Journal for Nature Conservation 65 (2022) 126115

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Journal for Nature Conservation

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Review

How are biodiversity and carbon stock recovered during tropical forest
restoration? Supporting the ecological paradigms and political
context involved
Carolina M. da Rosa *, Marcia C.M. Marques
Laboratório de Ecologia Vegetal, Departamento de Botânica, SCB, Universidade Federal do Paraná, Caixa Postal 19031, 81581-980 Curitiba, Paraná, Brazil

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

Keywords: To meet agendas for biodiversity conservation and mitigation of climate change, large-scale restoration initia­
Carbon stock tives propose ecological restoration as an alternative that can reconcile these two objectives. In ongoing
Diversity ecosystem restoration, increased diversity is always associated with increased productivity (and consequent
Ecosystem service
carbon stock), which is among the most important ecosystem functions. The ecological paradigm of this asso­
Productivity
ciation is that ecosystem biodiversity (B) is positively related to both ecosystem functions and services (EF and
Tropical forest recovery
ES). However, BEF and BES relationships vary spatially and temporally, which makes understanding these re­
lationships relevant and important for practical restoration actions. In this study, we asked how biodiversity and
carbon stock recovery occurs during tropical forest restoration. We reviewed literature of the relationships be­
tween BEF and BES in the context of ecological restoration and asked whether ecological restoration can recover
both. In addition, we conducted a metadata analysis of studies on the recovery of biodiversity and biomass in
regenerating tropical forests (n = 83) to find the best model that describes this relationship. In general, studies
showed that ecosystem biodiversity and productivity are positively related, and that restoration can recover
both. We found an asymptotic and positive correlation between biodiversity and biomass in tropical forests,
suggesting limitation of the mutual gains of these two ecosystem properties during restoration. We discuss these
results in the context of ecological theory and the practice of ecological restoration.

1. Introduction basing criteria on ecological, economic and social contexts (SER, 2004).
The main purpose of ecological restoration is to recover biodiversity and
The current trend of surpassing planetary limits for biodiversity loss ecological functions, separately and together, (Lamb, Erskine & Par­
and climate change (Steffen et al., 2015) has stimulated the use of large- rotta, 2005; Wright et al., 2009) of degraded ecosystems. Thus, biodi­
scale ecological restoration as a mitigation tool (Valikchali, Pourma­ versity and ecosystem functions are usually considered together in
jidian & Darvishi, 2014). For instance, the current global agreements for restoration projects, even though the putative relationship between the
biodiversity conservation, climate change mitigation (Paris Agreement) two is often neglected. In the seminal study by Bradshaw (1984; Ap­
and world sustainability (Sustainable Development Goals) view resto­ pendix 1, Fig. A1), restoration is conceived of an action that results in
ration as a tool with which to achieve those proposed goals (Leadley the gradual increase in ecosystem structure (species and complexity)
et al., 2014; UN, 2015; UNFCCC, 2015). Increasingly, ecological resto­ and function (biomass and nutrient content), and which are linearly
ration is receiving attention and will be among the main focal points of related. Thus, an increase in the number of species (or functional
global environmental agendas with the UN Decade on Ecosystem groups) is followed by, or associated with, a concomitant increase in
Restoration (UN, 2020). Consequently, understanding how (and ecosystem functions. Yet, the exact relationship between ecosystem
whether) ecological restoration is an efficient method for achieving all structure (biodiversity) and ecosystem function (productivity, decom­
global goals is the subject of intense scientific debate. position, nutrient cycling) and how it operates across different restora­
The first step for ecological restoration planning is to define goals, tion strategies and scales remains uncertain. The hypothesis has been

Abbreviation: BEF, Biodiversity-Ecosystem Function.

