# What are we planting in ecological restoration of tropical open ecosystems? A systematic review of controlled experiments

Natália F Medeiros, Elise Buisson, Fernando a O Silveira

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Natália F Medeiros, Elise Buisson, Fernando a O Silveira. What are we planting in ecological restoration of tropical open ecosystems? A systematic review of controlled experiments. Acta Botânica Brasílica, 2025, 74 Brazilian Botany Conference, 39, pp.e20250067. 10.1590/1677-941x-abb-2025-0067. hal-05408958

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# What are we planting in ecological restoration of tropical open ecosystems? A systematic review of controlled experiments

Natália F. Medeiros<fnref n="1" />* ID, Elise Buisson<fnref n="2" /> ID, Fernando A. O. Silveira<fnref n="1" /> ID

## ABSTRACT

Tropical grasslands, savannas, and shrublands comprise open ecosystems that are experiencing growing anthropogenic pressures and lack theoretical and practical knowledge on ecological restoration. Natural regeneration in these ecosystems is very slow and often unfeasible, making planting and seeding increasingly important restoration techniques. We conducted a systematic review to investigate the richness, origin, and functional groups of the plant species used to restore tropical open ecosystems. Identifying species and their functional groups used in restoration experiments allows us to better plan, implement, and monitor restoration. We identified 220 plant species introduced to restore tropical open ecosystems in 37 controlled restoration sites located in Brazil, Australia, the USA, Kenya, Venezuela, South Africa, and Congo. Nearly 55% of plant species were shrubs or trees, and 80% were native to the country where they were used. Strikingly, 51% of the plants were used only once. We found no evidence of biotic homogenization in the restoration of tropical open ecosystems. Only six reintroduced species were classified as endangered, suggesting an untapped potential for enhancing conservation-oriented restoration. Our results indicate that despite the predominant use of native species, woody species remain disproportionately planted compared to herbaceous species, which constitute the bulk of diversity in grasslands and savannas.

**Keywords:** active restoration, grassland, savanna, shrubland, tropics, tropical ecosystem, tropical grassy biome

<footnote n="1">¹ Universidade Federal de Minas Gerais, Departamento de Genética, Ecologia e Evolução, Centro de Síntese Ecológica e Conservação, Belo Horizonte, MG, Brazil.</footnote>

<footnote n="2">² Avignon Université, Institut Méditerranéen de Biodiversité et d'Ecologie, CNRS, IRD, Aix Marseille Université, IUT d'Avignon, AGROPARC, France.</footnote>

<footnote>*Corresponding author: nataliaf.medeiros93@gmail.com</footnote>
Introduction

Ecological restoration is a global priority in the context
of the emergent climate and biodiversity crises. Global
restoration efforts are increasingly supported by the United
Nations Decade on Ecosystem Restoration (2021-2030),
but a forest-centric view still dominates the scientific,
governmental, and media arenas (Silveira *et al.*, 2022).
Restoration efforts in tropical open ecosystems (ancient
grasslands, shrublands, and savannas) are relatively scarce
despite increasing degradation levels (Buisson *et al.*, 2022).
Open ecosystems are characterized by graminoids, herbs,
forbs, and shrubs. For instance, grasslands have no trees, and
some savannas are known to have seven herbaceous plants
for each wood species (Ribeiro & Walter, 2008). Moreover,
open ecosystems feature a unique ecosystem functioning
driven by soil properties, fire regimes, and large herbivores,
which play important roles in shaping vegetation (Buisson
et al., 2019). Open ecosystems are also biodiversity-rich,
with many endemic and threatened species (Veldman et al.,
2015; Bond, 2019; Buisson et al., 2019). Therefore, there is a
pressing need to restore open ecosystems, aiming to protect
their high biodiversity through effective management
of their main ecosystem drivers, including disturbance,
hydrological, and edaphic properties.

