The ephemeral forests of southern Costa Rica

Damaged ecosystems don’t recover overnight, but sometimes that’s all the time that they get. CCSD scientist Leighton Reid describes new research about tropical secondary forests in southern Costa Rica, including how long these young forests last, what’s at stake, and how we can keep them around longer.

Regrowing tropical forests on marginal farm lands is one of the main ways that humans can prevent runaway climate change. With ample moisture and long growing seasons, tropical trees often can grow quickly and pull large amounts of carbon out of the atmosphere, storing it in their wood and keeping it from trapping heat. At the same time, young forests provide habitat for plants and animals and improve water quality for humans, among many other benefits.

But even in a moist, tropical climate, trees don’t grow instantly. Typically, it takes many decades for a recovering forest to stock up all of the carbon that it can hold. And it can take even longer for some plants (like orchids) and animals (like antbirds) to return. If a forest starts to grow back, but then someone cuts it down again, these time-dependent benefits never accrue.

In other words, the hopes and expectations that many people have for young tropical forests depend on young tropical forests growing old. So do they? Our new study suggests not.

San Vito & Coto Brus Valley

The Coto Brus Valley and Talamanca Mountains in southern Costa Rica. Photo by J. Leighton Reid.

To find out how long secondary forests persist, I teamed up with Matthew Fagan, a landscape ecologist at the University of Maryland Baltimore County, and Rakan Zahawi, director of the Lyon Arboretum, as well as two students, James Lucas at Washington University and Joshua Slaughter at UMBC.

We studied a set of historical, aerial photos from southern Costa Rica, which covered the time period from 1947-2014. Previously, Zahawi and colleagues had classified which areas in each photo were forest and which areas were farms or other non-forest land uses. By comparing the maps they made for each year, we were able to see where and when new forests appeared and how long they remained as forest before they were converted to some other land use (mostly farms).

The young forests did not last long. Half of the new forests disappeared before they were 20-years old. And 85% were cut down before they were 54-years old. Larger forests and forests near rivers lasted longer.

One hectare forest fragment, Coto Brus, Costa Rica

An isolated forest fragment surrounded by cattle pastures in southern Costa Rica. Photo by J. Leighton Reid.

First, the bad news. Twenty years is not even close to the amount of time it takes for a young forest to become as diverse as an old-growth forest. For example, vascular epiphytes like orchids and bromeliads take more than 100 years to fully recover in young forests.

Carbon storage will also take a hit. If forests elsewhere in Latin America are as ephemeral as forests in southern Costa Rica, then carbon stocking over the next thirty years may be reduced by an order of magnitude.

Ephemeral forests could just be a problem in Costa Rica, but another study shows that secondary forests in eastern Peru have even shorter lifespans. There, secondary forests are cleared at a rate of 3-23% per year. Compared to that, the 2-3% per year rate of loss in southern Costa Rica is considerably better. And that’s not a good thing. Clearly we need more research on secondary forest persistence from other places.

There is some good news, though. Even though many new forests were short-lived, the ones that survived were predictable. And if we can predict where new forests will survive, we should also be able to help them survive longer. Larger forests and forests close to rivers were cut down less often than small forests and forests far from rivers. This suggests that restoring large, riparian forests could be a smart investment.

Gulfo Dulce from Fila Cruces - Coto Brus, Costa Rica

Forests and cattle pastures in southern Costa Rica. Photo by J. Leighton Reid.

Governments and other organizations can also help forests persist by creating incentives for long-term forest management, providing resources to enable long-term management, and ensuring that local people will be able to enjoy the benefits that old forests provide.

We hope that this work will lead to stronger restoration commitments. Right now, dozens of countries are setting big targets for forest restoration. For example, in 2012 Costa Rica committed to restore a million hectares of degraded land by 2020 (an area about one fifth the size of the country). There is a great opportunity for Costa Rica and other ambitious countries to plan for long-term forest restoration.

If we can begin to restore a million hectares of forest by 2020, why not plan to restore a million hectares of 100-year old forest by 2120?

Melissa's Meadow, Las Cruces Biological Station, Costa Rica

A trail through secondary forest at the Las Cruces Biological Station in southern Costa Rica. Photo by J. Leighton Reid.

For more information on this research, you can read our open-access paper in Conservation Letters or watch a video of Leighton Reid presenting to the Association for Tropical Biology and Conservation back in June. Additional papers on restored ecosystem persistence are available here and here. This work is a product of the PARTNERS (People and Reforestation in the Tropics: a Network for Research, Education, and Synthesis) Working Group on Spatial Prioritization. Funding was provided by grant DEB-1313788 from the U.S. National Science Foundation’s Coupled Human and Natural Systems Program.