* Corresponding author.
E-mail address: cmrosa92@gmail.com (C.M. Rosa).

https://doi.org/10.1016/j.jnc.2021.126115
Received 5 May 2021; Received in revised form 4 December 2021; Accepted 6 December 2021
Available online 21 December 2021
1617-1381/© 2021 Elsevier GmbH. All rights reserved.
C.M. Rosa and M.C.M. Marques Journal for Nature Conservation 65 (2022) 126115

proposed that more species are required for constant ecosystem func­ restored to halt biodiversity loss (UN, 2015).
tions, and even more for ecosystem stability (Loreau et al., 2001), and With these agreements in mind, restoration is conceived of as being
some models suggest that the ecosystem function can become stabilized the best (or only) way to halt, and reverse, the extreme rates of biodi­
at a given species diversity level (Naeem, 1998). Theoretically, there­ versity loss and global warming (Bullock et al., 2011). As a consequence,
fore, ecosystem processes - and the ecosystem services they generate other initiatives established pacts for large-scale restoration, such as the
(Costanza et al., 1997; de Groot et al., 2010) - are dependent on biodi­ Bonn Challenge, which is a global effort to restore 350 million hectares
versity (Hooper, Legendre, & Condit, 2005), and so restoration is pro­ by 2030 (IUCN, 2011a), and the 20 × 20 Initiative, of Latin American
posed as a means by which both can be recovered or reestablished and Caribbean countries to restore 20 million hectares by 2020 (WRI,
(Bullock, Aronson, Newton, Pywell & Rey-Benayas, 2011). Nonetheless, 2017b), and the African Forest Landscape Restoration Initiative
we do not yet know the true efficiency of large-scale restoration to (AFR100), to restore 100 million hectares by 2030 (WRI, 2017a), and
mitigate the current climate and biodiversity crises (Bustamante et al., finally, Brazil’s Atlantic Forest Restoration Pact, to recover 15 million
2019). Therefore, the fundamental issue behind restoration objectives is hectares by 2050 (Rodrigues, Brancalion & Isernhagen, 2009).
that we must come to understand how biodiversity and ecosystem Following these initiatives, additional approaches have been rolled out,
functions and services are related. such as the Forest Landscape Restoration initiative, whose goal is to
Here, we discuss the ecological paradigms of the theories of recover ecosystem functionality and improve human well-being (IUCN,
biodiversity-ecosystem functioning (BEF) and biodiversity-ecosystem 2011b), and the New York Declaration on Forests (UN, 2014), which
service (BES), in the context of ecological restoration. We reviewed aims to decrease forest losses, to restore forests and croplands, and to
literature of the relationships between ecosystem biodiversity and reduce greenhouse gases by 4.5–8.8 billion tons by 2030 (UN, 2014).
functionality in the context of ecological restoration, to determine Thus, ecological restoration, for its many mitigating effects, is a
whether ecological restoration is able to recover both diversity and worldwide concern and has been prioritized by most countries. For these
function. In addition, we conducted a metadata analysis of studies on the reasons, the UN declared 2021–2030 as the Decade of Ecosystem
recovery of biodiversity and biomass in regenerating tropical forests to Restoration, aiming to halt degradation of ecosystems, and to restore
find the best model that describes that relationship. These results pro­ them to their former states, to achieve these global goals. Following the
vide important insight for the international debates of possible diver­ proposal with 70 signatory countries, this UN resolution is building a
gence in the agendas that focus either on biodiversity conservation or on strong, broad-based global movement to increase restoration actions
climate change mitigation (Epple et al., 2016). and to put the world on track for a globally sustainable future. In sum­
mary, ecological restoration has never before achieved such worldwide
2. Global political context recognition, and which underscores that we need to better understand
how gains in biodiversity and ecosystem functionality are related during
The most important recent global agreements on sustainability restoration.
explicitly indicate that restoration is the path to reach their goals
(Chazdon et al., 2016). Aichi’s Targets, established at the 10th COP of 3. Biodiversity and productivity relationships (BP): Revisiting
Convention on Biological Diversity (CBD) in 2010, had 193 signatory ecosystem functions (BEF) and services (BES) theories