Active restoration approaches, including species
reintroduction and active management, can yield better
restoration outcomes in open ecosystems, as natural
regeneration in these ecosystems is extremely slow or may
never happen, depending on the degree of degradation
(Nerlekar & Veldman, 2020). Consequently, planting
and seeding native species have been employed as main
techniques aimed at enhancing the resilience of open
ecosystems and contributing to the maintenance of
biodiversity of restored sites (Wiederhecker et al., 2024).
In addition to species reintroduction, other main active
restoration strategies in open ecosystems include tree
cutting, invasive species control, livestock and wild herbivore
management, prescribed fires, and soil acidification (Buisson
et al., 2019; Haddad et al., 2021; Buisson et al., 2022;
Pilon et al., 2023; Lira-Martins et al., 2024). Therefore,
the efficiency of open biome restoration ultimately relies
on active approaches, with planting and/or seeding native
species playing important roles in restoring diverse and
resilient ecosystems (Lyon et al., 2023).

The use of native species in restoration has been
recommended for decades, but non-native species .are
still commonly used, even in forest restoration, which
has received greater attention over the years (D'Antonio
& Meyerson, 2002; Simões et al., 2024). The benefits of
non-native species in restoration can be due to their faster
growth and the income provision to landowners (timber,

forage, fruit). Despite this, the choice for non-native species
may also be constrained by the low availability of native
species on the market (Vidal *et al.*, 2020; Faria *et al.*, 2025),
which is especially true for herbaceous species, the main
component of open ecosystems (de Castilho Silva *et al.*,
2024). Furthermore, despite recent attempts to synthesize
the common practices in the restoration of tropical open
ecosystems (Pilon *et al.*, 2023; Medeiros *et al.*, 2024), a
knowledge gap remains about the identity and richness
of plant species used in active restoration approaches.
Assessing the species pool used in restoration efforts in
tropical open ecosystems can provide important insights
into the effectiveness of these initiatives. In particular,
it can help evaluate their potential to recover the diverse
herbaceous layer, protect endangered species, and mitigate
biotic homogenization.

To address this gap, we investigated the restoration
species pool used to restore tropical open ecosystems.
Specifically, we examined the richness, origin (native or
non-native), and growth forms. Under the hypothesis that
restoration practices follow the vegetation structure of
open ecosystems, we expect grasses, forbs, and shrubs to
be the most commonly reintroduced species, with a lower
proportion of trees to match the open biome vegetation
structure. Moreover, under the assumption that restoration
is grounded in ecological theory (Barbosa-Dias et al., 2025),
we expect the preferential reintroduction of native species
instead of non-native ones. Finally, to address the potential
of conservation-oriented restoration in open ecosystems
(Volis, 2019), we examined the conservation status of the
reintroduced species.

**Materials and Methods**

To assess the richness and functional groups of plant
species used in active open biome restoration - defined as
the world's ancient grasslands, shrublands and savannas
(Bond, 2019) -, we carried out a systematic literature review
of peer-reviewed articles using Web of Science and Scopus
databases using the following keywords: ("grassy biome"
OR "grassland" OR "meadow" OR "prairie" OR "rangeland"
OR "savanna"* OR "steppe" OR "shrubland" OR "cerrado")
AND ("afforestation" OR "afforested" OR "restoration" OR
"restored" OR "recovery" OR "recovered" OR "reforestation"
OR "reforested" OR "rehabilitation" OR "rehabilitated"
OR "revegetation" OR "revegetate") AND ("tropical" OR
"subtropical"). Details of the search engine and paper
screening are provided in Medeiros *et al.* (2024). Hence, we
only included controlled restoration studies, which means
studies comparing data obtained in field experiments
(laboratory and greenhouse research were excluded)
with a preserved and/or degraded site. However, here, we
used only a subset of studies, including only those that applied active restoration techniques, including species reintroduction as well as seedling planting and direct seeding (see List S1). Studies must also provide a list of reintroduced species directly on degraded open ecosystems. We retrieved geographic coordinates and the identification of the reintroduced species, and used the Kew database (POWO, 2024) to correct species names and record the origin (native or non-native, depending on where they were planted or seeded), life cycle (annual or perennial), and growth form. We completed the growth form for 11 species using the Reflora database (FFB, 2024) because such information was not available in the Kew database. Besides, we assumed that all shrubs and trees were perennials, and we used the IUCN Red List (IUCN, 2024) to retrieve species conservation status. We conducted an ANOVA test to assess the richness of different growth forms and a GLM to evaluate the origin of planted species used in open biome restoration. All analyses were performed in the R program (R Core Team, 2025) using *dplyr*, *tidyr*, and *ggplot2* packages. We made an alluvial graph to present the relationship between plant growth forms and the biome in which they were planted. To do so, we needed to assign only one growth form to each plant species; thus, we adopted a conservative approach in which, for example, if a plant species could assume an herbaceous or subshrub form, we assigned it as herb, while if it could assume a shrub or tree form, we assigned it as shrub. Hence, we classified as herbs all herb, herb-climbing-subshrub, herb-shrub, and herb-subshrub-shrub because the species could be herbaceous. Following the same criteria, we classified as

