Forest Ecology and Management 553 (2024) 121628

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Forest Ecology and Management

journal homepage: www.elsevier.com/locate/foreco

Restoration plantings in the Atlantic Forest use a small, biased, and

homogeneous set of tree species
Crislaine de Almeida a, *, J. Leighton Reid b, Renato A. Ferreira de Lima c,
Luis Fernando Guedes Pinto d, Ricardo Augusto Gorne Viani e
a
Programa de Pós-graduação em Recursos Florestais, Universidade de São Paulo, Escola Superior de Agricultura Luiz de Queiroz, Av. Pádua Dias, 11, Piracicaba, SP
13418-900, Brazil
b
School of Plant and Environmental Sciences, Virginia Tech, 185 Ag Quad Lane, Blacksburg, VA 24061, USA
c
Departamento de Ciências Biológicas, Escola Superior de Agricultura Luiz de Queiroz, Universidade de São Paulo, Av. Pádua Dias, 11, Piracicaba, SP 13418-900,
Brazil
d
Fundação SOS Mata Atlântica, Av. Paulista 2073 - Horsa 1 - Conj. 1318, São Paulo, SP 01311-300, Brazil
e
Centro de Ciências Agrárias, Universidade Federal de São Carlos, Rodovia Anhanguera, km 174, Araras, SP 13600-970, Brazil

A R T I C L E I N F O A B S T R A C T

Keywords: Investigating which species are planted in active restoration initiatives is useful for identifying underrepresented
Biodiversity conservation species and functional groups, which in turn can inform and improve the practice and policies of forest resto­
Biotic homogenization ration. The Atlantic Forest is one of the global hotspots for biodiversity conservation and a region where many
Forest restoration
forest restoration initiatives have been carried out in recent decades. We conducted a taxonomic and functional
Functional groups
Landscape restoration
analysis of a database with 1073 forest restoration plantings implemented from 2002 to 2018 in the Brazilian
Nurseries Atlantic Forest. We assessed how well restoration plantings represent the dispersal syndromes, successional
groups, and the ratio of nitrogen-fixing, and threatened species, based on a comparison with 268 Atlantic Forest
remnants. Comparisons were made for the entire biome as well as three subtypes: Atlantic Rainforest, Araucaria
Forest and Seasonal Forest. We found 423 tree species planted in the restoration efforts, which represent less than
8 % of the Atlantic Forest tree flora. In comparison with the forest remnants, restoration plantings have a greater
proportion of nitrogen-fixing tree species (17 % in plantings vs. 9 % in remnants) but underrepresent non-pioneer
(67 % vs. 85 %), animal-dispersed (56 % vs. 74 %), and threatened species (18 species found in plantings vs. 83
in remnants). Our results indicate a bias towards N-fixing, non-animal-dispersed, and pioneer species in resto­
ration plantings and towards using the same set of species across different regions of the Atlantic Forest. We
recommend actions to increase the representation of the native tree flora in Atlantic Forest restoration plantings,
especially concerning inclusion of animal-dispersed and threatened species and groups of species that are less
likely to colonize restoration sites, such as large-seeded animal-dispersed species.

1. Introduction consequences for global biodiversity and society.

Society is currently making unprecedented investments in ecological
restoration to address the interrelated global challenges of climate
change, mass extinction, and desertification (UNFCCC, 2020). Dozens of
nations have committed to restore hundreds of millions of hectares over
the current decade. There are many ambitious forest restoration pro­
grams established across the world (Haase and Davis, 2017; Aronson
et al., 2020), and we have just started the United Nations Decade on
Ecosystem Restoration (Abhilash, 2021; Ceccon et al., 2020). How this
restoration is implemented could have far-reaching, long-term
Planting native tree seedlings is the most used forest restoration
technique in many tropical regions (Lamb, 2011; Rodrigues et al., 2011;
Holl, 2012; Palma and Laurance, 2015). After planting, some species
may become absent from a restoration site because of the early death of
their seedlings while many other new species may recruit via seeds
arriving from the surrounding landscape (Ashton et al., 2014; Dimson
and Gillespie, 2020; Suganuma and Durigan, 2021). Thus, the long-term
tree species composition in a restoration site frequently differs from that
on the day of planting. Nonetheless, knowing the set of planted species is
important, not only for the impacts they have on subsequent forest

* Corresponding author.