Rules of thumb for tropical forest restoration

Sometimes farmlands quickly regrow tropical forests on their own, but other times they don’t. Dr. Karen Holl, a professor at the University of California Santa Cruz, gives some rules of thumb for when we can save money on tropical forest restoration by letting nature do the work, and when we may need to invest in tree planting.

Ambitious targets are being set to restore tropical forest because of their importance in storing carbon, regulating water cycles, conserving biodiversity, and supporting the wellbeing of people who live in tropical countries. For example, the 20 × 20 Initiative aims to restore 20 million hectares of tropical forest in Latin America by 2020. This represents an area slightly smaller than the country of Ecuador. One big question is: How are we going to restore forests at this scale with limited funds?

One of the cheapest ways to restore forest is to let nature do the work and leave forests to recover on their own. This works in some sites where forests regenerate quickly. In other cases, usually sites that have been used intensively for agriculture, the land may be covered by tall grasses (up to 3 meters, or 10 feet high) for years. Our past research shows that even within a small region, the rate of natural forest recovery varies greatly.


Natural forest recovery is highly variable in southern Costa Rica, even after a decade of recovery. Left: slow recovery on a former farm, still dominated by non-native grasses, with an open canopy and little tree recruitment. Right: speedy recovery on a former farm, with virtually no grass cover, a closed canopy, and diverse tree recruitment. Photos by Andy Kulikowski.

So, how do we predict which sites will recover quickly and which ones need some help in the form of clearing pasture grasses and planting trees? If we could develop some rules of thumb it would help land managers to more efficiently allocate scarce restoration funds.

To answer this question, we drew on our long-term study on tropical forest restoration in southern Costa Rica. We have research plots at 13 different sites where we removed the land from agriculture and let the forest recover on its own. Each year we measure grass cover, tree canopy cover, and how many and what species of new tree seedling establish in the plots. We have also quantified the forest cover surrounding the plots, the nutrients in the soil, and how long cows had grazed the sites in the past.

We found that two easy-to-measure variables explained on average two-thirds of variation in forest recovery 7 years later; those were the amount of grass cover and tree canopy cover measured after only 1.5 years. Plots that had more canopy cover and lower grass cover early on had a closed tree canopy and lots of forest tree seedlings from many species after nearly a decade. We were surprised that the amount of surrounding forest cover and soil nutrients did not explain much of the variation in forest recovery.


Rules of thumb for predicting tropical forest regeneration on farmlands. Forests grow back quicker when there is not too much grass, a little bit of shade, and many tree seedlings already present. Illustrations by Michelle Pastor.

Of course, our results need to be tested in other recovering tropical forests. But, if they hold true, this is good news! It means that land owners and managers just need to wait a year or two and then measure the tree canopy and grass cover. If some trees have established and are starting to shade out the grasses, land managers can use the low cost method of leaving the site to recover naturally. If the site is mostly a monoculture of dense grass, then the site is a good candidate to plant native trees. Planting trees takes more resources since it is necessary to clear around the native tree seedlings for a couple of years until they grow taller than the grasses. At least now there are some general guidelines to help chose where to invest the extra effort.

For more information, see our new paper in Applied Vegetation Science. This work was supported by the National Science Foundation.

How to grow instant fig trees to restore rain forests in Costa Rica

CCSD scientist Leighton Reid and Lyon Arboretum director Rakan Zahawi write about giant fig tree cuttings: how to make them and why some grow better than others.

Choosing the right species to include in a restoration project is a hard choice, but in the economy of nature, some species earn a bigger ROI than others. For example, Pacific sea otters maintain kelp forests by eating sea urchins, and wolves in Yellowstone National Park allow aspen groves to regenerate by scaring away tree-munching elk. These vital creatures are called “keystone species” because they hold ecosystems together, much like the keystone in an arch.


A keystone and three keystone species. (A) This small keystone holds up an arch in the Shoenberg Temperate House at Missouri Botanical Garden. (B) Sea otters are keystone predators in kelp forests. Photo by Marshal Hedin CC-BY 2.0. (C) Gray Wolves are keystone terrestrial predators. Photo by Gary Kramer USFWS CC-BY-NC 2.0. (D) A keystone fig tree feeding a Knobbed Hornbill in Sulawesi, Indonesia. Photo by T. R. Shankar Raman CC BY-SA 3.0.

