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.

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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.

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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.

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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.

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

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

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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.

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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%.

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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.

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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.

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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.

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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.

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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.

Tale of two Highlands Part I: Horton Plains, Sri Lanka

This post is contributed by Dr. James Aronson, a restoration ecologist at MBG’s Center for Conservation and Sustainable Development, and his son Thibaud Aronson. James is also a researcher with the CNRS (National Center for Scientific Research) in Montpellier, France.

In Sinhalese Sri Lanka means “Resplendent Isle”, a fine name indeed for this tear-shaped island off the coast of southeastern India, just north of the equator. Last month I travelled with my son on a self-guided Natural History + Ecological Restoration visit, we are finding and photographing cloud forests and birds galore, like the endangered endemic Sri Lanka whistling thrush, Myophonus blighi, and the Kashmir flycatcher, Ficedula subrubra, which over-winters exclusively in the Sri Lanka highlands, from its very restricted breeding grounds in Kashmir, northern India.

We were also looking at the mosaic of grasslands, cloud forests, and lowland forests we find here from a restoration ecology perspective.  That means we’re trying to “read” the landscapes we see in terms of known transformations carried out during the British colonial era (1815 and 1948, when Sri Lanka was known as Ceylon), and since independence. The remarkable Horton Plains National Park is a mosaic of montane grassland (ca. 35%) and cloud forest (ca. 65%), encompassing the headwaters of three major rivers. It was declared a sanctuary in 1969 and elevated to national park status in 1988; it became part of a large UNESCO World Heritage site in 2010. In the central highlands of Madagascar, grasslands appear to occupy about 99% and most people assume they are anthropogenic…. This month, I’m travelling with Leighton Reid in the Central Highlands of Madagascar, and we will be blogging about this soon.

But, the history of preservation in the highlands here goes back a lot further, to the days when the Isle was part of the British empire, along with all of India. According to information we gathered at the extraordinary, and poorly known Hakgala Botanic Gardens, the great English botanist and explorer Joseph Dalton Hooker had advised the British government to leave all montane forests above 5000 ft. (ca. 1300 m) above sea level “undisturbed” and after 1873 the administration prohibited clearing and felling of forests throughout the central highlands. What a great idea that was! It is too bad there were not enlightened laws on hunting of wild animals as well. One Scottish officer in colonial service in Sri Lanka bragged he had shot and killed over 1400 elephants in Horton Plains and nearby. Today, there are none left there and, so far as we could determine, no plans to reintroduce them from the other remarkable parks, including Yalla and Uda Walawe….

So, what is the significance of the absence of elephants in this park? And, what else can we learn from past regimes and historic periods in Sri Lanka? For starters, we discover that conservation, and respect for other organisms goes back much further than the 19th century. Consider the sign at the entrance to Udawattakele Forest Reserve, near Kandy, one of the historic capitals from the long period of successive kingdoms the island had known prior to the European colonial chapter in Sri Lanka’s history:

O Great King, the birds of the air and the beasts have an equal right to live and move about in any part of this land as thou. The land belongs to the peoples and the other beings and thou are only the guardian of it.”

-Arahath Mahinda (a son of the emperor Asoka the Great, who brought Buddhism to Sri Lanka)

How would it be if we could revive that approach to the Web of Life in our own day and age?

So, what has Horton Plains National Park, with its grassland-forest mosaic, its tourists, and its absent elephants got to do with the Central highlands of Madagascar? For one thing, we can see that fire is a big ecological driver in both areas. The abundant arborescent Rhododendrons in Horton Plains tell a vivid tale in this regard.

Rhododendron arboreum subsp. zeylanicum at Horton Plains National Park. It appears to be fire-resistant and is the only tree species present in large areas of grasslands subject to fire.

Rhododendron arboreum subsp. zeylanicum at Horton Plains National Park. It appears to be fire-resistant and is the only tree species present in large areas of grasslands subject to fire.

On the grand scale of things, Sri Lanka’s Central highlands also resemble those of Madagascar’s since both are the crowns of a poor, emerging tropical island with small and very similar human population size (21 million vs. 24 million), despite being much nearly ten times smaller, and with over 30,000 years of human history, as compared to merely two millennia for Madagascar.

Horton Plains also has remarkable conservation value both for its biodiversity and the ecosystem services it provides to people. Also, as I said, it’s a mosaic of grasslands and cloud forest, that in the past was certainly much affected by both elephants and fire.

Finally, both Sri Lanka (along with the Western Ghats of southern India) and Madagascar count among the world’s biodiversity hotspots, easily visible in their fauna and flora, which is one of the main reasons why MBG researchers, and many others travel and work in Madagascar.

Now, let’s turn back to fires. A big fire hit Horton Plains in 1998, and there are serious invasions of two noxious, cosmopolitan weeds, namely Gorse and Bracken fern. Some control work is underway on the Gorse, but the Bracken fern is apparently not seen as being a problem. Rainbow trout were introduced in the 19th c. and apparently have displaced all native fish, and are taking a toll on native shrimp and no doubt other fauna.