countries, and targets were intended to reduce biodiversity losses
through 2020. Specifically, targets 14 and 15 explicitly state that eco­ To understand how restoring ecosystem can reestablish original
systems providing essential services should be restored and safeguarded, structure and function, we must return to key points in the evolution of
and that the contribution of biodiversity to carbon stocks must be BEF and BES theories (see methods for literature review in Appendix 2).
enhanced, thereby contributing to climate change mitigation. It was The relationship between biodiversity and ecosystem function has long
proposed that these actions should recover at least 15 per cent of global been a central issue that intrigued ecologists (Loreau et al., 2001), for
degraded ecosystems (Leadley et al., 2014). Thus, it was clearly recog­ which it was included as one of the 100 fundamental ecological ques­
nized that conservation alone is unable to provide for the maintenance tions that are extremely relevant for the future of ecology (Sutherland
of biodiversity and that restoration is also essential (Dobson, Bradshaw et al., 2013). It is generally agreed that biodiversity may influence both
& Baker, 1997; Moreno-Mateos, Power, Comín & Yockteng, 2012). ecosystem functions (productivity, soil stability and nutrient cycling)
Therefore, while understanding the important ecosystem effects of and ecosystem services (soil fertility, provisioning of plant products,
ecological restoration is still developing, the recognition that restoration erosion control, invasion resistance, pest and pathogen regulation and
is fundamental, and subsequent actions had important and significant water supply), and so it follows that biodiversity loss can drastically and
roles in mitigating threats to biodiversity (Aronson & Alexander, 2013). negatively affect ecosystem processes (Hector et al., 2007; Naeem et al.,
Based on those observations, the 11th COP Convention on Biological 1999; Quijas et al., 2010). Considering ecological restoration, the BEF
Diversity (CBD) in 2012 in Hyderabad emphasized the importance of approach allows us to evaluate restoration in the context of ecosystem
restoring degraded lands to achieve the Aichi Targets (CBD, 2012). function, and respond to several knowledge gaps in restoration ecology
While those targets have not yet been fully achieved, biodiversity con­ (Aerts & Honnay, 2011). Three possible BEF relationships are proposed:
servation goals post-2020 related to Climate Changes have been 1) a linear function, 2) an asymptotic curve, and 3) no effect (inde­
considered by the CBD. The Paris Agreement signed by 175 countries in pendence) of biodiversity on ecosystem function (Vitousek & Hooper
the 21th COP of United Nations Framework Convention on Climate 1993; Appendix 1, Fig. A2). Thus, a large part of the discussion about
Change (UNFCCC) aims to strengthen the global response to the threat of BEF involves finding the pattern of this relationship.
climate change, to keep the increase in global temperature below 2◦ C, to Early, and heated, debate about BEF concerned the biodiversity-
reduce greenhouse gas emissions, and to increase the capacity for productivity relationship (BP), in the 1960s. At that time, productivity
adaptation to climate changes (UNFCCC, 2015). To reach those goals, in was thought to be the cause of diversity (Preston, 1962), and a unimodal
addition to other actions, both restoration and adequate management of pulse (diversity increasing to its maximum, then decreasing) was the
forests (as carbon sinks to increase carbon storage) are necessary default position (Abramsky & Rosenzweig, 1984; Fox, 1985; Tilman,
(Griscom et al., 2017). Also, the Sustainable Development Goals (SDG), 1983), even under stressful conditions (Xiao & Chen, 2019). However,
launched in 2015 by the United Nations with 193 signatory member robust studies found this pattern in only 30% of the studies across
states, established goals to combat climate change and biodiversity loss. terrestrial and aquatic ecosystems (Waide et al., 1999), as well as finding
SDG 13 affirmed that increases in reforestation of degraded and positive (Abrams, 1988; Cusens et al., 2012) and negative linear (Owen,
degrading landscapes are necessary to combat global warming, and SDG 1988; Wang et al., 2001) relationships, as well as independence
15 emphasized that terrestrial ecosystems should be protected and (Cermeño et al., 2013; Wang et al., 1999). Thus, ecosystem patterns did