Plants can be keystone species too. Around the world there are about 800 species of fig trees, and they hold tropical forests together by providing food for a wide array of animals. On any given day, the busiest tree in a rain forest is likely to be a fig tree with fruits. Monkeys, birds, bats, and others gather at fig trees to eat, and in the process, they deposit seeds of other plant species that they have been carrying in their guts. This chain of events, repeated day after day, often turns the area beneath a fig tree into a hotspot of plant diversity.

A few years ago, we had an idea to plant keystone fig trees in young forests in Costa Rica. We wanted the figs to grow as fast as they could, so instead of planting seedlings, we planted cuttings – big ones. With help from our local collaborator, Juan Abel Rosales, we cut dozens of twelve foot-long branches from eight species of fig trees. We stripped off all of their leaves to keep them from drying out, and then we planted our figs trees in shallow holes.


Rakan Zahawi (delighted!) poses with a three year-old fig stake.

To our delight, many of the fig trees grew!

The ones that did the best came from a special group, the subgenus Urostigma. Many figs in this group have a unique life strategy. They begin their lives in the top of a tree when their tiny seeds are deposited on a branch by a bird or some other animal. As they grow in the treetop, they send long roots down to the ground, and these roots harden and fuse together, forming a lattice-like trunk. Over time, these figs kill their host trees by taking most of the water, nutrients, and light. They also keep the host tree from growing outwards, giving them the nickname “strangler figs”. Maybe the ability to transform a flimsy, dangling root into a solid trunk is related to these figs being able to grow from cuttings.

To find out how well our planted fig cuttings might survive over the long-term, we also tracked down some fig cuttings that we had planted in 2004. We were happy to learn that out of the trees that survived for their first three years of life, all of them were still thriving a decade later.

Full disclosure: planting large cuttings is not a new idea.  Farmers in many parts of the tropics plant trees this way to create ‘living fences’ – with all of the normal fixings like gates and barbed wire, but with a row of living trees instead of dead posts. The advantages for farmers are many – their fences don’t rot and fall apart (that happens quickly in the tropics); the trees provide shade for cattle; they have a constant source of new fence posts (by cutting off a limb); and in some cases they can feed the young shoots to livestock.

Big cuttings have big benefits for restoration too. Not only are planted trees already several feet tall, you also get to skip the pricey nursery phase, and, most excitingly, cuttings have a tendency to fruit quickly.

Some of our young fig trees are now making fruit, but we will have to wait a bit longer to see whether they start attracting more big animals and whether those animals carry more tree seeds into our young forests. For now, we can say that others who are interested in growing keystone figs for forest restoration may have the best luck by working with the stranglers.

For more information, please take a look at our open access paper on this project in Perspectives in Ecology and Conservation and prior blog posts here, here, and here.


How to grow an instant fig tree. (A) Remove a long, thin branch segment from an adult tree. The red arrow shows a cut branch. (B) Strip the cuttings of their leaves to keep them from drying out, then carefully transport cuttings so as not to damage cortical tissue. Here, cuttings are padded by a foam mattress. (C) Remove the bark from a ring on the cutting to promote root growth. Here, a ring is being cut about 20 cm (8 in) above the base so that it will be just below the soil surface when planted. (D) Dig a shallow hole and plant the cutting. Be sure that the cutting is firmly planted to prevent it from toppling, but take care not to compact the soil too much around its roots. Photos by Rakan Zahawi.


To plant or not to plant?*

What we think we know about how to restore tropical forests is getting a second look. A new paper produced by scientists in Missouri Botanical Garden’s Center for Conservation and Sustainable Development (CCSD), the University of Hawaii’s Lyon Arboretum, and the University of Maryland Baltimore County points out an important bias in recent studies.

How should we restore forests in places where they have been lost? This is one of the main questions that we study in the CCSD, so we were surprised last year when a big synthesis paper that compiled data from many earlier studies said that, when it comes to restoration, doing nothing was the same as doing something.

That’s only a slight exaggeration. The paper, by Renato Crouzeilles and several other scientists, said that letting a forest regrow on its own (that is, natural regeneration) was usually more successful than planting trees (that is, active restoration). Their conclusion was based on comparing many studies done throughout the world’s tropical forest regions.

Apples to oranges

The problem with this paper (and several like it) was that the set of studies looking at natural regeneration were not really the same as the set of studies looking at tree planting. The natural regeneration studies focused on forests that already existed, while the tree planting studies focused on a wider range of sites, many of which started with no forest. In other words, the natural regeneration studies had already been filtered to exclude places with a weak ability to grow forests.


Comparing tree planting studies to pre-existing forests is like comparing apples and oranges. Photo by Michael Johnson CC BY 2.0.