E-mail address: crislainealmeid@gmail.com (C. de Almeida).

https://doi.org/10.1016/j.foreco.2023.121628
Received 2 September 2023; Received in revised form 29 November 2023; Accepted 30 November 2023
Available online 14 December 2023
0378-1127/© 2023 Elsevier B.V. All rights reserved.
C. de Almeida et al.
recovery, but also because these may represent most of tree species
present in later years, even two or three decades after, particularly at
sites with limited natural regeneration capacity (Murcia, 1997; Beltrán
et al., 2022). Compiling and evaluating the pool of species used in
restoration plantings in a region or biome is useful to identify missing
and underrepresented species and functional groups in restoration ef­
forts (Holl et al., 2017; Brancalion et al., 2018). Some groups, such as
vascular epiphytes and slow-growing tree species dispersed by gravity or
large mammals recruit very slowly in forest restoration plantings (Reid
et al., 2016; Holl et al., 2020; Suganuma and Durigan, 2021). These
groups may be confined to remaining forests if not introduced by active
restoration initiatives (Martínez-Garza and Howe, 2003). Thus, the
knowledge on the species planted for restoration is crucial to guide the
creation and improvement of public policies aimed to stimulate seed
collecting, seedling production, and the inclusion of missing species and
functional groups that fail to recruit in restoration plantings.
In recent decades, many restoration initiatives have been carried out
in the highly diverse Atlantic Forest (Rodrigues et al., 2009; Shaw, 2019;
de Siqueira et al., 2021). The Atlantic Forest is a hotspot for biological
conservation, with high plant endemism (Fiaschi and Pirani, 2009; de
Lima et al., 2020b), threatened by losses of species and carbon stocks
(Joly et al., 2014; de Lima et al., 2020a). Thus, our objective was to carry
out a descriptive and critical analysis of the tree species planted in the
southern part of the Brazilian Atlantic Forest in the last two decades.
Brancalion et al. (2018) showed that animal dispersed tree species with
larger seeds are underrepresented in Atlantic Forest plantings, compared
to what is observed in remnant forests. To advance this topic and pro­
vide new insights, we made taxonomic considerations on what is planted
and evaluated results within regional forest types rather than only
generalizing to the whole Atlantic Forest. In addition, we evaluated
whether nitrogen fixing and pioneer tree species are under or over­
represented in forest restoration plantings compared to remnant Atlantic
Forest fragments. Animal-dispersed (Viani et al., 2015; Camargo et al.,
2020; de Almeida and Viani, 2021), early-successional (Martínez-Garza
et al., 2013; de Almeida and Viani, 2019) and nitrogen-fixing (Nichols
et al., 2001; Siddique et al., 2008) species play key roles for accelerating
tropical forest restoration processes. Finally, we also investigated the
presence of threatened and endemic tree species in the pool of species
planted as well as of the species that combine higher-than-average po­
tential for carbon storage, ecological interactions, and biodiversity
conservation (de Lima et al., 2020a), because carbon storage and tree
species conservation are both expected outcomes of forest restoration
plantings (Brancalion et al., 2018; Di Sacco et al., 2021).
Tree seedlings for forest restoration purposes are produced by
nurseries, which frequently limit the pool of species available for forest
restoration at regional scales (Jalonen et al., 2018; Ladouceur et al.,
2018; Vidal et al., 2020). The restoration species pool (i.e. the set of
species available for restoration) is mainly composed of the most
regionally abundant species, while rare and threatened species, whose
seeds are more difficult to obtain, tend to be less available or, even,
completely absent in nurseries (Hoffmann et al., 2015; Vidal et al.,
2020). Thus, we hypothesized that non-pioneer, and animal-dispersed
species in general are underrepresented in restoration plantings,
because pioneer and abiotic-dispersed species are more easily found in
the edges of the degraded forest fragments used for seed collection and
have a higher seed production (Martínez-Garza and Howe, 2003; Santos
et al., 2008). Furthermore, we expected that the abundance and fre­
quency of threatened and endemic species are lower in restoration
plantings than in remaining forest fragments since endemic species have
restricted spatial distribution (Mori et al., 2018) while threatened ones
are usually rare and/or commonly not found in the edges of degraded
forest fragments.”