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C.M. Rosa and M.C.M. Marques Journal for Nature Conservation 65 (2022) 126115

not conform to expectations due to this wide variability (Mittelbach because diversity alone may not recover ecosystem functions in per­
et al., 2001). Subsequently, in the 1990s, diversity driving productivity turbed ecosystems (Parain et al., 2019).
began to be debated and tested (Moorthi et al., 2008; Naeem et al., 1994;
Schläpfer & Schmid, 1999; Striebel et al., 2012), leading to the current 4. Restoration and biodiversity recovery
consensus that both directions of the BEF are interrelated, but context-
dependent (Cardinale et al., 2009; Gross and Cardinale, 2007). At that In general, biodiversity loss has five main causes: 1) habitat degra­
time, several ecological drivers of BP were identified, including type of dation (including fragmentation), 2) biological invasion, 3) over­
ecosystem function (Maureaud et al., 2020), herbivore density (Declerck exploitation, 4) pollution, and 5) diseases (McGill et al., 2015; Vitousek
et al., 2007; Hillebrand & Lehmpfuhl, 2011; Yee & Juliano, 2007), et al., 1997; Wilcove et al., 1998). Restoration, in these situations, can
community assembly rules (Steiner & Leibold, 2004), trophic level in­ resolve (partially or completely) losses, but in different ways (Halme
teractions (Rakowski & Cardinale, 2016; Thébault & Loreau, 2006), et al., 2013; Jordan III, 1997; Young, 2000). For example, ecological
spatial (Braschler et al., 2004; Chase & Leibold, 2002; Dodson, Arnott, & restoration creates new habitat (Jordan III et al., 1988), establishes el­
Cottingham, 2000; Li et al., 2019) and temporal (Dodson et al., 2000) ements of the landscape that improves connectivity (Tambosi &
scales, environmental (Costa et al., 2016) and regional processes (Bis­ Metzger, 2013), and protects fragments from edge effects (Brancalion
choff et al., 2005). Also, a negative BEF relationship was noted in et al., 2013), all of which contribute to biodiversity recovery in degraded
invaded communities (Wu et al., 2018), demonstrating that invasions and fragmented habitats. Restoration strategies based on planting native
can change these relationships. Thus, abiotic and biotic conditions can species may increase resilience and resistance to invasive species (Wil­
modify the shape of the BEF relationship (Guerrero-Ramírez & Eisen­ son, 2013), thereby decreasing biodiversity losses due to biological in­
hauer, 2017). vasion (Simberloff & Vitule, 2014). Restoration projects that support
Soon, a mechanistic approach to BP was proposed. Highly- economic development and creation of sustainable livelihoods protect
productive areas can have greater species richness (Preston, 1962) biodiversity, because unemployment is often associated with forest
because the increase in available energy (due to productivity) allows degradation and overexploitation (Cao et al., 2017). Finally, restoration
more species in the food web (Connell & Orias, 1964; MacArthur, 1955). of degraded lands can reduce the effects of air and water pollution and
This mechanism explains the effects of productivity on diversity. increase biodiversity (Lee et al., 2007; Wong, 2003). Additionally,
Alternatively, the selection effect (i.e., the greater chance of more pro­ restoration projects can make ecosystems more diverse and resistant to
ductive species existing in more diverse environments), and niche disease (Carnus et al., 2006). Thus, restoration is likely to often be a very
complementarity (i.e., species with different functions occupying effective method to mitigate the impacts of human activity on both
different niches) explained the effects of diversity on productivity biodiversity (Qin et al., 2016), and to recover degraded ecosystems
(Omidipour et al., 2020; Tilman, 1999). (Srivastava & Giri, 2020).
BP began to be more relevant when anthropogenic drivers were While restoration is expected to limit the effects of the main causes of
included while examining ecosystem function (Liang et al., 2016). Not biodiversity loss, species numbers can increase throughout the restora­
only ecosystem functions, but also ecosystem services were shown to be tion process (Brown & Lugo, 1990; Guariguata & Ostertag, 2001). Evi­
affected by biodiversity. The relationship between biodiversity and dences indicate that plant species richness (Liebsch et al., 2007;