To understand the problem, it is helpful to look back at the history of tropical forest restoration research. For many years, scientists who wanted to know about how forests recover after a disturbance (like a hurricane or logging) would go out and find several forests that had been recovering for different amounts of time. If you take forests that are 5, 10, and 20 years old, you can try to compare them to each other in order to see how a forest might change over 20 years. In contrast, tree planting studies usually start with a piece of land that has no trees on it. Scientists who want to know how trees grow on this land will plant some and then observe their survival and growth over time.  These trees may or may not create a forest there, as the land can vary in quality.

So where does that bring us with respect to this study? If you compare a forest that already exists with another potential forest where planted trees may or may not survive and grow well, it’s a safe bet that the pre-existing forest will have taller trees. It has a head start over the planted forests, and we argue in our paper that the comparison is not a fair one.

This means that letting forest regrow on its own is not always a better option than planting trees. In fact, there are many places – like overgrazed pastures, mine sites, and other heavily degraded lands – where forests have been cleared and most likely will not be able to grow back on their own.


Comparison of natural regeneration (foreground) and active tree planting (background) to restore a cattle pasture in southern Costa Rica. Tree seedlings planted on the hillside are just visible in the 2005 image. The yellow circle indicates a person for scale. After nine years, active tree planting had produced a forest, whereas natural regeneration was stalled. Overgrown pasture grasses covered the ground. Natural regeneration is highly variable, so this example is not representative of all situations. Photos courtesy of Karen Holl.

 Same team!

While we were not convinced by studies that said that natural regeneration is better than tree planting, we also don’t want to take any options off the table. Natural regeneration and tree planting are not mutually exclusive – in fact, they are highly complementary. Our practical advice is that if you want to get forest back, the best option is to see if natural regeneration can do the trick before you invest in tree planting. Or better yet, set up a paired experiment comparing the two strategies at the same site.

*Thanks to Erle Ellis for coming up with the title for this blog post. For more information, please see our open-access paper and press releases at EurekAlert, UMBC News, and Science Daily.


Special Feature: Ecological Restoration in a Changing Biosphere

The following is an introduction by Leighton Reid and James Aronson to a special feature in Annals of Missouri Botanical Garden about ecological restoration in a changing biosphere. The eight papers described are derived from presentations last October at the 65th Annual Plant Symposium. The full issue can be found here.

Restoration efforts will affect large areas of the planet and hundreds of millions of people over the coming decades, but what will these actions look like, and what will they achieve? Debate continues about what constitutes appropriate restoration targets in our human-dominated and ever more rapidly changing world, and the outcome of this debate will impact the actions taken to conserve biodiversity, sequester carbon, and improve human livelihoods at large spatial scales. This special issue brings together eight scientific, historical, and journalistic perspectives to address these two critical questions about ecological restoration in a rapidly changing biosphere.

In the post-COP22 world, when all three of the UN’s “Rio Conventions” call for scaling up and mainstreaming of ecological restoration (UNCBD 2012; UNCCD 2015; UNFCCC 2015), and dozens of governments have made ambitious restoration commitments (IUCN 2016), it is clear that restoration programs will affect hundreds of millions of hectares – and as many people – over the coming decades. At the same time, we find ourselves in an era of unprecedented change where climate, ecological baselines, and future land-use changes are highly uncertain (Steffen et al., 2015). This raises the crucial question: What will large-scale restoration activities look like in the coming years?

Unsurprisingly, there are differences of opinion about the future of restoration and how to scale it up and integrate it with larger programs in an era of major, anthropogenic changes. Hobbs et al. (2011; pg 442) observe that “…the basic principles and tenets of restoration ecology and conservation biology are being debated and reshaped. Escalating global change is resulting in widespread no-analogue environments and novel ecosystems that render traditional goals unachievable. Policymakers and the general public, however, have embraced restoration without an understanding of its limitations, which has led to perverse policy outcomes.” [Emphasis added]

This perspective has received considerable attention (ESA 2016) and also pointed criticism (Murcia et al., 2014). Aronson et al. (2014; pg 647) retort that “…Restoration includes a wide range of practical possibilities for dealing with transformed ecosystems, including rehabilitation, reclamation, and remediation. Some will bring the ecosystem back to its historical trajectory, some will bring back only some attributes, but the intention is that the end product is better than the degraded ecosystem. Importantly, a label such as novel ecosystem implies no need for further intellectual exertion – and ignores the growing science of the young discipline of ecological restoration.” [Emphasis added]

The debate goes on about what we are trying to restore (Hobbs, 2016; Kattan et al., 2016; Miller & Bestelmeyer, 2016), with implications far beyond academia. Billions of dollars are now being spent to rehabilitate and restore degraded ecosystems, sometimes at large scales, and the science of restoration ecology must adapt to be integrated in larger planning and management schemes, wherein conservation, management, and restoration will all take place.