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Forest Ecology and Management 553 (2024) 121628
2. Material and methods
2.1. Study site
The Atlantic Forest originally covered an area of about 1.5 million
km2, in Argentina, Paraguay and mostly in Brazil, in a wide range of
environmental conditions (Ribeiro et al., 2009). Variation in tempera­
ture, rainfall regime and in latitude and altitude drive differences in its
species composition (Oliveira-Filho and Fontes, 2000), and conse­
quently the formation of different Atlantic Forest subtypes. We consid­
ered three forest types: Atlantic Rainforest, Seasonal Forest, and
Araucaria Forest (Fig. 1, Fig. S1).
The Atlantic Rainforest is on the coast of Brazil, and is characterized
by highlands with altitude ranging from 0 to 2800 m. The climate is
humid subtropical without a dry season, with an average annual pre­
cipitation greater than 2200 mm and an average annual mean temper­
ature around 17 ◦ C (Scudeller et al., 2001; Joly et al., 2012). The
Seasonal Forest is on the hinterland highlands in the center and south­
eastern interior of Brazil (Oliveira-Filho and Fontes, 2000). This vege­
tation type is under a seasonal climate with a dry season in the winter,
from April to September (Morellato et al., 2000; Oliveira-Filho and
Fontes, 2000). Average annual rainfall is 1000–1500 mm while average
annual mean temperature is 20–26 ◦ C (Behling, 2002). Finally, the
Araucaria Forest is dominated by the coniferous Araucaria angustifolia
(Marques et al., 2004), and it is found in the highlands of Southern Brazil
and in the south of the Serra da Mantiqueira. The climate is temperate
and humid, without dry periods, with an average annual precipitation
between 1400 and 2200 mm (Hueck, 1953). The average annual mean
temperature is 12–18 ◦ C, and winter temperatures approach 0 ◦ C
(Nimer, 1989; Oliveira-Filho, 2009). We used the Brazilian official de­
limitation of the Atlantic Forest regions (IBGE, 2004) to place each
planting in a given forest type. By this classification, our study included
958 plantings in the Seasonal Forest, 81 in the Atlantic Rainforest and 34
in the Araucaria Forest (Fig. 1).
2.2. Databases
We combined databases of the “Click Árvore” and “Florestas do
Futuro” restoration programs, coordinated by the NGO SOS Mata
Atlântica (https://www.sosma.org.br) in Brazil. These databases
contain lists of trees planted from 2002 to 2018, in 1073 Atlantic Forest
restoration sites (Fig. 1, Table S1). Seedlings for these plantings came
from 29 nurseries. Although SOS Mata Atlântica encourages the use of
high biodiverse, the participating nurseries had free choice for seed
collection and seedling production. The database includes the number of
trees by species planted in each restoration site, and as such it does not
contain information about planted trees that established or non-planted
species that recolonized sites naturally.
We also used data stored in the TreeCo database (de Lima et al.,
2015, 2020b), that compiles information on species abundances and
functional traits in eastern South America using scientific publications,
including monographs, dissertations, theses, and national and interna­
tional scientific articles. From the TreeCo database, we selected 268
forest remnants described as belonging to the Atlantic Forest Biome,
from 140 scientific publications (Fig. 1) which had 1) inventories
including trees with diameter at breast height >5 cm, 2) no evidence of
disturbance and, 3) available information on the inventoried forest type.
We included in our analyses a subset containing 85 Atlantic Rainforest
sites, 27 Araucaria Forest sites, and 156 Seasonal Forest fragments sites,
totaling a sampling effort of 207 ha and 286,660 trees (Fig. 1, Table S2).
