Planting trees recovers 70 years’ worth of dead wood carbon pools in less than two decades

By Estefania P. Fernandez Barrancos, a PhD candidate in Biology at the University of Missouri – St. Louis and a fellow of the Whitney R. Harris World Ecology Center. Her most recent research paper in Forest Ecology and Management is freely available through March 9th.

When most people walk through a forest the last thing they probably look at is dead vegetation, and unless you are an avid mushroom harvester you probably don’t even notice dead logs. However, dead wood stores an important amount of carbon. An amount important enough that if dead wood disappeared it could promote more changes to our already rapidly changing climate.

Mushrooms on a dead log. Photo: JL Reid.

Dead wood is also a crucial habitat for many organisms such as fungi, insects, and birds. Many insects and fungi use dead wood as a source of food and nutrients, and several species of birds are only able to nest in dead logs.

A Resplendent Quetzal (Pharomachrus mocinno) exiting its nest inside a standing dead log to go harvest food for its fledglings. Photo: Estefania Fernandez.

Anthropogenic disturbances, such as logging and deforestation, can significantly decrease the amounts of dead wood present on the forest floor, sometimes leading to losses of up to 98% of dead wood. The implications of dead wood loss are potentially warmer temperatures due to the release of carbon contained in dead wood as well as the loss of habitat that is critical to many forest organisms. Tropical ecosystems contain some of the most biodiverse habitats on Earth, yet they are among the ecosystems that suffer the most from anthropogenic disturbance. For example, most forests in the county of Coto Brus in Southern Costa Rica, our study area, were transformed into cattle pasture or coffee plantations in the 1950s-1980s. Today, the landscape consists of a mosaic of cattle pasture, coffee plantations, and small forest remnants.

Deforestation to create farms and cattle pastures has decreased the amount of dead wood in southern Costa Rica. Photo credit: JL Reid.

Forest restoration is the process of assisting the recovery of an ecosystem that has been damaged or destroyed (SER International Standards) and it has a high potential to reverse the problem of dead wood loss through different strategies. In the Tropics, the most common restoration strategies are passive and active restoration. Passive restoration consists of allowing an ecosystem to recover with minimal to no human input.  In contrast, active restoration consists of assisting the ecosystem in its recovery through actions such as tree planting.

Old-growth forest (A) and and two restoration treatments: tree plantations (B) and natural regeneration (C). Old-growth forests are ≥100 years old. Plantations and natural regeneration were 16-17 years old at the time of the study. Photos:  Juan Abel Rosales & Estefania Fernandez.

Recently, I studied the pattern of dead wood re-accumulation through time after disturbance in southern Costa Rica as well as the effectiveness of passive and active restoration at recovering dead wood as it is found in undisturbed forests. To evaluate dead wood accumulation through time, my team and I surveyed dead wood volumes inside 35 forest patches of increasing ages (from 3 to over 100 years old) that were former coffee plantations. We evaluated the effectiveness of active vs. passive restoration at recovering dead wood by surveying dead wood volumes inside 17-year old passive and active restoration plots and inside nearby old-growth forests. Our passive restoration treatment was represented by natural regeneration plots around which fences were established to exclude cattle and where vegetation was allowed to re-establish naturally. Our active restoration treatment was represented by restoration plantations, where seedlings of two native (Terminalia amazonia and Vochysia guatemalensis) and two naturalized (Inga edulis and Erythrina poeppegiana) tree species were planted 17 years ago to facilitate the re-establishment of vegetation. Our reference ecosystem included nearby old-growth forests over 100 years old.

Juan Abel Rosales measures the diameter of dead logs in order to estimate their volume in an old-growth forest in Southern Costa Rica. Photo: Estefania Fernandez.
To measure the diameter of dead, rotting logs, we measured the distance between two tent poles set vertically along the logs’ edges. Photo: Estefania Fernandez.
Jeisson Figueroa Sandí establishes a transect to evaluate dead wood inside a forest fragment. Photo: Estefania Fernandez.

We found that dead wood recovers following a logistic shape through time in our study area: volumes are low initially, increase rapidly, and then plateau. The low volumes of dead wood at the beginning of succession could be explained by the fact that most of the wood remains are typically harvested by local inhabitants after lands are abandoned in our study area. As pioneer trees recolonize abandoned coffee plantations and subsequently die, they produce dead wood. As the forest grows older, there is a mix of short-lived pioneer trees and long-lived trees which contribute to large amounts of dead wood on the forest floor through branchfall and their own deaths.

Dead wood volumes as function of forest age in a chronosequence of secondary forests in southern Costa Rica. Blue dots represent the raw data (i.e. course woody debris, or CWD, volumes per hectare). The red line represents the predicted values from a generalized linear model plotted using a smoothing function. Eight outliers that were included for the analysis where CWD volume per transect was ≥125 m3ha-1 were removed for better visualization. CWD volumes in plantations (purple dot), natural regeneration (yellow triangle) and five nearby old-growth forests (green dot) are also represented. Mean CWD volumes per hectare for each restoration plot (n=5) and corresponding 95% confidence intervals are shown.

We also found that restoration plantations contain 41% of dead wood amounts found in old-growth forests, whereas natural regeneration only contained 1.7% of dead wood volumes found in old-growth forests. The extremely low recovery of dead wood in natural regeneration might be explained by the fact that our natural regeneration plots were dominated by exotic grasses which typically hamper tree colonization. If there are no trees growing in the plots, there cannot be dead wood either. This is an important finding, because it shows that restoration plantations area a faster and more efficient way to recover dead wood in this fragmented, pasture-dominated landscape, even though this restoration strategy might be more time consuming and expensive due to the costs and time of planting seedlings.

Overall, our study unveils an important forest process, showing that dead wood carbon pools recover following a dynamic logistic pattern through time in this Neotropical forest region. Knowing that dead wood is 50% carbon, our findings allow us to predict carbon stocks in Neotropical forests more accurately. Our study also shows that restoration plantations accelerate the recovery of dead wood carbon pools in this Neotropical ecosystem, and potentially promote the preservation of dead wood-associated biodiversity.

For more information, see our recent paper in Forest Ecology and Management, which is freely available online through March 8th, 2022.

The importance of knowledge of cultural values for dryland ecological restoration: Lessons from Argentine Patagonia

Fernando Farinaccio, Eliane Ceccon and Daniel Pérez, describe the importance of documenting cultural values, in the use of native flora, as a contribution to the restoration of drylands. Fernando is a researcher at the Laboratory for the Rehabilitation and Restoration of Arid and Semi-arid Ecosystems (LARREA), Argentina. Eliane is a researcher at the Regional Center for Multidisciplinary Research at UNAM (National Autonomous University of Mexico), Mexico, and Daniel is the scientific director of LARREA.

NB. LARREA belongs to the Faculty of Environmental and Health Sciences of the National University of Comahue, Argentina, where sixteen researchers and collaborators study selection of species for the recovery of sites with severe disturbance, seed-based restoration, interactions between exotic and native species, agroecological systems, and restoration-based education.

The extreme socio-ecological transformation and degradation of vast areas of arid Argentinean Patagonia has its origin in the 1880s when the Argentine government carried out an official program of extermination of all Indigenous Peoples (ignominiously called the “Desert campaign”). The goal was to consolidate political dominance over the coveted territories and to expand livestock production.

As elsewhere, this genocide led to a tragic loss of human lives and the uprooting and dispossession of native inhabitants who had lived in and managed this country for millennia prior to the arrival of the Europeans – most of whom had little or no understanding of the natural dynamics of this arid and semiarid territory or in the lives and cultures of the peoples who lived there.