On 8 Oct 2016, we convened a panel of six scientists, one historian, and a journalist, all with long-standing involvement in the field of restoration ecology. The goal was to discuss ecological restoration in a changing biosphere at the 63rd Annual Fall Symposium at Missouri Botanical Garden. Each speaker has contributed a paper to this special issue.

The first set of papers focus on the question: Has global change outpaced and rendered obsolete the so-called “classical” ecological restoration approach? Aronson et al. (2017) say no, far from it; for example, the historically-based reference system ‒ a pillar of ecological restoration to date ‒ is more valid than ever and can indeed be adapted to landscape and higher levels of complexity. They emphasize that while restoration ecology has produced many useful ecological models, a participatory approach and consensus-building among stakeholders are crucial at these higher levels of integration. Falk (2017), in contrast, says yes: global change calls for radical rethinking of ecological restoration. He focuses on ponderosa pine forests in the southwestern US, which are undergoing a major, climate change-induced biome shift from forest to shrub land, and he concludes that a shift towards resilience-based management is necessary to supplement traditional ecological restoration. Meine (2017) takes the middle ground through an historical analysis; he notes that Aldo Leopold (1887-1948) would likely have concluded that a simple “yes” or “no” was inappropriate and that ecological novelty is neither novel nor absolute.

Whereas the first group of papers asks what we should restore, the second group focuses more on how we will restore at larger spatial and temporal scales. Brancalion & van Melis (2017) suggest that to bridge the gap between science and practice, we need to innovate; rather than refining current approaches, restoration ecologists must look outside of their disciplinary silos for fresh solutions to contemporary dilemmas. One source of new insights will be through joint research between scientists and practitioners. To this end, Holl (2017) presents several new directions for tropical forest restoration research (graduate students – take note!). She emphasizes that for research to best inform practice, it should be conducted at large spatial and temporal scales, research projects should be undertaken jointly with stakeholders, and resulting knowledge should be shared across regions. Chazdon (2017) argues that natural regeneration, more than any other method, is the key for scaling up to efficient forest and landscape restoration, and she emphasizes the need to identify priority areas where natural regeneration is maximally feasible and minimally competitive with alternative land uses. Finally, Reid et al. (2017) argue that however we restore ecosystems, we should plan to make them last; the longevity of restored ecosystems, they suggest, is variable, often finite, and determined to some degree by stakeholder preferences, environmental attributes, and the umbrella of governance. These papers emphasize tropical forest restoration, particularly in Latin America, which is appropriate given this biome’s global importance, yet the topics addressed will be of interest to readers with experience in many ecosystems.

The last word (for this special issue, at least) is left to Paddy Woodworth (2017), an international journalist with broad and optimistic perspectives on ecological restoration (Woodworth, 2013). Looking across the contributions, he observes that the words we choose have meaning and cautions against the use of the word “restoration” for anything less than the process of assisting the recovery of an ecosystem that has been degraded, damaged, or destroyed (SER 2004).

We hope that readers from many backgrounds, including researchers, practitioners, and policymakers, will find this special issue worth pondering as they move forward with our collective task to progress towards a more sustainable, just, and desirable future.



Aronson, J., J. Blignaut & T. B. Aronson. 2017. Conceptual frameworks and references for landscape-scale restoration: Reflecting back and looking forward. Annals of the Missouri Botanical Garden 102(2): 188–200.

Aronson, J., C. Murcia, G. H. Kattan, D. Moreno-Mateos, K. Dixon & D. Simberloff. 2014. The road to confusion is paved with novel ecosystem labels: a reply to Hobbs et al. Trends in Ecology & Evolution 29: 646-647.

Brancalion, P. H. S. & J. van Melis. 2017. On the need for innovation in ecological restoration. Annals of the Missouri Botanical Garden. 102(2): 227–236.

Chazdon, R. L. 2017. Landscape restoration, natural regeneration, and the forests of the future. Annals of the Missouri Botanical Garden. 102(2): 251–257.

ESA (Ecological Society of America). 2016. Ecological Society of America announces 2016 award recipients. The Bulletin of the Ecological Society of America 97: 337-351.

Falk, D. A. 2017. Restoration ecology and the axes of change. Annals of the Missouri Botanical Garden. 102(2): 201–216.

Hobbs, R. J. 2016. Degraded or just different? Perceptions and value judgements in restoration decisions. Restoration Ecology. doi: 10.1111/rec.12336.