2.3. Species information
First, we checked the spelling and synonyms for all the species names
occurring in the plantings and remnants lists, using the Flora do Brasil,
2020 database (Flora do Brasil, 2020). Then, we classified all tree
C. de Almeida et al.
Fig. 1. Official limits for the Atlantic Forest subtypes in Brazil, and the location of the studied restoration plantings and remnant forests. Despite some plantings and
remnants lying outside of the overall limits shown in the map, all of them are Atlantic Forest patches.
species according to their seed dispersal syndrome, successional group,
and nitrogen fixation status. For the dispersal syndrome classification,
species were defined as animal-dispersed or non-animal dispersed
(anemochoric or autochoric) based on our recognition of their fruit
types. We classified species as pioneer or non-pioneer (early and late
secondary species) considering previous information of the Treeco
database and our knowledge. Overall, pioneer trees are small-seeded,
fast-growing and short-lived trees that germinate on forest gaps while
non-pioneer species are large-seeded, long-lived and slow-growing trees
that germinate under the forest shade (Swaine and Whitmore, 1988).
Species were classified as nitrogen-fixing or non-nitrogen-fixing using a
global nitrogen status database (Tedersoo et al., 2018). Species listed in
Tedersoo ( et al. (2018) as Rhizobia, likely Rhizobia, and Frankia were
considered nitrogen fixing, while species listed as unlikely Rhizobia were
considered as non-nitrogen-fixing.
Species were classified according to their extinction risk as vulner­
able, endangered, critically endangered, or not endangered based on the
Brazilian Official List of Threatened Plant Species (MMA (2000)).
Knowing whether threatened species are being planted in restoration
initiatives is useful to infer the role these plantings play for tree species
conservation. Additionally, we classified species planted in forest
restoration following the results of three recently published articles that
categorized Atlantic Forest species 1) by their capacity to colonize forest
restoration areas (Suganuma and Durigan, 2021, restricted to a few
species they evaluated), 2) as endemic (de Lima et al., 2020b), and 3) by
combing a higher-than-average potential for carbon storage, ecological
interactions, and biodiversity conservation (de Lima et al., 2020a).
2.4. Data analysis
We evaluated the number of species shared between restoration
plantings and remnants. We calculated absolute frequency and relative
abundance of each species in restoration plantings and in forest rem­
nants. We also calculated the percentage of trees and individuals by
dispersion syndrome, successional group, nitrogen-fixing capacity, and
threat of extinction. Then, to infer if some groups are over or under­
represented in restoration plantings, we compared the proportions of
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Forest Ecology and Management 553 (2024) 121628
each category within a given functional group between plantings and
remnants, for species and individuals, using Chi-square tests. All ana­
lyses were performed for all forest types combined and for each forest
type. Additionally, we created a list with all species, from both plantings
and remnants. Then, we used a Chi-square test to compare the propor­
tion of animal versus non-animal dispersed species within the groups of
pioneers and non-pioneer species, using the whole set of species, to
know whether the prevalence of one of these successional groups in
plantings may affect the representativeness of species and trees of a
given dispersal syndrome in the restoration initiatives. Specifically, we
would like to test whether an overall preference for pioneer species
planting could lead to an underrepresentation of animal-dispersed tree
species. Finally, we calculated the percentage of higher-than-average
potential for carbon storage, ecological interactions, and biodiversity
conservation (de Lima et al., 2020a), and of endemic species (de Lima
et al., 2020b) found in the restoration plantings.
All analyses were performed in R version 3.5.1. (R Development Core
Team, 2018) and interpreted following the language of evidence (Muff
et al., 2021), where values <0.001 represent very strong evidence, 0.001 –
0.01 represent strong evidence, 0.01 – 0.05 represent moderate evidence,
0.05 – 0.1 represent weak evidence and 0.1 – 1 represent little or no evi­
dence. We applied the Bonferroni correction to all Chi-square tests.
3. Results
3.1. Taxonomic diversity of restoration plantings and remnants
A total of 423 species belonging to 66 families were found in the
database of Atlantic Forest restoration plantings, totaling about 21.7
million seedlings (Table S3 and S4). Overall, the five most frequent
species in Atlantic Forest plantings were, in descending order: Schinus
terebinthifolia (present in 94 % of the plantings), Ceiba speciosa (93 %),
Cedrela fissilis (88 %), Croton floribundus (86 %), and Guazuma ulmifolia
(85 %). The three most-abundant species were S. terebinthifolia,
C. speciosa, and Parapiptadenia rigida, which together made up 7 % of all
seedlings. The most-planted species overall were also the most-planted
ones in the Seasonal Semideciduous Forest, but differed slightly in the
C. de Almeida et al. Forest Ecology and Management 553 (2024) 121628

Atlantic Rainforest and in the Araucaria Forest (Fig. 2; Table S3).