Aguada San Roque, an isolated rural settlement of 160 inhabitants, extends over an area of 142,000 hectares in an arid basin called “Añelo basin” in northern Patagonia. It is characterized by high altitude variability, from 223 to 2258 meters above sea level, over a linear distance of 50 km. This town is in one of the most arid ecosystems of Argentina called ‘Monte’ (Busso and Fernández 2017). This ecosystem covers 20% (approximately 50 million hectares) of Argentina. The Monte has an annual average temperature of 12°C, with a high thermal amplitude and an annual temperature range from 40°C to −13°C (Coronato et al. 2017). The relationship between precipitation and potential evapotranspiration ranges from 0.05 to 0.5, indicating a strong water deficit.

“Jarillas” (Larrea spp.; Zygophyllaceae; creosote bush, in English) are the shrub species that give the typical appearance of most natural environments of Aguada San Roque. The dominant species of jarilla (L. divaricata, L. cuneifolia, and L. nitida) can reach approximately 2 meters in height when mature. For the attentive eye, it is probable that hybridizations between them have occurred and generated, among others, the striking “dwarf jarilla”(Larrea ameghinoi), that only reaches 20 to 30 cm in height.

Photo 1. Larrea divaricata. Typical of the natural environment around Aguada San Roque, Patagonia Argentina. Credit: Daniel Pérez.
Photo 2. “Dwarf Jarilla“ (Larrea ameghinoi).A species present in some areas of the Añelo basin. Its biology and reproduction  are very poorly known, but like all Creosote bush species in southern South America and the arid regions of North America, they have enormous influence on ecosystem functioning. Credit: Daniel Pérez.

Despite the aridity of the Añelo Basin, where it rains only 150 mm (6 inches) a year on average, with some years of only 50 mm, the beauty of nature is starkly visible to those who pay attention to details, and its mystery is slowly being revealed through scientific studies of the surprising and wonderful  strategies of plant and animal adaptations to aridity and drought. For example, Grindelia chiloensis (Asteraceae) known as “yellow love” or “honey-eyed” surprises and intrigues with its sticky stems, leaves, and flowers, all bearing so much resin that it is perceptible to the slightest touch. This trait is the result of biochemical efforts to manufacture organic compounds to avoid water loss. Fully 1/3 of the dry weight biomass of individual Grindelia shrubs is made up of these dense resins that allow it to adapt and thrive under the most arid and – importantly – degraded environments.

Photo 3. Detail of the flower of Grindelia chiloensis. Credit: Paul Alvarez.

A species that has probably been benefiting from the advance of wind deposits that multiply due to overgrazing is the “Patagonian lily” (Habranthus jamesonii; Amaryllidaceae). This plant is only noticeably visible in spring, as it develops from bulbs that remain under the sand during periods of unfavorable weather.

Photo 4. Habranthus jamesonii plant and flower in a sandy environment near Aguada San Roque. Credit: Daniel Pérez.

A plant that is almost white in color due to saline exudates is Atriplex lampa; Chenopodiaceae; a member of the widespread arid lands Saltbush genus that rewards the watchful eyes of the desert dwellers (Photo 5). This species has a profuse annual production of fruits with two small bracts that act as ‘wings’(Photo 6).

Photo 5: Atriplex lampa, typical of Monte desert landscapes, with fruits (almost yellow) in spring. Credit: Daniel Pérez.
Photo 6: Fruits of Atriplex lampa. Two bracts act as wings, facilitating their flight and dispersal by the wind. Credit: Paul Alvarez.

In very saline and clayey soils of our region, Halophytum ameghinoi (Halophytaceae) is very common. This species accumulates water in its stems and leaves as a strategy to withstand droughts. Their colors vary from intensely red to green tones during the juvenile and adult growth phases (Photo 7).

Photo 7: Juvenile individual of Halophytum ameghinoi. The increase in salty soils due to degradation will probably increase the amount of natural habitat for this species. Credit: Daniel Pérez.

Sadly, Aguada San Roque, like all the neighboring settlements, is seriously affected by long-standing desertification and degradation processes. Recently, the exploitation of large deposits of shale gas and oil, using fracking technology in the geological formation called “Vaca Muerta”, has revitalized economic activity, but also has induced a new and severe wave of environmental damage both underground and on the surface.

Photo 8: The preparation of land for the extraction of hydrocarbons entails a tremendously brutal action that spells disaster for biodiversity, ecosystem ‘health’ and, ultimately, human health and wellbeing. Credit: Daniel Pérez.
Photo 9: The action of the goats is not perceived with the same sensation of negative impact as that of the heavy machines engaged in fracking. However, overstocking of domestic livestock also causes irreversible damage. Credit: Daniel Pérez.
Photo 10: Frequent dust storms are one of the consequences of overstocking livestock. Aguada San Roque. Credit: Fernando Farinaccio.
Photo 11: A barchan dune, an example of the intense erosive processes in the vicinity of the Aguada San Roque settlement. These natural processes are exacerbated by overgrazing and intense hydrocarbon extraction activity. Credit: Eliane Ceccon.

Therefore, in this region, it is essential to plan and carry out ecological restoration and rehabilitation projects and programs that take into account the harsh socioeconomic conditions of the local population and include them in the process from the beginning. Fully 24% of the inhabitants – all of whom are of “criollo” origin – live in stark poverty, and more than 30% are illiterate. Life for these people is truly precarious, with little or no easy access to potable water and gas, and only 15% have electricity in their homes. Despite these conditions, the families that live there show an admirable desire to find ways of life that will allow them to continue inhabiting these arid lands.

Photo 12: Irma and Adalberto are owners of more than 9000 hectares of arid lands dedicated to raising goats in the Aguada San Roque area. They were unable to finish their basic studies in school and they have very limited income from the goats that they sell in informal markets. They are typical puesteros, or small scale farmers, of the region. Credit: Eliane Ceccon.
Photo 13: Irma roams the arid lands trying to prevent predators such as pumas (Felis concolor) and foxes (Lycalopex culpaeus) from attacking her goats, while directing and herding them to the few locations that can provide intermittent supplies of forage and water. Credit: Eliane Ceccon.

Therefore, due to the dire socioeconomic conditions mentioned above, it is necessary to conceive and launch sustainable restoration and rehabilitation projects that in addition to recovering ecological processes and functioning must also offer tangible goods and services to the local human population. In this sense, what we call “productive restoration” may be the most appropriate strategy, since it aims to recover soil productivity and offer products for the local population, along with some of the elements of the structure and function of the pre-disturbance ecosystem. (This is comparable to ecological rehabilitation as the term is used in the Society for Ecological Restoration Primer; SER 2004).

As mentioned, a critical key to successfully developing productive restoration projects in San Roque and other settlements in Argentinean Patagonia is to know and understand the socio-ecological context of the local population, in cultural, educational, health, and socio-economic terms, and also the values that local people assign to native plant species. We carried out surveys and interviews among the local inhabitants and visits to each of their landholdings, which allowed us to evaluate the knowledge and the value that they gave to the local flora, and their interest in cultivating native (and introduced) species in future restoration projects. The ecological attributes of selected species, and their importance for the productive restoration were obtained through a literature review. This review arises as part of Fernando Farinaccio’s PhD work. For more details and information, read his open access paper in Ecosystems and People.

Photo 14: Sometimes family settlements are located in places where there is an outcrop with easy access to groundwater (for example, a natural spring).This settlement recently benefited from government subsidies to improve water storage, allowing them to purchase and install the two water tanks shown here. Credit: Eliane Ceccon.