ecosystem services tends to be positive (MEA, 2005), while also scale- Saldarriaga et al., 1988; Tabarelli & Mantovani, 1999), along with
dependent (Chisholm et al., 2013; Luo et al., 2019) and varies with phylogenetic (Qin et al., 2016) and functional (Cadotte et al., 2011;
climate, soil and elevation (Di Marco et al., 2018). This is because Marcilio-Silva et al., 2016; Purschke et al., 2013) diversity all increase
biodiversity plays different roles in its influence on ecosystem services, during restoration. This is equally true for animals, including insects
sometimes acting as regulator of some ecosystem processes, but also as a (Piper et al., 2009), amphibians (Brodman et al., 2006; Hilje & Aide,
service in itself (Mace et al., 2012). In short, strong evidence indicates 2012), reptiles (East et al., 1995), birds (Catterall et al., 2012), and
that biodiversity positively influences ecosystem functions and services mammals (Kalies et al., 2012).
(Balvanera et al., 2006; Rita & Borghetti, 2019; Tilman et al., 2012). Evidence abounds for increased biodiversity with ecological resto­
Consequently, we can deduce that diversity must also positively influ­ ration. In both aquatic and terrestrial ecosystems world-wide, restora­
ence biomass and carbon storage (Díaz et al., 2009). After perturbation, tion has been shown to increase biodiversity by 44% (Benayas et al.
therefore, biomass is more conserved in more diverse communities than 2009) and by 43% in Chinese terrestrial ecosystems (Huang et al., 2019)
in less diverse communities (Tilman, 1996). Thus, more diverse com­ over that of the degraded ecosystems that were restored. Rates of
munities are more stable and resistant to disturbance (Gross et al., biodiversity recovery are often even greater in tropical terrestrial eco­
2014). systems (250%, Benayas et al., 2009) or agrosystems (68%), with re­
Despite the many studies of BP in lakes, wetlands and grasslands, BP covery from 54% in vertebrates to 79% in invertebrates (Barral et al.,
has been neglected in forests (Ojha & Dimov, 2017; Vilà et al., 2007). 2015). Recovery of diversity in forests varies from 15 to 84% (Crou­
Yet, evidence of positive BP in forests include Mediterranean scle­ zeilles et al., 2017, 2016), and in tropical forests recovery is around 53%
rophylous (Vilà et al., 2007), temperate and mixed (Lei et al., 2009; (Shimamoto et al., 2018). Rates of biodiversity recovery vary over life
Paquette & Messier, 2011), and tropical forests (Häger & Avalos, 2017; forms and type of ecosystem. For example, restoration increases inver­
Lasky et al., 2014; Poorter et al., 2015). That positive relationship was tebrate biodiversity by 108% in Chinese forests (Ren et al., 2017), while
found with different measures of biodiversity, including species rich­ aquatic invertebrate biodiversity in wetlands increased by 15% (Meli
ness, diversity indices, and functional and phylogenetic diversities et al., 2014).
(Henry et al., 2010; Ojha & Dimov, 2017; Potter & Woodall, 2014; Clearly, it is important to recognize that restoration influences
Zhang et al., 2011). With the few studies of BP in forests, the trend is for biodiversity recovery, but that recovery depends on the kind of chal­
positive and asymptotic relationships (Liang et al., 2016). During forest lenge posed by the type of degradation, the restoration strategy under­
restoration, the positive BP trend is pervasive (Bu et al., 2014; Salisbury taken, time that the ecosystem has suffered degradation, the type of
& Potvin, 2015), even though it depends on the successional context landscape, taxonomic groups, and climate where degradation has
(Lasky et al., 2014). However, the greater evidence for positive BP occurred (Crouzeilles et al., 2016; Huang et al., 2019; Ren et al., 2017).
trends should be treated cautiously as this consensus may be caused by Although restoration can be a very positive tool for recovery, conser­
publication bias if (quite possibly) studies demonstrating the expected vation workers must also be realistic and recognize that the conditions in
result are more easily published (Braga et al., 2017). Thus, the rela­ which degradation occurred will determine the magnitude of its effec­
tionship requires additional study and decisions based on those results tiveness (Hobbs, 2013).
should be carefully considered. We should also include climate change