shrubs all shrub, shrub-tree, subshrub, and subshrub-shrub (Medeiros *et al.*, 2025). All other graphs were made using the ggplot2 package in the R program.

## Results

The surveyed papers provided 37 restoration experiments from 29 studies that met our criteria for inclusion (see list S1). Nine sites from six studies only provided information on the richness of species planted without providing a specific species list (e.g., mixed grassland species, native species, high number of species), and were excluded from our analyses. Most experiments were concentrated in Brazil (43%), Australia (22%), the USA (19%), followed by Kenya (5%), Venezuela (5%), South Africa (3%) and Congo (3%). Moreover, we had nine experiments in grasslands, 16 in savannas and only three in shrublands.

Globally, 220 species from 45 families were reintroduced, with Poaceae (60 species), Fabaceae (47 species) and Asteraceae (14 species) standing out as the most common families (Fig. 1; Supplementary Material - Table S1). Regarding species growth form, only 45.6% of the species were herbaceous and 53.8% were shrubs or trees, reintroduced mainly into savannas, particularly in Brazil, Venezuela and Congo (Figs 2 and 3). Besides, we have not found a difference between herbs and woody species (Supplementary Material – Fig. S1; p>0.05). Regarding species life cycles, we found 180 species perennial, 12 annuals, eight annual/perennial, and two annual/biennial species. Information was missing for 18 species. As we expected, nearly 80% of the species were native to the

<figure><img src="image_2.png" /><figcaption>**Figure 1.** The richness of plants used in tropical open ecosystem active restoration research experiments through planting or seeding. Plant richness is grouped by families. Family used only once are not represented on this figure and includes: Acanthaceae, Annonaceae, Araliaceae, Boraginaceae, Caryocaraceae, Convolvulaceae, Dilleniaceae, Eriocaulaceae, Euphorbiaceae, Gentianaceae, Haloragaceae, Lycopodiaceae, Malpighiaceae, Metteniusaceae, Moraceae, Ochnaceae, Plantaginaceae, Polygalaceae, Rutaceae, Santalaceae, Smilacaceae, Solanaceae, Urticaceae, Verbenaceae, Vochysiaceae.</figcaption></figure>

<figure><img src="image_3.png" /></figure>
<figure><img src="image_4.png" /><figcaption>**Figure 2.** The growth-form of the plants used in tropical open ecosystem active restoration research experiments through planting or seeding experiments by country.</figcaption></figure>