Local knowledge and use value of the native flora

Puesteros that we interviewed identified a total of 44 multipurpose species, of which 38 were native. Among the most frequently mentioned native species, Prosopis flexuosa var. depressa, Atriplex lampa, and Larrea spp., were considered by puesteros to have the highest potential and promise to restore and rehabilitate their fields and landholdings. The main reasons were not only ecological, but also the multiple uses of the plants, such as providing high quality fodder for livestock, and firewood for heating and cooking.

Photo 15: A portion of a plantation of Atriplex lampa (a nutritious and palatable native shrub) carried out in 2012 in a degraded area near Aguada San Roque. A recent study has proposed this species as a “framework species” for dryland ecological restoration (Pérez et al. 2019). Credit: Laboratory for the Rehabilitation and Restoration of Arid and Semi-arid Ecosystems, National University of Comahue, Argentina.

Ecological attributes for the reintroduction and reinforcement of populations of the plant species most valued by puesteros

According to studies carried out locally, the most valued species show high and easy germination (with rates of >60%) and are relatively easy to propagate in plant nurseries (see Farinaccio et al. 2021). In addition, some of them have shown high success in terms of survival and growth in field experiments (>70%) (see Pérez et al. 2019; 2020). These species are attractive because they are food sources for vertebrates and invertebrates, and also offer thermal refuge and nest sites for seed dispersers (Farinaccio et al. 2021).

Characteristics of puesteros‘ home gardens

Home gardens are traditional agroforestry systems supporting subsistence of poor rural families, and they are usually located near people’s homes. These home garden shave also been the cradle for selection, domestication, diversification, and conservation of elements of flora and fauna, and the preservation of cultural values. In the puestero’s home gardens, a total of 44 species were identified, of which 85% were exotic, and used to obtain forest products (from afforestation), 47% for shade and other amenities, and only 40% to obtain forage, food, and medicine.

Photo 16. Puesteros often use exotic trees in their home gardens. The most frequently used species are Eucalyptus spp., Populus spp., and Tamarix ramosissima, all of which are used for shade and wind breaks. Credit: Fernando Farinaccio.
Photo 17. In some home gardens, small areas marked off with wooden or iron fences are used for the production of fruit trees, medicinal species, and forage (A). The cultivation of species for food consumption is also carried out (B), and in some cases, these species are protected from inclement weather (e.g., intense winds, and extreme low and high temperatures), through the construction of small greenhouses (C). Credit: Fernando Farinaccio.

Conclusions

The socio-ecological, economic, and cultural contexts of the Aguada San Roque community showed an unfavorable well-being panorama. Likewise, the extensive livestock production system, on which all puesteros’ depend for their subsistence, added to the intense hydrocarbon activity (fracking), have triggered an irreversible desertification process. In this context, local people recognize a low percentage of useful native species and prefer to use a large proportion of exotic species.Similar results have been documented in other studies in drylands of Argentina and the world. The low results regarding the use of native species by the local inhabitants, and the preference in the use of exotic species, show a loss of traditional ecological knowledge, which could be a consequence of the above-mentioned historical occupation of arid Argentinean Patagonia. However, they expressed motivation and interest in sharing their historical practices with restoration actions with multipurpose native species. Beyond this unfavorable panorama, the puesteros expressed motivation and interest in carrying out restoration and rehabilitation actions with multipurpose native species. The three species most frequently mentioned by the puesteros (Prosopis flexuosa var. depressa, Atriplex lampa, and Larrea spp.), were all successfully established in ongoing restoration pilot studies.

This study proposes that the interpretation of the historical, social, cultural, and ecological reality of local people is fundamental before undertaking ecological restoration and rehabilitation programs. “Top down” programs may not be successful if the local inhabitants’ needs, desires, and proposals are not taken into account. A restoration-based education program can help implement these projects successfully. The program may promote the strengthening of local capacities and the rescue of traditional knowledge; increase collective learning, to ultimately restore the historical links between local people and the native, natural ecosystem.

References cited and additional reading

Busso, C.A., O.A. Fernández. 2017. Arid and semi-arid rangelands of Argentina. In: Gaur, M.K., V.R. Squires, editors. Climate variability impacts on land use and livelihoods in drylands. New York: Springer InternationalPublishing; p. 261–291.

Coronato, A., E. Mazzoni, M. Vázquez, F. Coronato. 2017. Patagonia: una síntesis de su geografía física. Santa Cruz (Argentina): Editorial de la Universidad Nacional de la Patagonia Austral. ISBN 978-987-3714-40-5.

Farinaccio, F.M., E. Ceccon, D.R. Pérez. 2021. Starting points for the restoration of desertified drylands: puesteros’ cultural values in the use of native flora. J Ecosystem & People. 17:476-490. https://doi.org/10.1080/26395916.2021.1968035

Pérez, D.R., F.M. Farinaccio, J. Aronson. 2019. Towards a Dryland Framework Species Approach. Research in progress in the Monte Austral of Argentina. J. Arid Environments 161:1-10. https://doi.org/10.1016/j.jaridenv.2018.09.001.

Pérez, D.R., C. Pilustrelli, F.M. Farinaccio, G. Sabino, J. Aronson. 2020. Evaluating success of various restorative interventions through drone- and field-collected data, using six putative framework species in Argentinian Patagonia. Restoration Ecology. 28:44-53. https:// doi: 10.1111/rec.13025.

SER (Society for Ecological Restoration International Science & Policy Working Group). 2004. The SER International Primer on Ecological Restoration.https://www.ser-rrc.org/resource/the-ser-international-primer-on/.

30 dump truck loads of coffee pulp help restore a Costa Rican rainforest

Rakan “Zak” Zahawi is the executive director of the Charles Darwin Foundation in Galápagos, Ecuador. He and his collaborator, Rebecca Cole, partnered with a coffee processing plant to repurpose farm waste and help restore a rainforest. Read more about the project in an open access article in Ecological Solutions and Evidence.

From the very first time I saw the results of the orange peel project on the ground back in 2004 I was sold! What a brilliant idea I thought – use the waste products generated from the production of orange juice (and any related citrus products) to regenerate degraded habitats where expansive dry forests were once found. The idea was Dan Janzen’s, an ecologist at the University of Pennsylvania who has worked in northern Costa Rica for the better part of 50 years. At the time I was working for the Organization for Tropical Studies (OTS) and given that I work as an ecologist in forest restoration, a colleague thought I might be interested.

One of thirty dump truck loads of coffee pulp, locally called brosa, spread on former farmland to restore rainforest in southern Costa Rica. Photo credit: Rebecca Cole.

The idea is simple, take truckloads of agricultural waste (in this case orange peels) and spread them in a layer ~0.5 m thick across hectares of extremely degraded land dominated by forage grasses. Under the tropical sun this layer generates an enormous amount of heat, and in the process of ‘cooking’ down it asphyxiates and kills the forage grass that is notoriously difficult to eradicate. At the same time, birds and other seed dispersers visit the site, attracted by the abundant larvae helping to decompose the material. The net result is a lot of organic material and nutrients and many seeds dispersed combining to help jump-start the recovery of a degraded habitat and return it to a forested state.

I never forgot that visit and over the years that I worked in southern Costa Rica as Director of the Las Cruces Biological Station (a field station run by OTS) I always thought of trying the study there. The difference was that there was no orange production in the region but another agricultural byproduct was widely available – coffee pulp waste! I wondered – could the results of the orange project be replicated with another agricultural waste product? While the idea was always on my mind, it took more than a decade for me to actually test it after Rebecca Cole, a long-term research colleague who was based at the University of Hawaii expressed interest in collaborating.

Before: a former cattle pasture in southern Costa Rica. Photo credit: Rebecca Cole.
During: coffee pulp piled half a meter high over the experimental plot. Photo credit: Rebecca Cole.
After: the area piled high with coffee pulp rapidly grew into a secondary forest (left) while the control area remained covered in pasture grass, as it has been for decades (right). Photo credit: Rebecca Cole.