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C.M. Rosa and M.C.M. Marques Journal for Nature Conservation 65 (2022) 126115

5. Restoration and recovery of ecosystem functions and services future (Chokkalingam & de Jong, 2001; Holl, 2017, 2002). To examine
BEF relationships and how biodiversity and carbon stock are recovered
Many ecosystem functions can be recovered during the restoration during tropical forest restoration, we reviewed studies of ongoing
process, including nutrient and biogeochemical cycles (Amazonas et al., ecological restoration in tropical forests. From these studies we extrac­
2011; Macedo et al., 2008), water regulation (Simões et al., 2002; ted two ecological indicators (species richness and basal area) and
Stromberg, 2001), carbon sequestration (Philipson et al. 2020; Shima­ modeled their relationships. We used species richness because it is the
moto et al., 2014), decomposition (Smith & Chadwick, 2014) and most clear and typical measure of biodiversity (Purvis & Hector, 2000).
pollination (Forup, Henson, Craze, & Memmott, 2008; Williams, 2011). Similarly, basal area is strongly correlated with aboveground biomass,
Typically, restoration recovers ca 25% of ES of degraded lands and therefore, a good indicator of the amount of carbon fixed from the
world-wide (Benayas et al., 2009). Recovery rates vary among ecosys­ atmosphere (Brown & Lugo, 1990). We searched the Web of Science,
tems, types of ecosystem function and services, and restoration strate­ Science Direct and Scopus through December 2020 (with no other date
gies. For example, in agrosystems, restoration increased ES by 120% limit). Search terms were: “tropical forest“, “restoration” or “regenera­
(Barral et al., 2015), and in passive restoration strategies, increased by tion“ or “recovery” or “succession“, “basal area”, “species richness“, and
319% (Ren et al., 2017) over the degraded conditions. Restoration of “plant” or “vegetation“. Details of the literature review and analyses are
wetlands increased delivery of provisioning, regulating and supporting in Supplementary Material (Appendix 3).
ecosystem services by 36% (Meli et al., 2014) and regulating services for In 83 plots in tropical regions (Appendix 3, Table A1), comprising 44
tropical forests by 52% (Shimamoto et al., 2018). While variable, it is studies in 21 countries (Fig. 2), we found that the relationship between
clear that restoration can improve both ecosystem functions and species richness and basal area was best described by a Michaelis-
services. Menten model (Sr = 0.219; AIC = 8.0474; Table A2), followed by
Productivity and carbon sequestration are among the most important linear and quadratic models (Appendix 3, Table A2). The Michaelis-
ecosystem functions and services that tend to be recovered by restora­ Menten model has two parts: the first (t = 2.42; p = 0.018) indicates
tion in terrestrial ecosystems (Aide, 2000; Dixon et al., 1994; Parrotta initial growth, and the second (t = 1.68; p = 0.096) indicates asymptotic
et al., 2012). Productivity is measured as biomass, where carbon typi­ stabilization. When compared to the linear model (based on AIC), there
cally comprises 50% of the total mass that plants accumulate (He et al., was a significant result (p < 0.001). That happens because there are
2005; Lacerda et al., 2009). Thus, the atmospheric excess of carbon may more data at the part one when compared to part two. We observe a
be sequestered and accumulated in ecosystems during restoration (Cao tendency to stabilization at second part. We expected that the second
& Woodward, 1998; Jones & Donnelly, 2004; Lal, 2004; McGuire et al., part should contain older forests, but the oldest secondary forest in our
2001; Newell & Stavins, 2000). This is especially true during early model is 70 years. Nevertheless, considering the biological approach,
successional stages of restoration, when fast growing species are more the Michaelis-Menten model is more meaningful and represents better
abundant and carbon sequestration proceeds more rapidly (Montagnini this relationship between carbon and diversity in ecological restoration.
& Porras, 1998; Shimamoto et al. 2014; Sierra et al., 2012). Effectiveness of restoration depends on the setting of the region that
The main effect of carbon sequestration in ecosystems is that CO2 was perturbed, as we previously noted, and demonstrated here in this
decreases in the atmosphere (Alves et al., 1997; Sierra et al., 2012), analysis. The original studies were in regions with different kinds and
thereby mitigating climate changes (Holl & Zahawi, 2014; Locatelli magnitudes of perturbation, including logging, mining, agriculture
et al., 2015; van der Sande et al., 2017). Clearly restoration can be an (including domesticated animals), cyclones and fire. Restoration type
important strategy that responds efficiently to the alterations in regional also varied, and included natural regeneration, active restoration, and
and global climate (Harris et al., 2006), and function as a carbon offset agroforestry, all of which influence BEF relationships (Gonzalez et al.,
for many years (Silver et al., 2000). Thus, because restoration of 2020). Disturbance regimes and restoration methods, all important in­
degraded lands is associated with greater carbon uptake and storage, it fluences of diversity and carbon storage in tropical forests (Cardinale,
may be a key global strategy to mitigate global climatic change Nelson & Palmer, 2000; Lindner & Sattler, 2011; Zanini et al., 2021),
(Houghton et al., 1993). and this variety contributes to model fit.
The best-fit M-M model is positive and asymptotic and relates basal
6. Restoring biodiversity and biomass in tropical forests: A case area to tree diversity, and so productivity is associated with biodiversity
study