<figure><img src="image_5.png" /><figcaption>**Figure 3.** The proportion of herbaceous, shrub and tree species (total of 220 species) used in active restoration of tropical open biomes: grassland (nine experiments), shrubland (three experiments) and savanna (16 experiments). It is important to note that we adopted a conservative approach to classify the plant growth form. Therefore, we classified as herbs all herb, herb-climbing-subshrub, herb-shrub, and herb-subshrub-shrub because the species could be herbaceous. Following the same criteria, we classified as shrubs all shrub, shrub-tree, subshrub, and subshrub-shrub. Finally, we classified as trees the species that can only be trees. The numbers at the left panel indicate the number of species introduced in each functional group, with herbaceous species representing 45.6%, shrubs 28.8% and trees 25%.</figcaption></figure>

<figure><img src="image_6.png" /></figure>
country (Fig. 4; Supplementary Material - Fig. S2, p < 0.001), indicating that few non-natives are used to restore open ecosystems. Most non-native species reintroduced belonged to Poaceae (12), Fabaceae (3) and Gentianaceae (1) families, including invasive, fast-growing invasive African grasses such as *Urochloa eminii* and *Urochloa dictyoneura* planted in Venezuela, and forage legumes such as *Stylosanthes scabra* and *Stylosanthes hamata* planted in Australia. Ten of the 37 restoration experiments used at least one non-native species, usually without a clear justification. Moreover, we found that only 24 species were planted in more than five

restoration experiments, while most species (107 plant species) were used only once (Fig. 5). Among species with available IUCN classifications, most were listed as 'Least Concern' (76 species). Additionally, four species were classified as 'Vulnerable' (*Eucalyptus populnea*, *Dipteryx alata*, *Mimosa caesalpiniifolia*, *Metrosideros polymorpha*), three species as 'Endangered' (*Cynodon dactylon*, *Amburana cearensis*, *Andropogon leucostachyus*), two species as 'Near Threatened' (*Eucalyptus camaldulensis*, *Pterogyne nitens*) and one species as 'Data Deficient' (*Myracrodruon urundeuva*). IUCN classification was unavailable for 145 species.

<figure><img src="image_7.png" /></figure>

**Figure 4.** The origin of plants used in the 28 restoration experiments of tropical open ecosystem active restoration through planting or seeding. Note that we found another nine experiments of planting/seeding species in tropical open ecosystems that were not represented here because they did not provide the plant species identification.

<figure><img src="image_8.png" /></figure>

**Figure 5.** The bar graph represents how many times a single species was used in different tropical open ecosystem restoration experiments. The circle graph represents the growth-form proportion of the most used species (those used in five to eight experiments).

<figure><img src="image_9.png" /></figure>
Discussion

Our review is based on a relatively low number of
studies and *in situ* restoration experiments, highlighting
the scarcity of controlled restoration studies of tropical
open ecosystems using standardized, reproducible and
clear protocols (Medeiros *et al.*, 2024). This emphasizes
the importance of improving and expanding restoration
science to support policies that promote and assist the
restoration of open ecosystems worldwide (Lyons *et al.*,
2023). Despite a relatively high richness of plant families
retrieved in restoration experiments, we still need to
increase species richness in the restoration species pool
for open ecosystems, as most experiments reintroduced
less than three species from all plant families. Tropical
open ecosystems are extremely species-rich (Murphy
*et al.*, 2016), and increasing taxonomic, functional and
phylogenetic diversity in plantings is likely to promote
more biodiverse and resilient restored ecosystems (Buisson
*et al.*, 2022; Mazzochini *et al.*, 2024). For example, grasses
are important in open ecosystems, but forbs also have an
important, yet underappreciated role in these ecosystems,
as they promote biodiversity by supporting pollinators and
herbivores (Siebert *et al.*, 2024). Thus, we highlight the need
to diversify the restoration species pool and encourage the
reintroduction of forbs in open biome restoration (Silveira
*et al.*, 2022; Nerlekar *et al.*, 2024).