We found 1891 tree species in the selected remnants, with 1327 in
the Seasonal Forest, 1309 in the Atlantic Rainforest and 409 in the
Araucaria Forest. The most abundant and frequent species in the rem­
nants, overall and within forest subtypes, were generally different from
those in restoration plantings (Table S5). Of the 20 most abundant
species in the remnants, 15 were non-pioneer and dispersed by animals
and only one, Croton floribundus, was also within the 20 most abundant
species in restoration plantings. The richest families in the plantings
were Fabaceae, with 104 species (25 %), and Myrtaceae, with 36 species
(9 %; Fig. S2; Table S4). Those families were also the richest families in
the remnants: Myrtaceae with 273 species (14 %) and Fabaceae with
223 species (12 %; Fig. S2; Table S6).
There were 93 threatened species (vulnerable or endangered) in
remnants’ species list, and 18 in plantings’ species list. For the restora­
tion plantings, threatened species represented 4 % of the species and 4 %
of the seedlings. Cedrela fissilis was the most-abundant threatened spe­
cies, and it was the only threatened species among the top-20 most-
planted species overall and in the top-five in the Atlantic Rainforest. Fig. 3. Threatened species in Brazilian Atlantic Forest restoration plantings and
Only three threatened species, C. fissilis, Psidium myrtoides, and Zeyheria percentage of plantings in which each threatened species was found. The
tuberculosa were present in > 50 % of the plantings (Fig. 3). However, 99 number in front of the bar of each species is its relative abundance ( %) in the
% of the plantings (1055) had at least one threatened tree, with the total amount of seedlings planted in the databased we consulted.
number of threatened species in each planting varying from 0 to 12
(Table S7). We also found two exotic species in the restoration plantings: proportion of nitrogen-fixing tree species (15 % vs. 8 %) but under­
Psidium guajava, with 128,177 seedlings, found in 444 (41 %) sites and represent in around 20 % points non-pioneer (67 % vs. 85 %), and
Tecoma stans, present in five (0.5 %) plantings and totaling 651 in­ animal-dispersed (50 % vs. 71 %) species. When the comparison is made
dividuals (Table S1). considering the number of trees, overall, the proportion of animal
Species overlap among forest subtypes was higher for plantings than dispersed trees drops 21 % points in plantings compared to remnants (50
for remnants (Fig. 4A). A total of 360 species were shared between vs. 71 %), while proportion of pioneer (33 vs. 12 %) and nitrogen-fixing
remnants and plantings, representing 19 % of the species in the rem­ trees (19 vs. 9 %) were more than the double in restoration plantings.
nants and 85 % in the plantings. No more than 20 % of the species found The proportions of animal dispersed species (Fig. 5A) and individuals
in remnants in our dataset were planted in Atlantic Forest restoration, (all p-values < 0.0001; Table S8) were lower in restoration plantings
overall (16 %) and for all forest subtypes individually (Fig. 4B). than in remnants, overall and for each forest type. Although the pro­
portion of pioneer species seemed similar between restoration plantings
3.1.1. Functional groups in restoration plantings and reference forests and remnants (Fig. 5b), the former had proportionally more pioneer
Overall, restoration plantings have almost the double of the

Fig. 2. Ten most-abundant species planted in 1073 forest restoration plantings (left, blue) and found in 238 selected Atlantic Forest remnants (right, green), overall
(A, B) and for Atlantic Rainforest (C, D), Araucaria Forest (E, F) and Seasonal Forest (G, H). The percentage is the sum of relative abundance for the 10 most
abundant species.

4
C. de Almeida et al.
Fig. 4. Tree species sharing among Atlantic Forest A) subtypes for restoration plantings and for remnants and B) overall and for forest type between restoration
plantings and remnants. Percentages in B are the percentage of species found in the other condition: for example, for all type of forests, 89 % of the planted species are
shared with remants while 19 % of the species in remnants are shared with plantings.
trees, overall and for each forest type (all p-values < 0.0001; Table S8).