With funds secured from the March Conservation Fund, we setup a modest pilot study with a 35 × 45 m plot buried half a meter deep. That’s 30 dump trucks – or 360 m3 of material! As with the orange peel study, this land was primarily degraded pasture and would have been slow to recover on its own. We monitored this and an adjacent similar-sized plot for 2 years and the results were nothing short of spectacular. While the control treatment languished with overgrown grasses with a few shrubs, the coffee waste plot was completely transformed. The grass was smothered and in its place a patch of young trees. All species were pioneers but they are nonetheless critical to the recovery process – and the fact that they dominated the entire plot was really promising. With time it is hoped that more mature forest species will come into this system and establish – and with a young canopy of pioneers providing a little shade, the conditions are perfect for this to happen!

Drone image showing the area where coffee pulp was dumped (left) and the control plot (right) after two years. Photo credit: Rakan Zahawi.

This study is a small pilot project, but the results speak for themselves. So does the coffee industry! Every year, millions of tons of coffee pulp waste are generated and finding a way to not only dispose of this waste in an ecologically sound manner, but also use it for habitat recovery is a win-win for everybody. It is exceedingly rare for industry to be able to pair up so seamlessly with conservation and restoration that it is hard to believe. Of course, there are hurdles – such as governmental regulations that manage such waste products, but the potential here is enormous.  And the next challenge before us is to see if we can bring this idea to scale and test the methodology across big areas of degraded habitat in the tropics. We will keep you posted!

Read more about this project in a recent open-access article published in Ecological Solutions and Evidence.

Do we really need to plant a trillion trees? Tree islands are an ecologically and economically sound strategy to facilitate tropical forest recovery

Karen Holl (UC Santa Cruz) and Leighton Reid (Virginia Tech) describe lessons learned from a 15-year study of tropical forest restoration in southern Costa Rica. Their new paper is published in the Journal of Applied Ecology.

It seems that everybody from business people to politicians to even Youtubers is proposing that we should plant millions, billions, or even trillions of trees. They cite a host of reasons, such as storing carbon, conserving biodiversity, and providing income. These efforts should be done carefully and with a long-term commitment to ensure that the trees survive and to prevent unintended negative consequences, such as destroying native grasslands, reducing water supply in arid areas, or diverting attention from efforts to reduce greenhouse gas emissions.

Another important question is whether we really need to plant that many trees to restore forest. In a new paper in the Journal of Applied Ecology, we summarize some the lessons we have learned about a different approach.

Volunteer plants tree seedlings in one of our plantations in southern Costa Rica. Photo: Karen Holl

Over 15 years ago, we set up an experiment in southern Costa Rica to test whether planting small patches or “islands” of trees could speed up forest recovery for a lower cost than typical tree plantations. The idea is to plant small 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. As a result, over time these tree islands spread as they grow and facilitate the establishment of a lot more trees.

Compared to tree plantations, the tree island approach has two major benefits. First, it better simulates the patchiness of natural forest recovery. Second, it costs much less than planting rows and rows of trees.

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 15 years it appears to provide a good balance. Figure modified from Corbin & Holl (2012).

In our experiment, we planted tree islands that covered about 20% of a 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). We repeated this set-up at 15 sites in 2004-2006.

Over the past 15 years, we have monitored the recovery of vegetation, litterfall, nutrient cycling, epiphytes, birds, bats, arthropods, and more. Our data reveal a few key lessons about how to restore tropical forests more ecologically and economically.

First, our data show that planting tree islands is as effective as bigger tree plantations, despite cutting costs by around two-thirds. Compared to plantations, tree islands have similar recovery of nutrient cycling, tree seedling recruitment, and visitation by fruit-eating animals. Both tree islands and plantations speed up tropical forest recovery compared to letting the forest recover on its own. After 15 years, cover of trees and shrubs in the island planting plots has increased from 20% to over 90%.

Artist's depiction of three tropical forest restoration treatments: natural regeneration, tree islands, and plantation.
Drawing of our three treatments showing a few trees establishing in the natural regeneration plots, the tree island merging canopies merging in the island plots, and the rows of trees in the plantation. Artist: Michelle Pastor.

Second, we have found that larger tree islands are more effective than smaller islands in enhancing the establishment of fauna and flora, as larger tree islands attract more birds and shade out competitive grasses.

Third, while tree islands cost less than plantations, some landowners won’t use the tree island approach because the land looks “messier” than orderly tree plantations. Some people prefer to plant lots of trees that are valuable for timber or fruit, rather than having the diverse suite of species that are typical of a tropical forest. So, the tree island planting strategy will be more suitable in cases where the goal is to restore forest.

Natural recruitment of trees seedling in the understory of a canopy of planted trees.

Our results and those of others show that the tree island planting approach holds promise as a cost-effective forest restoration strategy in cases where there are seed sources nearby to colonize and animals to disperse them, and where the spread of tree islands is not likely to be slowed by fire or invasive species. But we need more long-term studies to judge whether tree islands will be effective in other tropical forest ecosystems and to test other questions, like how the particular tree species used affect forest recovery, or what is the best distance to leave between tree islands.

More broadly, our study shows that tropical forests can recover some species quickly but it will take many decades, or longer, for forests to fully recover. So, preserving existing rain forests is critical to conserve biodiversity and the services that intact forests provide to people.

Yes, carefully-planned tree planting can help accelerate tropical forest recovery. But, in many cases we don’t need to plant trees everywhere. Rather we should use restoration strategies that encourage trees to plant themselves.

To learn more about our research, read our new article in the Journal of Applied Ecology, visit our websites (Holl Lab, Reid Lab), or watch a 7-min. video below.

Karen Holl describes the tree planting restoration approach and our long-term experiment in southern Costa Rica.
Los investigadores principales describen el método de applied nucleation y nuestro experimento a largo plazo en el sur de Costa Rica.

Hard times for hemiepiphytes: Aroids have trouble making a comeback in second-growth forests

Estefania Fernandez Barrancos is a PhD student and Christensen Fellow at the University of Missouri St. Louis, where she is affiliated with the Harris World Ecology Center and the Center for Conservation and Sustainable Development at the Missouri Botanical Gardens. Estefania has previously written about how to restore bromeliad populations. Here she describes a recent study asking how well hemiepiphytic aroids recover in secondary forests in Panama.

Most people know aroids as the familiar swiss cheese plants found growing in hotels and shopping malls. But few people realize that the aroid family (Araceae) is the fifth most diverse plant family on Earth. These plants provide essential food and refuge for birds, bats, insects, and primates in tropical forests throughout the world.

Like many other plants, aroid populations are dropping because the rainforests where they live are being converted into farms. My new research shows that aroids are also slow to recolonize new forests that become available.

City Aroid Country Aroid

City aroid (left, Monstera deliciosa in a building), country aroid (right, Monstera sp. in a Colombian forest). Photo sources: Left Maja Dumat CCBY 2.0; Right – Thomas Croat via Tropicos.

Before I describe that research though, here is some botanical jargon for the uninitiated. Epiphytes (a.k.a. air plants) are plants that grow on other plants (but not as parasites). Hemiepiphytes are plants that grow on other plants but only for part of their lives. Many aroids are hemiepiphtyes because they start life in the soil of the forest understory and grow until they find a tree. Then they climb up the tree and live above the ground, but they always keep a connection to solid earth.

To study their recovery, I surveyed hemiepiphytic aroids in native tree plantations (9-years old), natural secondary forests (8-14-years old), and mature forests (>100-years old) near the Panama Canal. These forests are part of Agua Salud – a tropical forest restoration experiment led by the Smithsonian Tropical Research Institute. In the dense forest, I found aroids by looking for their stems coming down from the trees, then I followed the stem with binoculars until I found their leaves, which helped me identify the species. In all, I surveyed 1479 trees this way.