BP relationships in ecological restoration are especially important for

tropical forests and so must be understood. Tropical forests are the most
biologically diverse ecosystems world-wide (Brown, 2014; Holl, 2002)
and for years have been suffering from deforestation (Mittermeier et al.,
2004). Tropical forests provide many ecosystem services, including
carbon sequestration (Lugo & Brown, 1992; Sierra et al., 2012), water
purification (Ellison et al., 2012), and soil-fertility maintenance (Ditt
et al., 2010), among others. Tropical forests tend to be carbon sinks
(Lugo & Brown, 1992; Ngo et al., 2013), that sequester huge amounts of
carbon, storing around 45% of terrestrial carbon, 11% of soil carbon
pool, as well as contributing 50% of global net primary productivity
(Bonan, 2008; Brown & Lugo, 1982; Chazdon et al., 2016; van der Sande
et al., 2017). Specifically, forests of tropical America have the capacity
to sequester 46% of total carbon in tropics, while tropical Asian forests
can sequester 34%, followed by tropical Africa at 20% (Brown, 1996).
Thus, in addition to the extreme biodiversity found in tropical regions
(Brown, 2014), the very great rates of primary productivity provide
equally great potential to sequester or conserve large quantities of car­
bon (Gillman et al., 2015).
Tropical forests, because of their importance and rapid deforestation, Fig. 1. Michaelis-Menten model: relationship between relative species richness
are (or should be) the main focus for ecological restoration in the near and relative basal area in tropical forests ongoing restoration (n = 83 plots).

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C.M. Rosa and M.C.M. Marques Journal for Nature Conservation 65 (2022) 126115

Fig. 2. Map showing locations around the world of the 44 tropical forest studies (21 countries) used in the case study.

during restoration in tropical forests (Fig. 1). The type II model (pro­ Acknowledgements
posed by Vitousek & Hooper 1993) is similar, in which saturation in
ecosystem function occurs (Appendix 1, Fig. A2). Asymptotic carbon We thank Jerônimo Sansevero and Jean Vitule for their suggestions.
storage is likely to be due to the way in which niche complementarity We are grateful to Regiane S. da Cunha and Elivane S. Capellesso for
and selection influences BEF (Tilman, 1999). their help in preparing the figures. We thank Larissa M. F. Manfrinato,
Thus, as new species with complementary functions are added dur­ Camila M. da Rosa and James R. Roper for language revision.
ing restoration, carbon storage will continue to increase until it reaches a
maximum value when species function redundancies no longer add Funding
productivity in the forest being restored. Asymptotic relationships in
BEF and BES during restoration also occur in other ecosystem functions Support was provided by the Brazilian Education Council (CAPES)
and services, including regulation of water flow, provision of clean with the fellowship to CMR (Grant Code 001), and by the Brazilian
water, and maintenance of soil fertility (Benayas et al., 2009; Hector & Research Council (CNPq) with the grant to MCMM (Grants 202897/
Bagchi, 2007; Hobbs, 1992; Naeem, 2006; Wright et al., 2009). 2016-3; 303356/2019-7).

7. Conclusion Appendix A. Supplementary material

In this study we discuss the relationships between biodiversity, Supplementary data to this article can be found online at https://doi.
ecosystem functions and services in the context of ecological restoration. org/10.1016/j.jnc.2021.126115.
Ecological theories that underlie BEF and BES relationships were from
studies in different systems, seldom from studies of tropical forests. The References
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