The mismatch between the planted species pool and the
reference open biome, like savannas and grasslands, was
identified here. Only 45% of the species were herbaceous,
despite savannas having an herb:wood species ratio of
nearly 7:1 (Ribeiro & Walter, 2008; de Castilho Silva et al.,
2024). Herbaceous species dominance in open ecosystems
is often neglected in open biome restoration, exemplified
by a 1:1 herbs and woody species ratio (Supplementary
Material – Fig. S1; p>0.05). Savannas were the main type
of biome targeted in restoration experiments. This biome
comprises open-canopy ecosystems with sparse and low-
structure trees (Bond, 2019) that have no benefits from
increasing tree density, which results in the loss of shade-
intolerant herbaceous species (Wieczorkowski & Lehmann,
2022). A positive example is the use of shrubs to restore
shrublands in the USA. To better align restoration efforts
with open ecosystems ecology, we reinforce that trees and
shrubs should be more carefully used, respecting their
proportions in the native open ecosystems, especially in
grasslands and savannas (Buisson et al., 2022).

Most of the reintroduced species were native; however,
the reintroduction of common non-native species raises
concerns, as they are fast-growing invasive grasses and
forage legumes. Combining fast-growing grasses and
N-fixing species usually stems from the idea that the
restoration of open ecosystems needs to increase vegetation
cover and improve soil nutrients, a misconception derived
from forest restoration (Silveira et al., 2022). For example,

many plant species in open ecosystems do not form
mutualistic interactions with mycorrhizal fungi because
they have evolved root specializations (Abrahão *et al.*, 2019),
thus not requiring fertilization or inoculation treatment
(Silveira *et al.*, 2022). Frequently, experiments had no clear
justification for using non-native species. Perhaps, outdated
paradigms from the 1990's such as ideas stemming from
forest restoration (Lovera & Cuenca, 1996; Cuenca *et al.*,
1998), may help explain the use of non-native species in
some cases. Another possible reason for using non-native
species is the fact that legislation often fails to recognize
the value of native herbaceous species in recovery and
rehabilitation projects (Pimenta & Fonseca, 2021). Finally,
another possibility is the paucity of native seedlings in the
market (Ladouceur & Mayfield, 2017).

Regarding the species life cycle, we found that most of
the species were perennial, which is consistent with the
evolution of strategies to deal with natural disturbance
regimes common to open ecosystems, such as recurrent
fires and grazing (Bond & Midgley, 2001). Many open
ecosystems are characterized by species with persistent
niches, which describe the strategies that allow species to
survive and grow under high disturbance regimes (Bond
& Midgley, 2001). However, woody species were the most
commonly reintroduced species (Fig. 2), which contrasts
with our expectations and supports the idea that the
restoration of grasslands and savannas is disconnected from
ecological theory (Barbosa-Dias et al., 2025). To overcome
this misconception, focus should be given to herbs and
forbs in grasslands and shrubs in shrublands through the
development of cultivation and propagation protocols (Vidal
et al., 2020).

Concerns have been raised about the extent to which
common restoration practices favour widespread and easily
grown species with high survival and propagation knowledge
promoting biotic homogenization in restored sites (Holl et
al., 2022). In some cases, restored forests can be floristically
more similar to one another than to reference ecosystems
(Almeida et al., 2025). In tropical open ecosystems, the idea
of biotic homogenization is not supported, given that a few
herbaceous species have been reintroduced in more than
five experiments. However, homogenization may have gone
undetected because of low species richness in open biome
restoration globally.

Conservation-oriented restoration stems from the idea
of reintroducing threatened species as a complementary
approach to traditional restoration (Volis, 2019). Most of the
reintroduced species had an IUCN classification unavailable
or were listed as 'Least Concern', which points out that
open biome restoration has an untapped potential to move
towards conservation-oriented practices by reintroducing
threatened plant species and integrating restoration
efforts into broader conservation efforts (Volis, 2019).
Besides, the species pool used in the active restoration of
open ecosystems through species reintroduction remains

<figure><img src="image_10.png" /></figure>
relatively small compared to the high plant biodiversity and endemism of tropical grasslands, shrublands and savannas (Murphy *et al.*, 2016). Our results agree with those of Vidal *et al.* (2020), who found an underrepresentation of savanna specialists in nurseries, which may lead to afforestation and to other negative consequences for the native diversity. This result calls for further experiments that explore and enhance the diversity of plant species reintroduced in open biome restoration, reducing the dissimilarity between restored sites and reference ecosystems.