Hobbs, R. J., L. M. Hallett, P. R. Ehrlich & H. A. Mooney. 2011. Intervention ecology: Applying ecological science in the Twenty-first Century. Bioscience 61: 442-450.

Holl, K. D. 2017. Research directions in tropical forest restoration. Annals of the Missouri Botanical Garden. 102(2): 237–250.

IUCN (International Union for Nature Conservation). 2016. Bonn Challenge commitments. Date accessed: September 22, 2016

Kattan, G. H., J. Aronson & C. Murcia. 2016. Does the novel ecosystem concept provide a framework for practical applications and a path forward? A reply to Miller and Bestelmeyer. Restoration Ecology 24:714-716.

Meine, C. 2017. Restoration and “novel ecosystems”: Priority or paradox? Annals of the Missouri Botanical Garden. 102(2): 217–226.

Miller, J. R. & B. T. Bestelmeyer. 2016. What’s wrong with novel ecosystems, really? Restoration Ecology 24: 577-582.

Murcia, C., J. Aronson, G. H. Kattan, D. Moreno-Mateos, K. Dixon & D. Simberloff. 2014. A critique of the ‘novel ecosystem’ concept. Trends in Ecology & Evolution 29: 548-553.

Reid, J. L., S. J. Wilson, G. S. Bloomfield, M. E. Cattau, M. E. Fagan, K. D. Holl & R. A. Zahawi. 2017. How long do restored ecosystems persist? Annals of the Missouri Botanical Garden. 102(2): 258–265.

SER (Society for Ecological Restoration). 2004. The SER primer on ecological restoration. Date accessed: September 28, 2009

Steffen, W., W. Broadgate, L. Deutsch, O. Gaffney & C. Ludwig. 2015. The trajectory of the Anthropocene: the great acceleration. The Anthropocene Review 2: 81-98.

UNCBD (United Nations Convention on Biological Diversity). 2012. UNEP/CBD/COP/DEC/XI/16. Date accessed: 12 December 2016

UNCCD (United Nations Convention to Combat Desertification). 2015. Land matters for climate: reducing the gap and approaching the target. Date accessed: 12 December 2016

UNFCCC (United Nations Framework Convention on Climate Change). 2015. Paris agreement.  Date accessed: 12 December 2016

Woodworth, P. 2013. Our Once and Future Planet. University of Chicago Press, Chicago.

Woodworth, P. 2017. Meeting  the twin challenges of global change and scaling up, Restoration needs insights from the humanities as well as analysis from science. Annals of the Missouri Botanical Garden. 102(2): 266–281.

Tree islands for tropical forest restoration: the outlook is rosy after 10 years

Planting tree islands has many of the benefits of larger plantations, but entails significantly less cost. Karen Holl (University of California, Santa Cruz), Leighton Reid (Missouri Botanical Garden), and Zak Zahawi (American University of Beirut) describe recent findings on tree seedling recruitment in a long-term experiment in southern Costa Rica.

Over the past few years there have been a growing number of commitments at the global, national and regional scale to restore forests because of their importance to conserve biodiversity, sequester carbon, reduce erosion, and provide goods and services to people. For example, Initiative 20×20, led by the International Union for the Conservation of Nature, aims to restore 20 million hectares of tropical forest by 2020, an area roughly equivalent to the size of Uruguay or Nebraska.

A common strategy to restore forests is to plant trees. But, the big question is: where will the money come from to plant billions of trees when there are so many pressing needs? As restoration ecologists, we started thinking about how we could most efficiently allocate resources to get the best bang for the buck and restore the largest area of forest.


Trade-offs in forest restoration strategies. Planting fewer trees leaves more to chance and can require more time, but tree plantations are more expensive and leave a bigger ecological footprint. Our study tests an intermediate option, and after 10 years it appears to provide a good balance. Figure modified from Corbin & Holl (2012).

Starting over 10 years ago, we set up a large-scale tropical forest restoration experiment in southern Costa Rica to test two ideas.

First, we tried planting tree “islands”. The idea is to plant groups of trees that attract birds and bats, which disperse most tropical forest tree seeds. The tree canopy also shades out light-demanding grasses that can outcompete tree seedlings. In one experimental treatment, we planted tree islands that covered about 20% of 50 × 50 m plot of former cattle pasture. We compared that to plots where no trees were planted (natural recovery) and to the more intensive (and more typical) restoration strategy of planting trees in rows throughout the plot (plantation).