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Estefania Fernandez (below) and field assistant Carlos Diaz (above) look for aroids in a mature forest tree in Panama.

I found out that there were virtually no aroids in secondary forests or plantations. I recorded more than 2000 aroids from at least ten species growing on trees in mature forest, but in secondary forests and plantations I found less than 1% as many aroids and only three species.

Why do aroids have recovery troubles?

One reason for the lack of aroids could be that seeds from adult aroids in mature forests can’t reach the new forests. This seems unlikely because all of the secondary forests and plantations in my study were close to mature forests full of aroids, less than one kilometer away. Also, birds that are present in secondary forests are known to eat aroid fruits and disperse their seeds.

Another reason could be that the young forest canopy is too open for aroid seeds to germinate and grow. Unlike most plants, some aroids start out life growing away from light and towards darkness. (This has another great word: skototropism). It seems counterintuitive since most plants need light. But it is actually a good strategy. By growing away from light, aroid seedlings are more likely to run into a tree, which they need to climb up into the canopy and get to the light that they need to photosynthesize. So it is possible that there is too much light in the young forests and it keeps the aroid seedlings from finding a host tree.

Whether dispersal or establishment limits aroids in secondary forests, it is likely that more time will help. As forests become older and darker and birds bring in more seeds, aroid populations should eventually begin to recover. My research suggests that there is a considerable lag time required for aroids to recolonize disturbed habitats such as secondary forests and plantations.

More importantly, my study highlights how important it is to hold onto old forests. Forest restoration is a poor substitute for mature forest conservation. To the extent that we can prevent older forests from being cut down, it will help preserve many species of aroids as well as other plant and animal species that are threatened by habitat loss.

Aroid being pollinated by scarab beetles at Barro Colorado Island, Panama. Source: www.aroid.org.

You can read more about Estefania’s research in her new open-access paper in Tropical Conservation Science, or on other posts from Natural History of Ecological Restoration (here and here).

Virtual field trip to the Guajira desert and the Serranía de Macuira in northern Colombia

James and Thibaud Aronson describe the natural and cultural context of a little-known area of northern Colombia, home to the Wayuu people and a microcosm of arid lands worldwide.

Colombia is one of the world’s seventeen megadiverse countries.  In a few hours of travel, one can go from the sweltering Amazonian lowlands to the snow-capped peaks of the Andes. It even has a true desert, a small peninsula called la Guajira, shared with Venezuela, which constitutes the northernmost point of South America.

For most of the last 50 years, the Guajira was notoriously dangerous, principally because of drug trafficking, but things have improved in recent years. We traveled there last month, shortly after the first big rains the region had received in several years. ​ And we found that it’s a poignant example of the plight of drylands globally and their peoples.

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The Guajira peninsula, in northern Colombia, including the authors’ itinerary.

Our trip actually began in Panama, which was part of Colombia until 1903. While much smaller, Panama is also a country of contrasts. Much of the Pacific coast used to be covered in seasonally dry tropical forest, and some fragments persist today in and around Panama City itself, while the forests of the Caribbean slope, a mere 50 km away, are much wetter. A curious switch occurs near the Colombian border, where the wet forests then extend down the Pacific coast of Colombia and Ecuador – the famous Chocó-Darien rainforest, one of the wettest and most diverse tropical forests on Earth.

Meanwhile, the seasonally dry forests continue along the 1,000 km long Caribbean coast of Colombia and give way to semi-desert and then true desert (annual rainfall < 250 mm), lined by a coast with mangrove forests, and a series of lagoons and bays where flamingos and ibises add a shock of color.

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Mangroves in Bahia Hundita, Alta Guajira, showing desert woodland with tree cacti (Stenocereus griseus) and various legume trees growing on the sandstone bluffs in the background.

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Roseate spoonbills, great egrets, and a white ibis sharing a coastal wetland near Uribia.

As if this wasn’t enough contrast, halfway along the Caribbean coast rises the Sierra Nevada de Santa Marta, Colombia’s tallest mountain range, reaching 5,700 meters (18,700 feet) above sea level at the highest peak. It takes only about two hours to drive from its foothills, where toucans and monkeys chatter in the majestic trees, to Riohacha, the gateway to the desert.

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A brown-throated three-toed sloth (Bradypus variegatus) hanging by one arm in a Cecropia tree in Tayrona National Park, at the base of the Santa Marta mountains.

Alta Guajira’s desert trees and woodlands

The Alta Guajira is arid indeed, but it hosts trees, remarkable both in their exuberant diversity and their abundance, considering the high temperatures and meager rainfall. We saw what we consider true desert canopies, such as we have described in other posts. However, no desert flora exists in isolation, and indeed the kinship to the ecosystem type known as Seasonal Dry Tropical Forest (SDTF; see map above) seems to be strong.

The dominant trees of the Guajira are species of Prosopis, Caesalpinia, Vachellia (formerly part of Acacia s.l.), Parkinsonia and other legume genera, accompanied by Bursera, Capparis relatives, Bignoniaceae, and other species common in the dry forests of Central and South America, and 3 kinds of tree cacti (Stenocereus, Pilocereus, and Pereskia), growing close together, often covered in climbing vines. In particular, it was interesting to see bona fide desert woodlands dominated by two well-known legume trees, Prosopis juliflora and Vachellia farnesiana, which are widespread and often strongly invasive in other parts of the world, but not here! Fascinating biogeographical and ecological questions abound in this poorly explored region, many of which are relevant to conservation and restoration.

Regarding  landscape ecology in the region, the vegetation is curiously like a patchwork, alternating between dense desert woodlands, nearly pure tree cacti stands, sometimes with a dense grass cover, and sometimes not, and frequent saline flats where nothing grows. In our opinion, the human element, namely land and resource use history, is paramount to understanding what one sees when travelling here and trying to ‘read’ the landscapes.

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Mixed patch of tree cacti and spiny legume trees with a surprising amount of grass understory. Elsewhere under similar stands, for no clear reason, there is no grass cover at all.

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A track of the Alta Guajira, near Nazareth, at the base of the Macuira hills where the notorious Prosopis juliflora, known in Colombia as Trupillo, is so exuberant and long-lived it forms a natural tunnel above this track.

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Prosopis juliflora colonizes newly exposed beach dunes, in areas where the shoreline is receding. Here, at Camarones, it occurs alongside Calotropis procera, a woody weed of the Apocynaceae known in English as giant milkweed, and familiar throughout the Caribbean islands, the Middle East and drylands of Africa. It survives because of its toxic milky latex where most other plants get eaten out by livestock.

Other standouts are the beautiful Palo de Brasil, Haematoxylum brasiletto, with its unusual fluted trunks and Pereskia guamacho, an enigmatic ‘primitive’ tree cactus with true leaves and one of the most exquisite tasting fruits we know. This is one of the least well-known but most intriguing of all desert trees to our minds.

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Typical landscape of the northern Guajira desert woodlands, with an even-aged stand of one of the several neotropical legume trees known as Brazilwood: Haematoxylum brasiletto, or Palo de Brasil in Spanish.