Overcoming species propagation barriers, aiming at diversifying the restoration species pool of herbaceous species would be vital to promote the reintroduction of a more diverse array of plant species in the restoration of open ecosystems (Vidal *et al.*, 2020; Faria *et al.*, 2025). Despite the predominant use of native species, focus is still given to woody species in many places. The reintroduction of forbs along with grasses to restore biodiverse and more resilient open ecosystems should be encouraged. Poorly planned tree plantations pose a threat to local floras, resulting in plantings that are floristically more similar to one another than to the regional species pool, even in forest restoration (Almeida *et al.*, 2025). Moreover, the woody species focus undermines the key roles the herbs and forbs play in ecosystem functioning (Siebert *et al.*, 2024). The species choice should respect the vegetation structure of the target open biome, which means that the herb:woody species ratio should ideally match that of the native reference ecosystem.

## Supplementary Material

The following online material is available for this article:

**List S1.** List of the scientific papers included in the systematic review of seeded and planted species on tropical open ecosystems

**Figure S1.** Mean number of plant species planted in open biome restoration experiments, depending on their growth form: herbs (grass, forbs, graminoids) or woody plant species. (ANOVA; n = 29 experiments; p>0.05; F = 0.0706, mean ± SE).

**Figure S2.** Mean number of species planted in open biome restoration experiments, depending on their origin: native or exotic plant species. (n = 29 experiments; p<0.001; z value = 15.048, mean ± SE). GLM (formula = number_plants ~ origin, family = "poisson")

**Table S1.** List of the 220 plant species planted in Tropical Grassland Biome restoration experiments. Ecosystems: G = grassland, Sh = shrubland, Sa = savanna; IUCN status: DD = data deficient, EN = endangered, LC = Least Concern, NT = neat threatened, VU = vulnerable.

## Acknowledgments

We thank the comments by the editor and two anonymous reviewers significantly increased the manuscript quality.

## References

Abrahão A, Costa PB, Lambers H *et al.* 2019. Soil types select for plants with matching nutrient-acquisition and - use traits in hyperdiverse and severely nutrient-impoverished campos rupestres in Central Brazil. Journal of Ecology 107: 1302-1316. doi: 10.1111/1365-2745.13111

Almeida C, Reid JL, Lima RAF, Pinto LFG, Viani RAG. 2025. High-diversity Atlantic Forest restoration plantings fail to represent local floras. Perspectives in Ecology and Conservatio 23: 6-11. doi: 10.1016/j.peco.2024.12.001

Barbosa-Dias LG, Silveira FAO, de Marco Júnior P, Padilha DL. 2025. Open ecosystems restoration: a global review shows biases and mismatches between theory and practice. Restoration Ecology 33: e14307. doi: 10.1111/rec.14307

Bond WJ. 2019. Open Ecosystems: ecology and evolution beyond the forest edge. Oxford, Oxford Academic.

Bond WJ, Midgley JJ. 2001. Ecology of sprouting in woody plants: the persistence niche. Trends in Ecology & Evolution 16: 45-51. 10.1016/S0169-5347(00)02033-4

Buisson E, Archibald S, Fidelis A, Suding KN. 2022. Ancient grasslands guide ambitious goals in grassland restoration. Science 377: 594-598. doi: 10.1126/science.abo4605

Buisson E, Le Stradic S, Silveira FAO *et al.* 2019. Resilience and restoration of tropical and subtropical grasslands, savannas, and grassy woodlands. Biological Reviews 94: 590-609. doi: 10.1111/brv.12470

Cuenca G, de Andrade Z, Escalante G. 1998. Diversity of glomalean spores from natural disturbed and revegetated communities growing on nutrient-poor tropical soils. Soil Biology and Biochemistry 30: 711-719. doi: 10.1016/S0038-0717(97)00191-0