Second, we asked: is it only possible to restore forest near remnant forests or can you restore forest anywhere in the landscape? This is important information to help guide forest restoration efforts. To do this we set up our entire experiment at 13 sites, some of which were mostly surrounded by agricultural land and some of which were adjacent to the largest remaining forests in the region.

Then we monitored establishment of new tree seedlings in our research plots over a decade. We compared the number of seedlings, number of species, and types of species in the restoration plots with those found in the nearby forest to evaluate how well the forest is recovering.

The tree island planting method not only saves money on buying, planting, and maintaining seedlings, but it also results in a more heterogeneous distribution of trees, so it looks more like a natural forest.


Profuse tree seedling and sapling recruitment in the understory between two tree islands in southern Costa Rica.

We counted over 6000 tree seedlings, 88% of which have seeds that are dispersed by animals. On average there were many more tree seedlings in the tree island and plantation treatments than in the natural recovery plots. These results suggests that some tree planting helps the forest to recover faster, but that it is not necessary to plant the whole area with trees. The tree island planting method not only saves money on buying, planting, and maintaining seedlings, but it also results in a more heterogeneous distribution of trees, so it looks more like a natural forest.

Even though there were many tree seedlings in the island and plantation plots, on average there were less seedlings of tree species that have big seeds (>0.5 cm/0.2 inches across) compared to mature, reference forests. It seems that the larger-seeded species that are common in mature forests are much slower to colonize restored sites, likely because they are eaten and dispersed by a small number of larger animals, such trogons and agoutis. Many of those dispersers are less likely to visit early successional forest.


Small frugivores, small seeds. Most of the birds we see in these experimental plots are small-gaped omnivores (e.g., Yellow-bellied Elaenia, Elaenia flavogaster, left), but it usually takes large-gaped species to disperse larger seeds1. The figure at right shows the maximum fruit size that a bird species with a given gape size was able to consume in a cloud forest in central Costa Rica (modified from Wheelwright (1985)). In our experiment, small seeds were ubiquitous, but large seeds were mostly absent.

We were surprised that the amount of forest cover around the experimental plots had a weak effect on the number of seedlings establishing. In other words, isolated plots had just as many tree seedlings as plots right next to old-growth forests. We think that this is likely due to the fact that there are many trees in the agricultural landscape surrounding our plots; these trees include remnant trees, living fence rows, and riparian corridors. Trees in the landscape can serve an important role in both providing sources of seeds and stepping stones for the movement of seed-dispersing fauna. We anticipate that having forest nearby will be more important in future years as these forests build up greater diversity of rare, large-seeded species. Nonetheless, our results suggest that there are good prospects for restoring forests in many locations in this landscape.

Our key finding is that planting tree islands can be a cost-effective way to restore tropical forests at our study site in Costa Rica, but we hasten to note that the strategy should be tested in other locations, particularly areas with fewer forest elements in the surrounding countryside. Our study also demonstrates that tropical forests can recover some species quickly but it will take many decades, if ever, for forests to fully recover. So, preserving existing rain forests is critical to conserve biodiversity and the services they provide to people.


Diverse tree cover in an agricultural landscape in southern Costa Rica. Remnant trees in pastures, trees along fence rows, and riparian forests provide important sources of flora and fauna to speed up forest recovery.

1See Melo et al. (2009) for an example to the contrary: small-gaped animals dispersing fairly large fruits and seeds.

This work was supported by a grant from the National Science Foundation.

Ecological Restoration in a Changing Biosphere

If you were at the MBG Fall Symposium, we want to hear from you! How did the symposium change your perception of restoration? Send us an email at

On October 8th, Missouri Botanical Garden hosted its 63rd annual Fall Symposium. This year’s theme was Ecological Restoration in a Changing Biosphere. Author and journalist Paddy Woodworth moderated the day, and seven speakers presented contemporary perspectives on a core challenge in modern restoration ecology. Namely: in the post-COP21 world, when all three UN conventions call for scaling up and mainstreaming of restoration, it is clear that restoration will affect hundreds of millions of hectares – and as many people – over the coming decade. At the same time, we find ourselves in an era of unprecedented change where climate, ecological baselines, and future land-use changes are highly uncertain. This raises the question: What should large-scale restoration look like in the remainder of the 21st century?


2016 Fall Symposium speakers. From left to right: Peter Wyse Jackson, Curt Meine, Robin Chazdon, James Aronson, Leighton Reid, Pedro Brancalion, Karen Holl, Don Falk, Paddy Woodworth, and Jim Miller. Photo by Andrea Androuais.

Talks during the morning focused on tropical forests, where much of the international restoration dialogue is focused.