The proportion of nitrogen-fixing trees was greater in restoration
planting than remnants in all comparisons (p-value < 0.0001; Table S8),
while the proportion of nitrogen-fixing species was greater in restoration
plantings overall and for the Seasonal Semideciduous Forest (Fig. 5C).
Overall, the most-planted tree species were pioneer and non-animal
dispersed. In the top 10 most planted species, there were 10 pioneer
and seven non-animal dispersed species. There was a relation between
dispersal syndrome and successional group for the species in the datasets
(χ2 = 34; p < 0.001; strong evidence), with the non-animal dispersed
species being more frequent among the group of pioneer species.
3.1.2. Threatened and underrepresented species
The proportion of threatened species did not differ between resto­
ration plantings and remnants (Fig. 5D), but forest remnants had more
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Forest Ecology and Management 553 (2024) 121628
trees of threatened species than restoration plantings (overall 6,4 × 4,4
%; Table S8). In addition, 81 % of the threatened species found in the
remnants were absent in the restoration plantings. Seventy five percent
of the threatened species in the remnants (70 out of 93) were dispersed
by animals, while in plantings only 28 % (five out of 18) were dispersed
by animals.
Plantings incorporated only 16 of the 42 species indicated by Suga­
numa and Durigan (2021) as species that do not colonize restoration
plantings. Only one of these non-colonizer species planted was threat­
ened - Apuleia leiocarpa (vulnerable).
Only 33 % (81 out of 242) of the top higher-than-average potential
for carbon storage, ecological interactions, and biodiversity conserva­
tion species listed by de Lima et al. (2020a) were found in restoration
plantings (Table S3). These 81 species represented 22 % of all planted
seedlings, but only 3 % of planted trees were threatened and
C. de Almeida et al.
Fig. 5. Proportions of species within functional groups in Atlantic Forest restoration plantings (R) and remnants (F). AR = Atlantic Rainforest, AF = Araucaria Forest,
SF = Seasonal Forest. * ** indicate difference between comparisons with a p-value < 0.0001 and * indicate difference between comparisons with a p-value < 0.05,
both based on the Chi-square test.
recommended. Just 73 of the 1547 (4,7 %) species belonging to the
group of Atlantic Forest pure endemic species defined by de Lima et al.
(2020b) were found in the list of species planted in restoration, and these
73 endemic species represented 13 % of all planted trees.
4. Discussion
We analyzed the largest database on the species used in restoration
plantings that we have knowledge of so far, for a region where forest
restoration has advanced through intense research, policies, and large-
scale restoration initiatives (Crouzeilles et al., 2019). We found, in our
dataset, that 22 % of the tree species surveyed in remnant Atlantic Forest
fragments were actively introduced in forest restoration plantings.
Native tree species representation was even lower, only 8 %, when
compared to the species richness of the whole Atlantic Forest, which
encompasses 5044 tree species (de Lima et al., 2020b). Additionally, we
found that there was a greater overlap of the species planted for resto­
ration than for the species found in remnants when we compared the
species across different Atlantic Forest subtypes. This means that, in the
set of plantings we investigated, there is a tendency of planting the same
tree species in restoration efforts, without regard to local flora compo­
sition. Plantings can be the main strategy to reestablish forest cover in
severe degraded landscapes (Rodrigues et al., 2011), but if we plant
always the same species and some of them persist in the plantings over
time, then we favor biotic homogenization, i.e., an increase in the
taxonomic similarity of two or more biotas over a time interval – an
effect that has the consequence of reducing biodiversity (Olden and
Rooney, 2006; Lôbo et al., 2011; Palma and Laurance, 2015; Holl et al.,
2022).