D'Antonio C, Meyerson LA. 2002. Exotic plant species as problems and solutions in ecological restoration: a synthesis. Restoration Ecology 10: 703-713. doi: 10.1046/j.1526-100X.2002.01051.x

de Castilho Silva AW, Pontara V, Bueno ML, Villa PM, Walter BMT, Meira-Neto JAA. 2024. The bulk of a plant hotspot: composition, species richness and conservation status of the Cerrado herbaceous-subshrub flora. Folia Geobotanica 59: 39-49. doi: 10.1007/s12224-024-09451-y

Faria FS, Ordóñez-Parra CA, Granata L, Resende LV, Silveira FAO. 2025. Low-cost technology supports propagation of endemic species from a global biodiversity hotspot. Restoration Ecology 33: e70005. doi: 10.1111/rec.70005

Flora e Funga do Brasil. 2024. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/. 16 Dec. 2024

Haddad TM, Pilon NAL, Durigan G, Viani RAG. 2021. Restoration of the Brazilian savanna after pine silviculture: pine clearcutting is effective but not enough. Forest Ecology and Management 491: 119158. doi: 10.1016/j.foreco.2021.119158

Holl KD, Luong JC, Brancalion PHS. 2022. Overcoming biotic homogenization in ecological restoration. Trends in Ecology & Evolution 37: 777-788. 10.1016/j.tree.2022.05.002

IUCN. The IUCN Red List of Threatened Species. Version 2024-2. https://www.iucnredlist.org. 16 Dec. 2024

<figure><img src="image_11.png" /></figure>
Ladouceur E, Mayfield MM. 2017. The early response of subtropical tussock grasslands to restoration treatments. Restoration Ecology 25: 689-695. doi: 10.1111/rec.12491

Lira-Martins D, Xavier RO, Mazzochini GG et al. 2024. Soil acidification controls invasive plant species in the restoration of degraded Cerrado grasslands. Restoration Ecology 33: e14294. doi: 10.1111/rec.14294

Lovera M, Cuenca G. 1996. Arbuscular mycorrhizal infection in Cyperaceae and Gramineae from natural, disturbed and restored savannas in La Gran Sabana, Venezuela. Mycorrhiza, 6: 111-118. doi: 10.1007/s005720050115

Lyons KG, Török P, Hermann J et al. 2023. Challenges and opportunities for grassland restoration: a global perspective of best practices in the era of climate change. Global Ecology and Conservation 46: e02612. doi: 10.1016/j.gecco.2023.e02612

Mazzochini GG, Lira-Martins D, de Barros FV et al. 2024. Effects of grass functional diversity on invasion success by exotic grasses in Cerrado grasslands. Journal of Applied Ecology 61: 271-280. doi: 10.1111/1365-2664.14561

Medeiros NF, Ordóñez-Parra CA, Buisson E, Silveira FAO. 2024. Systematic review of field research reveals critical shortfalls for restoration of tropical grassy biomes. Journal of Applied Ecology 61: 1174-1186. doi: 10.1111/1365-2664.14640

Medeiros N, Buisson E, Silveira FAO. 2025. Replication Data for 'What are we planting in ecological restoration of tropical open ecosystems? A systematic review of controlled experiments'. https://doi.org/10.48331/SCIELODATA.YJE17N SciELO Data, V1.