  • Leighton Reid (Missouri Botanical Garden) opened with a presentation on restoration longevity – the idea that some restoration projects create ecosystems that persist for more than a century (e.g., Floresta da Tijuca), while other projects fail quickly. Dr. Reid argued that how long restored ecosystems persist is quantifiable, predictable, and manipulable, opening the possibility for more ambitious restoration planning.
  • Robin Chazdon (University of Connecticut and beyond) then spoke about forest landscape restoration, an approach that aims to regain ecological integrity and enhance human well-being in deforested, human-impacted, or degraded forest landscapes. Drawing on a wealth of large-scale studies, Dr. Chazdon made the case that natural forest regeneration is the most ecologically effective and economically feasible approach to forest restoration globally.
  • Karen Holl (University of California Santa Cruz) presented her take on research priorities for forest restoration in the Neotropics. She highlighted that researchers could make an impact by studying forest restoration at larger spatial scales, at longer temporal scales, and in collaboration with stakeholders. Improving information exchange and standardizing monitoring protocols were also among her top priorities. (Graduate students, take note!)
  • Dr. Pedro Brancalion (University of São Paulo) completed the morning session with a TED talk-style discussion of the linkages between science, technology, policy, and best practice in Brazilian Atlantic Forest restoration. Using Thomas Kuhn’s structure of scientific revolutions, Dr. Brancalion argued that restoration ecology is in a crisis period, in part because disciplinary research has predominantly created solutions at smaller spatial scales than the (growing) problems the discipline seeks to address. Perhaps restoration is ripe for a paradigm shift?

Dr. Pedro Brancalion (right) asks whether restoration ecology is ready for a new paradigm shift, as Paddy Woodworth (left) moderates. Photo by Robin Chazdon.

After lunch, the conversation turned towards a major academic debate in restoration ecology. Has global change outpaced the restoration approach? And is a new approach needed?

  • Curt Meine (The Aldo Leopold Foundation) drew on his long experience in the upper Midwest, and, in particular, his studies of author and environmentalist Aldo Leopold (1887-1948). He argued that Leopold avoided the simple polarities through which some contemporary restoration debates are framed. He viewed nature in a relative way, neither entirely wild, nor entirely domesticated in any given landscape. Although he practiced ecological restoration in some contexts, he also advocated soil conservation and sustainable agriculture – activities motivated by his core values, as expressed in The Land Ethic (1949).
  • James Aronson (Missouri Botanical Garden) followed with an elucidation of the reference ecosystem concept. Reference ecosystems, he noted, help determine the social and ecological vision for a restoration project or program – a critical issue for restoring historic continuity in degraded landscapes. Dr. Aronson described a family of restorative actions for achieving progress towards the reference system, drawing on examples from Jordan and South Africa. He argued we need to look deeper into the past and ponder our choices from many angles as we decide how to do more effective restoration at the landscape and larger scales.
  • Donald Falk (University of Arizona) delivered the keynote address. He painted a disturbing portrait: rapid climate change is driving a massive forest-to-non-forest transition in the southwestern United States. In particular, many ponderosa pine forests will not be able to persist in the future where they have been in the recent past and present. Perhaps restoration ecologists should transition too. Rather than “chasing the ambulance”, maybe we could get out ahead of disasters and ease transitions between stable ecosystem states. Anticipating ecosystem transitions could mitigate the loss of ecosystem functioning that accompanies major climate-driven forest fires, but it would require a shift in restoration thinking. Importantly, Dr. Falk noted that ecosystems do not care what words we use – ecosystems respond to actions.

With moderator Paddy Woodworth’s help, we finished the day with a panel discussion, inviting questions from the audience. Among the thoughts and questions that we were left with:

  • Is ecological restoration more difficult in places with greater population density?
  • Should restoration focus on policy, economic, or cultural motivations for engaging people?
  • Are values a better guide for land management than ecological history? Are the two complementary?
  • How can the reference ecosystem concept accommodate rapid biome changes, as we are seeing in the Southwestern USA?
  • What is the way forward to mainstream serious, multisectorial monitoring and evaluation with all these new factors to consider? Who will fund it?
  • To what extent can we move from restoring degraded ecosystems to avoiding degradation in the first place?
  • Can forest landscape restoration and natural forest regeneration bridge the gap between small-scale, past restoration experience and present, large-scale restoration needs?

PhD candidates Ricardo Cesar (University of São Paulo) and Leland Werdan (University of Minnesota) compare notes on seedling functional traits in dry tropical forest restoration. Leland was the recipient of the annual Delzie Demaree award. Photo by Robin Chazdon.


More than 150 people registered for the symposium. They came from three continents, five countries, and seven US states.