Despite those common names, this species is in fact only found wild along Caribbean coastlines from Colombia and Venezeula, all the way north to both coasts of Mexico. The scientific name is thus a misnomer. The most famous Brazilwood tree is another legume, Paubrasilia echinata (= Caesalpinia echinata) that once grew abundantly along the Atlantic coast of Brazil, as a large tree with a massive trunk, reaching up to 15 meters tall. Today, it’s almost entirely gone in the wild, and mostly planted in gardens and along roadsides. It was prized for the bright red dye obtained from the resin that oozes from cut branches or trunks. The dye was widely used by textile weavers in the Americas and Europe in the 17th-19th centuries. The tree also provided the wood of choice for high quality bows for stringed instruments and was widely used for furniture making as well. So important was its economic value that the country was named after it, originally Terra do Brasil (Land of the Brazilwood), later shortened to Brazil. Recently it was designated as sole member of a new genus, as part of a comprehensive revision of the entire genus Caesalpinia, carried out by an international team of experts.

It’s curious that H. brasiletto bears the same common name as P. echinata, since the two trees are nothing alike, apart from their red sap and heartwood. Little literature exists for H. brasiletto, and we are embarking on some detective work to shed some light on this puzzle. We go into detail as these are both relatively fast-growing trees with great economic as well as ecological value. They would both be excellent candidates for inclusion in ecological restoration work and are both in dire need of conservation efforts.

Wayuu: Alta Guajira’s Indigenous People

This desert also hosts a fairly large human population. The Guajira is the home of the Wayuu, Colombia’s largest surviving Indigenous group and, along with the Navajo, one of the last desert-dwelling peoples in the New World. These fiercely independent people, organized in 17 matrilineal clans, were never subjugated by the Spanish, and even today the Guajira region functions mostly in isolation from the rest of the country. As we were heading well off the beaten track, we needed a guide, a 4 x 4 jeep in good condition, and a skilled driver to navigate the meandering and unmarked desert paths.

Despite an ancient history of human presence, and some periods of intensive exploitation and intervention (such as a pearl harvesting boom that took place soon after European explorers arrived), the ecological condition of the region at the landscape scale is remarkably good. Indeed, apart from the salt works in the small town of Manaure, which produce two thirds of Colombia’s salt, and El Cerrejón, South America’s largest open-pit coal mine, in the south of the Guajira, there is no major industry.

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Typical traditional salt works at Manaure worked by hand by local men and women just as they have for generations.

And the isolated people who dwell here – fishermen, shepherds, and weavers – are right out of a Gabriel García Márquez story. Indeed the author, most famous for One Hundred Years of Solitude, grew up on Colombia’s northern coast, speaking both Spanish and the Wayuu language, Wayuunaiki. As we traveled deeper into the desert, we traversed small settlements with simple houses made of wood and yards surrounded by tree cacti hedges.

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The Wayuu village of Boca de Camarones, in the south of the Guajira peninsula, showing the living hedgerows of columnar cacti produced from tall stanchions. In the background, surrounding the homes, are Trupillos, and good specimens of Dividivi Libidibia coriaria (formerly called Caesalpinia coriaria).

This third caesalpinoid legume tree, closely related to the two Brazilwoods mentioned above, is the source of another lovely red dye, derived in this case from its pods. Until recently, there was an annual festival in Camarones, in honor of this formerly major economic plant product. The tree was also used as an important source of tannins. Like Paubrasilia echinata, it deserves more ethnobotanical and biogeographical studies.

Here, as in many other arid lands, goats and sheep are important for the Wayuu people, as a source of food and social currency. For example bride price during arranged weddings, and gifts for guests attending vigils of important elders and healers, are paid to this day in heads of live goats or sheep. Historically, mules and donkeys were very common as well, but now they are increasingly replaced by motorcycles.

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Small children following a flock of desert-hardy sheep in Boca de Camarones. The peaks of the Sierra Nevada de Santa Marta are visible in the background.

Crown jewel of the Alta Guajira

The crown jewel of this desert, its best kept secret, is the Serranía de la Macuira, a small mountain range (serranía meaning “small sierra” in Spanish) in the northeast of the peninsula. This miniature sky island is almost impossibly lush, thanks to moisture-bearing clouds that shroud its upper reaches. They feed streams that flow year-round, and sustain many kinds of trees that grow to well over 10 meters tall.

As one climbs the slopes of the Macuira, the humidity dramatically increases and the parched lowlands, with their desert woodlands, blend perceptibly into a seasonally dry tropical forest reminiscent of those we had seen in Panama. A little-known fact: seasonally dry tropical forests are the most endangered of all tropical forest types, and those in La Guajira are worthy of much greater research, conservation, and restoration.

Climbing higher still, the mid- and upper ranges of the Macuira seem like another world. Most astonishing of all, there is apparently an abrupt transition above 550 meters, and the higher reaches are covered in true cloud forest, with mosses, epiphytic orchids, tree ferns, and dozens of tree species that otherwise occur hundreds of kilometers away! This is probably the only place in the world where cloud forest is found less than 5 km from true desert. Fortunately – from a conservation point of view, but unfortunately for us – the upper peaks of all three peaks of the Macuira are sacred to the Wayuu, and completely off-limits, to native people and visitors alike. Try as we might, we were unable to get permission to hike up there.

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Seasonally dry tropical forest on the northeastern facing slope of the Macuira, where precipitation is much higher than in the surrounding lowlands occupied by desert woodlands.

Even though the whole Macuira is officially protected as a national park, the reality is more complicated. While walking inside the park, we encountered recently cut trees, the ubiquitous goats, and even a Wayuu man hunting birds with a slingshot in broad daylight. The beautiful continuous tree canopy covering most of the slopes stands in stark contrast to the severely eroded, nearly bare hilltops, on which stand small Wayuu homesteads. Still, the presence of clear ecotones speaks to mostly healthy landscapes.

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The severe erosion around a small Wayuu farm inside the Macuira National Park.

Alta Guajira’s ecological future

The pressures on the Guajira’s ecosystem health include a large mine (El Cerrejón, mentioned above), overgrazing by domestic livestock, and stark poverty facing the native people and more recent immigrants. But there are positive factors as well. There are progressive laws in Colombia related to ecological restoration. Moreover, since 2012, Colombia has a National Restoration and Rehabilitation Plan (pdf), as well as a Law of Remediation, which imposes large environmental offset payments from large-scale development projects (like hydroelectric dams) to underwrite conservation and restoration work. Moreover, the national park system, within its network of 56 protected areas, harbors populations of almost half of the 102 Indigenous peoples in the country, and in the case of Macuira, this is clearly not just a paper park idea.

Still, the national park (25,000 ha in size; officially designated in 1977), operates with a skeleton staff attempting to carry out an ambitious management plan (pdf) despite an insufficient budget. Staff and volunteers provide short tours to day-visitors, and maintain some fenced-off livestock exclosure plots, where they are studying natural regeneration. Daily interaction with the Wayuu living in the park appear to be harmonious, and indeed there is a clear sense that part of the Park’s mission is to restore and protect the Wayuu people’s natural and cultural heritage. Recently, the Instituto Humboldt, Colombia’s stellar national research institute, has established permanent plots in the Macuira range as part of a series of 17 plots including all the tropical dry forest types in Colombia. In the Macuira, this work is done in collaboration with botanists from the Universidad de Antioquia, in Medellin. Furthermore, researchers at Kew, the Smithsonian Institute, and many conservation NGOs are all developing collaborations with the Colombian government to explore and help the country move forward with green development.  The Missouri Botanical Garden also has long-standing MoUs for joint research with 3 different institutions in Colombia, with bright prospects for deepening cooperation in the future.

Like many Indigenous peoples around the world, the Wayuu are at a crossroads. Their language and some of their traditions are still alive and well, but others have already faded. There are few legal sources of income in the harsh desert, the ancestral Wayuu land. How will they manage in the future? What can they do to adapt?  Some, like our guide, José Luis, are trying to change mentalities, but they clearly need more help.  As throughout Colombia, there is clear and urgent need to build on the alpha-level studies already underway, and move onto applied ecology, agroforestry and land management programs, including community-based restoration programs and ecotourism in conjunction with the national parks.