Murphy BP, Andersen AN, Parr CL. 2016. The underestimated biodiversity of tropical grassy biomes. Philosophical Transactions of the Royal Society B 371: 20150319. doi: 10.1098/rstb.2015.0319

Nerlekar AN, Veldman JW. 2020. High plant diversity and slow assembly of old-growth grasslands. The Proceedings of the National Academy of Sciences 117: 18550-18556. doi: 10.1073/pnas.1922266117

Nerlekar AN, Sullivan LL, Brudvig LA. 2024. Grassland restorations must better foster forbs to facilitate high diversity. Restoration Ecology 32: e14214. doi: 10.1111/rec.14214

Pilon NAL, Campos BH, Durigan G et al. 2023. Challenges and directions for open ecosystems biodiversity restoration: an overview of the techniques applied for Cerrado. Journal of Applied Ecology, 60: 849-858. doi: 10.1111/1365-2664.14368

Pimenta MA, Fonseca A. 2021. To what extent are threatened plant species considered in impact assessment decision-making? Insights from southeastern Brazil. Environmental Impact Assessment Review 86: 106516. doi: 10.1016/j.eiar.2020.106516

POWO. Plants of the World Online. Facilitated by the Royal Botanic Gardens, Kew. https://powo.science.kew.org/. 16 Dec. 2024

R Core Team. 2025. R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria. https://www.R-project.org/.

Ribeiro JF, Walter BMT. 2008. As principais fitofisionomias do bioma Cerrado. In: Sano SM, Almeida SP, Ribeiro JF (eds.) Cerrado: ecologia e flora. Brasília, Embrapa Cerrados. p.152-212

Siebert F, te Beest M, Fynn R et al. 2024. Past, present and future of forbs in old-growth tropical and subtropical grasslands. Annual Review of Ecology, Evolution and Systematics 55: 395-421. doi: 10.1146/annurev-ecolsys-102722-022331

Silveira FAO, Ordóñez-Parra C, Moura LC et al. 2022. Biome Awareness Disparity is BAD for tropical ecosystem conservation and restoration. Journal of Applied Ecology 59: 1967-1975. doi: 10.1111/1365-2664.14060

Simões LHP, Guillemot J, Ronquim CC et al. 2024. Green deserts, but not always: A global synthesis of native woody species regeneration under tropical tree monocultures. Global Change Biology 30: e17269. doi: 10.1111/gcb.17269

Veldman JW, Overbeck GE, Negreiros D et al. 2015. Where tree planting and forest expansion are bad for biodiversity and ecosystem services. BioScience 65: 1011-1018. doi: 10.1093/biosci/biv118

Vidal CY, Naves RP, Viani RAG, Rodrigues RR. 2020. Assessment of the nursery species pool for restoring landscapes in southeastern Brazil. Restoration Ecology 28: 427-434. doi: 10.1111/rec.13096

Volis S. 2019. Conservation-oriented restoration – a two for one method to restore both threatened species and their habitats. Plant Diversity 41: 50-58. doi: 10.1016/j.pld.2019.01.002

Wieczorkowski JD, Lehmann CER. 2022. Encroachment diminishes herbaceous plant diversity in grassy ecosystems worldwide. Global Change Biology, 28: 5532-5546. doi: 10.1111/gcb.16300

Wiederhecker A, Ferreira MC, Rodrigues SB et al. 2024. Ten Years of directing seeding restoration in the Brazilian savanna: lessons learned and the way forward. Journal of Environmental Management 365: 121576. doi: 10.1016/j.jenvman.2024.121576

**Authors' Contributions:**

N.F. Medeiros and F.A.O. Silveira conceived the ideas, designed the methodology and led the writing of the manuscript; N.F. Medeiros collected and analysed the data. All authors contributed critically to the drafts and writing, and gave final approval for publication.

**Conflict of Interest:**

The authors have no conflict of interest to declare.

**Funding Information:**

We thank CNPq and FAPEMIG for providing scholarships and grants that support this research.

**Data availability:**

The datasets and the code used in the current study are available at the GitHub repository: git@github.com:nataliafmedeiros/planting_seeding_tgb.git and at SciELO Data: Medeiros N, Buisson E, Silveira FAO. 2025. Replication Data for 'What are we planting in ecological restoration of tropical open ecosystems? A systematic review of controlled experiments'. Available at: https://doi.org/10.48331/SCIELODATA.YJE17N, SciELO Data, V1.