The underrepresentation of native flora in restoration plantings re­
flects the underrepresentation found in forest nurseries (Vidal et al.,
2020), since forest nurseries filter the species pool available for resto­
ration (Ladouceur et al., 2018). Whether taxonomic diversity of native
species in individual restoration plantings is really needed, is still a
recurrent question (Castro et al., 2021; Guerin et al., 2021). Recently,
the discussion has moved more to functional than taxonomic represen­
tation, because of the impacts that some functional groups have on
initiating tropical forest restoration processes and catalyzing divergent
successional pathways in the understory (Ashton et al., 2014; Carlucci
et al., 2020). In this regard, we found that animal-dispersed species and
late successional trees are underrepresented in forest restoration
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Forest Ecology and Management 553 (2024) 121628
plantings, despite their relevance to restoration processes and biodi­
versity recovery (Reid et al., 2015). Animal-dispersed species play an
important role attracting seed-dispersers (Martínez-Garza et al., 2013),
which therefore increases restoration success (Viani et al., 2015). Our
results corroborate those of Brancalion et al. (2018) and add that this
pattern is widespread across different Atlantic Forest subtypes. This is an
alarming observation, since many of the large seed-dispersers have
become locally extinct in recent centuries, and some native
animal-dispersed tree species may not be able to colonize regenerating
forests if they are not actively introduced (da Silva and Tabarelli, 2000;
Guimarães Jr.. et al., 2008). Moreover, restoration plantings in Atlantic
Forest are not using most species that have already been found unable to
colonize forest plantings (Suganuma and Durigan, 2021). Thus,
increasing availability of animal-dispersed species and trees in restora­
tion plantings is needed, especially focused on those group of species
that will be unlikely to colonize restoration sites on their own. A higher
diversity in restoration implementation may not necessarily result in
higher taxonomic diversity (Guerin et al., 2021), but, if one of the goals
of restoring forests is to achieve taxonomic diversity and composition
conservation, better identifying the species that are incapable of colo­
nizing restoration sites on their own is an important step towards veri­
fying whether they are underrepresented in restoration plantings and
whether we need to favor their active reintroduction (Reid, 2015). If
reintroduction is required, it must be done in a carefully planned and
science-based manner, aiming to guarantee these species, which may be
sensitive to restoration conditions, will succeed over time and reproduce
in restoration sites (Godefroid et al., 2011).
We found a greater proportion of nitrogen-fixing trees in plantings
than in native Atlantic Forest remnants. Even though this is probably the
result of greater availability of Fabaceae seedlings in forest nurseries
(Vidal et al., 2020), the planting of nitrogen-fixing trees can improve soil
quality in areas undergoing restoration, accelerating tree growth and
canopy closure (Nichols et al., 2001; Siddique et al., 2008; Chaer et al.,
2011; Aleixo et al., 2020). Additionally, restoration plantings overall
had a greater representation of pioneer species and individual trees than
forest remnants, as previously found for other regions (Engert et al.,
2020). Pioneer species grow faster and are better competitors in
degraded sites dominated by invasive grasses than late successional
species are, therefore, they accelerate canopy cover formation (de
Almeida and Viani, 2019), reduce restoration costs and tend to be
preferred in restoration plantings. However, depending on landscape
C. de Almeida et al.
fragmentation and on restoration site limitations, the arrival of some
species from other successional stages may not occur in restoration
plantings, making plantings with only pioneer species impoverished and
degraded over time (de Souza and Batista, 2004; Martínez-Garza and
Howe, 2003). This issue could be balanced if more species and in­
dividuals of late successional stages naturally arrive at restoration sites
over time (Rother et al., 2019). Alternatively, late-successional species
could be introduced through enrichment plantings (Bertacchi et al.,
2016), a strategy still costly and not sufficiently studied (Mangueira
et al., 2019) to avoid the collapse of restoration sites when the
short-living pioneer trees die. Additionally, since animal-dispersed
species were less common for the group of pioneers than for
non-pioneer species, prioritizing pioneer species in forest restoration
plantings may account for a lower representation of animal-dispersed
species in plantings in comparison to forest remnants. Thus, a recom­
mendation is selecting and prioritizing animal-dispersed pioneer spe­
cies, such as suggested by the framework species method and other
approaches (Elliot et al., 2013; Viani et al., 2015, de Almeida & Viani,
2021).
