grow and spread to neighboring trees. The blight causes Ozark chinquapin trees to die back to the roots, from which multiple stems resprout to form a large shrub-like growth form. Once a widely dispersed canopy tree in upland Interior Highland forests, chinquapins declined in abundance and are now often found as multi-stemmed, blight-infected subcanopy shrubs.

Castanea ozarkensis seeds, with white radicles, from two maternal lines. One group’s seeds are twice the size of the others. Large seeds hold more resources, which can give seedlings a head start. But larger seeds also face a higher risk of being preyed upon by rodents.

Chinquapins were prized by many folks of the Ozarks for the nutrient-rich nuts they produced, and its rot-resistant woods were used to make furniture, railroad ties, and fence posts, among other products. Additionally, large mammals such as black bears were known to forage in the Ozarks in search for the nutritious nuts to help replenish their fat reserves for the upcoming breeding season. There is a collaborative effort among groups to restore the chinquapin throughout its’ former range once blight-resistant seeds become widely available, estimated to take approximately 20 – 30 years.

Once blight-resistant seed becomes available, understanding how chinquapin trees successfully regenerate in the wild will be the key to successfully restoring this species. Unfortunately, little is known about the ecology of the tree prior to blight infection. Like other nut-bearing trees in Ozark woodlands (e.g., oaks), we suspect that there may be several limiting factors to chinquapin seedling recruitment, such as seed predation, poor soil quality and/or poor light availability. In order to test these predictions, we planted 320 chinquapin seeds (from two distinct maternal origins from the wild) across ten experimental replicates at the Shaw Nature Reserve (Gray Summit, Missouri). The criteria for each replicate was that there had to be a shrub microhabitat, which was dominated by the common understory tree, eastern redbud (Cercis canadensis), and an open habitat, separated by at least three meters. Like other Ozark woodlands, prescribed fire is currently being used to restore woodlands at the Shaw Nature Reserve and will likely be a key component to successfully reintroducing chinquapins back into the wild. After enduring many scrapes and pricks from constructing mammal exclusion cages for half of the seeds, roughly two weeks after planting the seeds we had a bounty of little chinquapin seedlings emerge.

Castanea ozarkensis seedling, protected from marauding rodents by a wire cage.

We found that consumer treatment and microhabitat structure, as expected, influenced the rates of Ozark chinquapin seed emergence. Nearly all of the seeds that were exposed to small mammals were eaten or removed, even though seeds were buried a few centimeters in the soil. Interestingly, small mammals consumed seed at greater rates in shrub than open microhabitats. These results imply that understory vegetation structure determines where chinquapin seedlings can successfully recruit through its influence on small mammal behavior. Also, small mammals removed larger, and presumably more nutritious, seed at greater rates than smaller seed. Environmental factors, like light availability, did not affect seedling growth, but the short duration of the study may not have been adequate for the potential influence of these factors to become apparent.

A rodent successfully consumed this unprotected nut.

A thwarted attempt to breach a protective cage.

Our work contributes to a larger ongoing project by the Ozark Chinquapin Foundation to restore and conserve Ozark chinquapins. Given the high rates of seed removal, future restorationists will have to transplant chinquapin seedlings as opposed to seeds to successfully reestablish this species in the wild. However, once reintroduced seedlings grow to maturity and produce seed, our study suggests that microhabitat structure in Ozark woodlands will play a key role in determining the recruitment and growth rates of restored populations.