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Our Wayuu guide José Luis Pushaina Epiayu (on the right) and Macuira park ranger Ricardo Brito Baez-Uriana (on the left), talking about birds with a local Wayuu family.

What does the Black-faced Antthrush tell us about tropical forest restoration?

Anna Spiers (University of Colorado Boulder) describes a recent field experiment done with Emma Singer (Hamlin College) and Leighton Reid (CCSD) during an Organization for Tropical Studies Field Ecology Course in Costa Rica.

Bird diversity and forest restoration are synergistic. Birds facilitate forest regeneration through seed dispersal, pest control, and pollination. Forest restoration replenishes lost bird habitat by providing food, protection from predators, and suitable territory for breeding and nesting. Monitoring bird communities in a regenerating forest is an effective strategy to gauge the success of restoration.

While some birds are flexible regarding the quality of their habitat, others require a narrower set of conditions to survive. One such bird is the Black-faced Antthrush (Formicarius analis), a medium-sized, ground-dwelling insect-eater, easily distinguished by its plaintive song and chicken-like strut. The bird spends its days flipping over leaves and sticks with its bill to expose tasty ants, beetles, and other arthropods (and sometimes small vertebrates). A member of a bird family highly threatened by forest fragmentation (Formicariidae), the Black-faced Antthrush is known to disappear from small forest fragments and to struggle crossing even narrow strips of open space. Finding such sensitive birds in a regenerating forest is a positive signal that forest restoration is increasing habitat for forest-dependent species.

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Black-faced Antthrush (Formicarius analis) strutting across the rainforest floor. Image: Luke Seitz/Macaulay Library at the Cornell Lab of Ornithology (ML54054261).

Earlier this month, we did an experiment to find out how different forest restoration strategies affect the Black-faced Antthrush. Specifically, we tested whether the bird exhibited a stronger territorial response in tree plantations, naturally-regenerating secondary forests, or areas where patches of trees (tree islands) had been planted to stimulate forest recovery. We expected to find that birds would be more defensive of areas where trees had been planted, given that these areas had a more closed canopy and more leaf litter for the birds to pick through for arthropods.

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Leighton holds up a speaker to conduct a bird call playback. Unsurprisingly, there was no response in this scrubby, abandoned pasture (one of the control points in our experiment). Image: Martha Bonilla-Moheno.

To test the bird’s territorial response, we amplified a locally-recorded sound file of the bird’s vocalization and recorded its response. We noted how long it took for the bird to respond, how many notes it sang in response, and how close it approached the speaker. For this species, a short call with 4 notes is a “hello”, but a long call with upwards of 12 notes is a warning to let the other birds know that this territory is taken.

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Our study area at Las Cruces Biological Station in southern Costa Rica. Each of the two restoration sites contained a tree plantation, a natural regeneration area, and a “tree island” area where patches of trees were planted to kick-start forest recovery. Image: Google Earth 2018.

Antthrushes defended restoration areas where trees were planted

As we expected, Black-faced Antthrushes responded more quickly and more forcefully when we taunted them with calls broadcast from tree plantations and tree island plantings – an indication that they were expending more energy to defend these areas. However, we only found this at one of the two restoration sites. The other site was a veritable antthrush desert with not a single response during any of our trials. Leighton’s collaborator Juan Abel Rosales often finds Black-faced Antthrushes at both sites, but this second site is near a road and dogs occasionally wander into the regenerating forest, possibly causing birds to temporarily abandon this area.

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Black-faced Antthrushes responded quickly and with many tooting notes when we played their song to them from tree islands, plantation, and mature forest, but they responded not at all in abandoned pastures or in natural regeneration. The data representing restoration treatments are from one site only – at the other site we recorded no birds during any trials.

Tree islands and plantation had a couple of habitat features that natural regeneration lacked. First, the understory was more open, providing ground-dwelling birds with greater visiblity. Second, planted areas also had deeper leaf litter, and leaf litter is essential for a bird that makes a living flipping leaves to find its dinner.

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Understory comparison between natural regeneration (left) and a tree plantation (right). Both have been recovering for 15 years. Natural regeneration vegetation is thick and still grassy from pasture days. A closing canopy in the tree plantation produced a thinner, more visible understory, with lots of nice leaf litter, full of delicious arthropods.

So what does the Black-faced Antthrush tell us about forest restoration?

 It may be telling us two things. First, restored forests growing up alongside remnant ones can be valuable habitat worth defending. When birds spend time calling, that is time that they do not spend foraging, and they can pay a price with their energy budget. Second, tree planting may create habitat for these birds faster than natural forest regeneration – although natural regeneration is highly variable from site to site, and we only found a pattern at one site right next to an old-growth forest. Promisingly, we did not see a difference between tree islands and the tree plantation, which suggests that we could plant fewer trees and still see the return of a forest-dependent bird species within about 15 years.

For more information about the Islas Project (with the tree islands) see previous NHER posts here, here, and here. Thanks to Bert Harris for some of the ideas that we used in this project!

 

 

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.

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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.

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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.

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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.

 

Seedlings planted for Brazilian forest restoration are not representative of tropical tree biodiversity

A collaborative research project involving MBG’s Center for Conservation and Sustainable Development, the Tropical Silviculture Lab at the University of São Paulo, and the PARTNERS research coordination network highlights important differences between the native tree flora of the Brazilian Atlantic Forest and the species that are widely planted for ecological restoration projects.

The Brazilian Atlantic Forest is a global biodiversity hotspot. This designation denotes two things. First, the Atlantic Forest is exceptionally and uniquely biodiverse. Second, the biodiversity of the Atlantic Forest is exceptionally threatened. This once-vast biome historically stretched from northern Argentina to Brazil’s eastern tip in Rio Grande do Norte, but it is now reduced to about 12% of its original size, and most of what remains exists as small, isolated fragments.

During the past decade, a major, multilateral effort has been undertaken to staunch biodiversity loss by doubling the size of the Atlantic Forest through ecological restoration. The Atlantic Forest Restoration Pact is composed of more than 270 private companies, governments, NGOs, and research organizations. It aims to restore 15 million hectares of Atlantic Forest by 2050.

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The Atlantic Forest biome: a global biodiversity hotspot and the site of the most ambitious tropical forest restoration project on the planet. Map imagery from NASA via Wikimedia Commons.

Atlantic Forest restoration projects are characteristically thorough and well-documented. For example, they often include high diversity plantings more than 80 tree species. Yet until recently there had never been a systematic study to evaluate how well these restoration plantings represented the Atlantic Forest biodiversity they aimed to protect.

Dr. Pedro Brancalion is a professor at the University of São Paulo’s agricultural school in Piracicaba, Brazil, where he co-directs the Tropical Silviculture Lab. Five years ago, he approached me at a meeting of the Society for Ecological Restoration in Madison, Wisconsin, and over a beer he told me about a dataset that he thought could shed light on the how well Atlantic Forest restoration projects were conserving tree biodiversity. The dataset consisted of seedling donation records from the NGO SOS Mata Atlântica. Between 2002 and 2015, the NGO donated more than 14 million tree seedlings to 961 restoration projects. By comparing the species composition in these records to tree species living in mature forests, we could see what elements of biodiversity might be missing and how this could be affecting carbon stocking – an important factor in mitigating global climate change.

Even in high diversity plantings, many of the most threatened tree species were not included.

Last month, Pedro and our collaborative team published a paper in Conservation Letters describing our results. We found that restoration projects in the Atlantic Forest biome had included 416 tree species out of the >2,500 tree species known from mature and old-growth forest fragments. This is an impressive figure, but the team discovered that it reflects a highly biased subsample of the Atlantic Forest tree flora. The most under-represented species were those with large seeds that are dispersed by animals. Animal-dispersed trees make up as much as 89% of tree species in some parts of the Atlantic Forest and include some of the most threatened species.