Only 19 % of the threatened species (18 of 93) observed in the forest
remnants were included in our dataset of species used in restoration
plantings. This means that most threatened species are not planted, and,
when planted, are usually introduced in low abundances and are often
the same species throughout regions. Thus, Atlantic Forest restoration
plantings poorly contribute to the reintroduction of threatened species,
which may disappear over time if specific reintroduction programs are
not implemented (Tabarelli et al., 2005; Luber et al., 2016). This pattern
is partly caused by the difficulty in collecting seeds for many of the
threatened species (Hoffmann et al., 2015). In addition, most endemic
species are absent in restoration plantings, demonstrating the difficulty
of accessing and to introducing these species, which often dominate
remnant forest patches (Caiafa and Martins, 2010; Villa et al., 2019). If
one of the goals of ecosystem restoration is directly contributing to
species conservation, more attention should be given, through oriented
public programs and policies to encourage seeding collection (Goodale
et al., 2023) and inclusion of species with high-conservation value in
forest restoration plantings (de Lima et al., 2020a). However, to be
effective, these programs and policies should be carefully planned and
implemented to assure that those species will survive in the harsh
microclimate of restoration sites and that collection of their seeds for
seedling production will not threaten remaining native populations.
We conclude that despite recent advances and the emergence of
many technologies and initiatives for forest restoration, the number of
species used in Atlantic Forest restoration plantings still represents a
small subset of its tree flora. An important finding is that, at least in the
subset of plantings concentrated in Atlantic Forest southern portion,
planted species tend to be the same, independent of the variation in
species composition of natural forest subtypes. In addition, animal-
dispersed, and high-conservation value species are underrepresented
in restoration plantings. The diversity and composition of planted spe­
cies may be less important when most regional tree species can recolo­
nize a site naturally, however this capacity is doubtful for many rare,
threatened, and animal-dispersed species and at restoration sites iso­
lated from remnant seed sources. Thus, a step forward is investigating
their capacity to recruit forest restoration sites over time, in different
landscapes scenarios. Finally, we recommend that restoration initiatives
better incorporate regional variation of forest composition into their
reference models and target species lists. Although some authors have
recommended improving regional variance in restoration plantings
(Barbosa et al., 2017), our results indicate that there is still a long way to
go before Atlantic Forest restorations will truly represent the diversity
and complexity of this unique biome.
CRediT authorship contribution statement
Crislaine de Almeida: Conceptualization, Methodology, Formal
7
Forest Ecology and Management 553 (2024) 121628
analysis, Investigation, Writing - Original Draft, Writing - Review &
Editing. J. Leighton Reid: Formal analysis, Investigation, Writing -
Original Draft, Writing - Review & Editing. Renato Augusto Ferreira
de Lima: Data Curation, Writing - Original Draft, Writing - Review &
Editing. Luis Fernando Guedes Pinto: Data Curation, Writing - Review
& Editing. Ricardo Augusto Gorne Viani: Conceptualization, Meth­
odology, Formal analysis, Investigation, Writing - Original Draft,
Writing - Review & Editing; Supervision.
Declaration of Competing Interest
The authors declare the following financial interests/personal re­
lationships which may be considered as potential competing interests.
Crislaine de Almeida reports financial support was provided by Coor­
denação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil
(Capes) - Finance Code 001. Renato Augusto Ferreira de Lima reports
financial support was provided by Fundação de Amparo à Pesquisa do
Estado de São Paulo.
Data Availability
Data will be made available on request.
Acknowledgments
This study was financed in part by the Coordenação de Aperfeiçoa­
mento de Pessoal de Nível Superior - Brasil (Capes) - Finance Code 001.
We are thankful to the SOS Mata Atlântica nongovernmental organiza­
tion for aiding this research and to Virginia Tech for hosting Crislaine de
Almeida. Ricardo A. G. Viani is thankful to the grant #2022/13398-1,
São Paulo Research Foundation (FAPESP). The compilation of the spe­
cies abundances and information stored in TreeCo was funded by the
grant #2013/08722-5, São Paulo Research Foundation (FAPESP), but
many other agencies (e.g. e.g. Capes, CNPq, FAPEMIG, FAPERJ,
FAPESC, etc.) funded the hundreds of researchers that produced the
forest remnants data used in this study.
Appendix A. Supporting information
Supplementary data associated with this article can be found in the
online version at doi:10.1016/j.foreco.2023.121628.
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