The reason that large-seeded, animal-dispersed species are being used less often was probably related to the cost and challenges of collecting and growing seeds. Large-seeded, animal-dispersed trees are more expensive to purchase from nurseries than small-seeded or wind-dispersed species. Because they are energetically expensive to produce and are contained within large fruits, trees tend to produce large seeds in relatively low quantities, with just one or a few seeds per fruit. They are generally found in remote forest areas, and seed collectors have to compete for them with seed-eating animals, like peccaries and agoutis. Once large seeds are collected, they also take up considerably more space in storage and production facilities.

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In our analysis of animal-dispersed tree species, seed diameter explained 87% of the variance in seed price. Large seeds like those of Caryocar brasiliense were much more expensive than small ones, like Ficus guaranitica. Grid size: 1 mm. Photos reproduced from C. N. Souza Junior & P. H. S. Brancalion (2016).

The absence of large-seeded, animal-dispersed tree species in restoration plantings has important implications for biodiversity conservation. First, fewer large-seeded trees means less food for large birds, some of which eat mainly large fruits. Second, these species are sometimes overharvested for timber and have difficulty recolonizing forests from which they have been removed. So the fact that large-seeded, animal-dispersed trees are under-represented in restoration projects means that even if the ambitious restoration goals of the Atlantic Forest Restoration Pact are met, the increase in forest cover may not improve dispersal between fragmented populations of the most vulnerable species.

Large-seeded tree species also tend to store carbon more densely than small-seeded species. This tendency is related to large-seeded species growing slowly in the shady understory of the Atlantic Forest and their gradual formation of dense wood, which is rich in carbon. We simulated potential carbon stocking in restored forests and compared it to mature forests, and our results showed that under-representation of large-seeded, animal-dispersed trees could cause a 2.8-10.6% reduction in carbon storage. Based on the current price of carbon, this loss could represent $17-63 USD per hectare in lost carbon credits.

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Many Atlantic Forest restoration projects are quite isolated. A large seed would have a hard time reaching sites like this forest in a sugarcane matrix. Photo by Pedro Brancalion.

Reduced capacity for biodiversity conservation and carbon stocking sounds like bad news, and indeed it is not ideal. However, restoration ecology moves forward by identifying problems and seeking scientifically-based solutions to overcome them. Knowing that large-seeded, animal-dispersed trees are under-represented in restoration plantings means that we can turn our attention to innovative solutions.

For example, new policies could help bridge the gap between Brazil’s exceptional tree biodiversity and the relative paucity of species being used for ecological restoration. One way this could happen would be for the Brazilian government to subsidize the cost of producing large-seeded, animal-dispersed tree seedlings. This could be done through financial incentives or potentially by opening some forest reserves for seed harvesting, to make it easier for collectors to acquire these species. Facilitating uptake by reducing costs would be a carrot. A stick could be to legally mandate some representation of these species in future restoration plantings.

Market solutions may also exist. Based on our calculations, adding more large-seeded, animal-dispersed species to restoration plantings would increase carbon storage and carbon credits, offsetting the cost of the expensive seedlings and creating a net gain of $3-32 USD per hectare.

Banner image: Sterculia striata (Malvaceae). Photo by Mauricio Mercadante. CC BY-NC-SA 2.0.

Drivers of epiphyte recovery in secondary forests in southeastern Brazil

Alex Fernando Mendes is an undergraduate researcher in the Tropical Silviculture Lab at the University of São Paulo, Brazil. He describes his thesis project, undertaken in dozens of forest fragments in the endangered Atlantic Forest biome. Currently, Alex is analyzing his data as a visiting researcher in the Center for Conservation and Sustainable Development.

Historically, intensive agriculture in the Brazilian Atlantic Forest has caused large-scale deforestation of this biome. However, new legal requirements, land exhaustion, and the shifting priorities of farmers have recently allowed forests to regenerate on some formerly farmed lands. Given the unique nature of the Atlantic Forest, its high species endemism, and its potential for providing ecosystem services, the Tropical Silviculture Lab (LASTROP) at the University of São Paulo, coordinated by Prof. Pedro Brancalion and its partners, initiated a project in 2014 to better understand the structure and composition of these new forests.

However, forests aren’t made solely of trees. Among the plant components of a forest, there are others life forms such as epiphytes, lianas, and herbs that contribute to biodiversity and provide food, water, and shelter for many animal species. Epiphytes are plants that use other plants as support. Due to their sensitivity to environmental changes, epiphytes can be used as bioindicators. Therefore, we asked how these plants are doing in these young regenerating forests. And what landscape and local attributes facilitate or hinder their recolonization?

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Epiphyte species found in second-growth forests of the Atlantic Forest – A) Philodendron bipinnatifidum Schott; B) Lepismium houlletianum (Lem.) Barthlott.; C) Ionopsis utricularioides (Sw.) Lindl.; D) Catasetum fimbriatum (E. Morren) Lindl. & Paxton; E) Aechmea bromeliifolia (Rudge) Baker; F) Billbergia sp.

 

To try to answer these questions, we are studying the epiphyte communities in 40 second-growth forests (i.e., forests that were once completely cut down). We are considering three landscape drivers (distance from watercourses, distance from forest edge and forest cover in a 1-km buffer around the remnant) and four local drivers (previous land use, forest age, liana abundance, and tree basal area). We expect that forests close to watercourses would provide the moisture required by epiphytes. We expect to find more epiphytes further from the forest edge since forest cover in more conserved forests may limit their establishment. Since our forests regenerated from abandoned eucalyptus plantation and pastures, we want to check if the non-native eucalyptus could act as a filter preventing epiphytes recolonization. We also expect that older forests and forests with more basal area could house more epiphytes than young forests. Finally, field observations made us wonder if lianas could compete with epiphytes by occupying the same niche.

Of the more than 6,000 phorophytes (trees that could support epiphytes) sampled in these 40 forests we found 398 epiphytes belonging to 21 morphospecies distributed in 4 families (Araceae – 1 species, Bromeliaceae – 14 species, Cactaceae – 5 species, Orchidaceae – 5 species). Only three species, Tillandsia pohliana, Tillandsia tricholepis, and Ionopsis utricularioides, represented more than half (59.5%) of all epiphytes found in second-growth forests. The genus Tillandsia was expected to be abundant in these young forests, since these are disturbance-adapted species that can even be found growing on power lines in cities.

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Epiphytes of the genus Tillandsia (Bromeliaceae) are often found in extreme microenvironments in urban areas (Photos by A. Mendes, 2017).

Our analysis is in progress, but our preliminary observations suggest that forests closer to watercourses and closer to forest edge are more likely to have epiphyte recolonization than forests far from edge and watercourses. Forests regenerated on pastures have more epiphytes than those on abandoned eucalyptus plantation. Our dataset will soon be upgraded with new forest types: conserved and disturbed old-growth forests, and mixed tree plantings for forest restoration, totaling approximately 70 forests with epiphyte samples.

With this research, we hope to find out the local and landscape factors that contribute to epiphyte recolonization in second-growth forests. In practice, this will allow us to locate sites with limited potential for spontaneous colonization of this life form to take actions that promote colonization and establishment, such as introducing individuals. Finally, by identifying epiphytes species that are more sensitive to disturbance, we can focus our reintroduction interventions.

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Small, remnant forests are surrounded by cattle pastures in southern Brazil.

Note: The image of Philodendron bipinnatifidum featured at the top of this post was taken by David Stang.