Ecological Indicators 109 (2020) 105779

Contents lists available at ScienceDirect

Ecological Indicators
journal homepage: www.elsevier.com/locate/ecolind

Rainfall interception and plant community in young forest restorations T

a,b,⁎ b a,1
Fernando Ravanini Gardon , Renato Miazaki de Toledo , Bruno Melo Brentan ,
Rozely Ferreira dos Santosb
a
Department of Water Resources, School of Civil Engineering, Architecture and Urban Design, State University of Campinas, Rua Saturnino de Brito, 224 Cidade
Universitária “Zeferino Vaz”, Campinas, São Paulo CEP: 13083-889, Brazil
b
Department of Ecology, Institute of Biosciences, University of São Paulo, Rua do Matão, Travessa 14, 321, Cidade Universitária, São Paulo CEP: 05508-900, Brazil

A R T I C LE I N FO A B S T R A C T

Keywords: The conversion of tropical forests to human land-use threatens biodiversity conservation and the delivery of
Forest restoration many ecosystem services, especially water-related ecosystem services. In these landscapes, many investments
Plant community have been made to restore native forests and recover hydrological processes lost by deforestation. Rainfall in-
Rainfall interception terception is a key hydrological process for water-related ecosystem services’ maintenance, which plays an
Ecosystem services
important role in runoff, infiltration, erosion control, and flood regulation. We evaluated rainfall interception
Monitoring
over a 1-year period in eight restoration sites within the Brazilian Atlantic Forest. We used the interception
Tropical forest
function as an indicator of water dynamics recovery in actively restored forests. Monthly rainfall interception
was measured by 80 interceptometers distributed within the sites (10/site), and 24 pluviometers were installed
in open fields (3/site) close to the restoration sites to collect total precipitation (P) incidents over the sites. We
also measured plant community attributes involved in the interception process (density, basal area, tree species
richness, and the ratio of deciduous plants). The average rainfall interception reached 21.4 ± 3.9%, but a
significant variability was observed among sites. Results showed that 65% of the monthly interception collected
is below 30 mm.month−1. Basal area and species richness were forest attributes positively correlated to each
other and the most important in the interception process. The results show that actively restored forests can
reestablish rainfall interception rates similar to those of mature tropical forests in the short term (10 years). In
addition, more time or complementary interventions are needed for plant communities to reach expected at-
tributes’ values. Self-organized maps analysis showed a negative relationship between interception and the
proportion of deciduous plant individuals. We present information to support land-use policy decisions, as the
results revealed insights regarding the effects on the water cycle that may result from increasing forest cover. We
argue that restoring ecosystem services should be the main goal of restoration programs and determining if
hydrological processes are being effectively recovered by restoration actions is crucial for achieving water
sustainability.

1. Introduction
The conversion of tropical forests to human land-use leads to the
degradation of these natural ecosystems, threatening biodiversity,
ecological integrity, landscapes sustainability, and ecosystem services
maintenance (Cardinale et al., 2012; MEA, 2005; Naeem et al., 2012;
Wu, 2013).
Studies have shown that in tropical regions, the water cycle and the
consequent delivery of water-related ecosystem services (WES) such as
water flows regulation and water quality for human well-being, are
closely related to forest extension, structure, composition, and
ecological functions and processes (Ferraz et al., 2014; Hackbart et al.,
2017; Tambosi et al., 2015). In these landscapes with heavy pre-
cipitation, where deforestation and soil degradation are commonly
observed, precipitation reaches the unprotected soil directly and a small
portion infiltrates, but most of it flows quickly to the rivers. This dy-
namic contributes to the erosive processes, flood events, and alterations
in the physicochemical properties of freshwater (Defries and Eshleman,
2004; Tambosi et al., 2015). In addition, continued deforestation in the
tropics will increase climate change through reducing evapotranspira-
tion and precipitation rates, and elevating global temperature
(Lawrence and Vandecar, 2015).

⁎
Corresponding author.
E-mail address: fernandogardon@hotmail.com (F.R. Gardon).
1
Centre de Recherche en Controle et Automatique de Nancy, Université de Lorraine, 2 rue Jean Lamour, Vandoeuvres-les-Nancy 54500, France.

https://doi.org/10.1016/j.ecolind.2019.105779
Received 6 June 2018; Received in revised form 24 June 2019; Accepted 27 September 2019
1470-160X/ © 2019 Elsevier Ltd. All rights reserved.
F.R. Gardon, et al.
In this scenario, forests can mitigate the effects of water scarcity and
global warming (Ellison et al., 2017). In tropical landscapes, restoration
has been encouraged as a path to recovery of hydrological processes
and a strategic tool to ensure water sustainability and safeguard WES
(De Groot et al., 2010; Keeler et al., 2012; Seifert- Dähnn et al., 2015).
However, evaluations of hydrological indicators and benefits are still
scarce (Amatya et al., 2016) and it is difficult to predict WES recovery.
In this study, we aimed to evaluate the recovery of the rainfall in-
terception function by restored forests as an indicator of hydrological
functioning as it is first hydrological process in the reestablishment of a
forest ecosystem’s water cycle (Loescher et al., 2002; Pike and Scherer,
2003). Some authors describe it as a key process in the water cycle,
which ensures water redistribution and contributes to regulating WES
through soil protection and air humidity increase (Arcova et al., 2003;
Geißler et al., 2013; Goebes et al., 2015; Zimmermann et al., 2013).
Interception rates depend on the structure and composition of forests
(Crockford and Richardson, 2000); therefore, we also selected possible
indicators of rainfall interception based on plant community attributes.
We expect to offer useful information on evaluations of restoration
success thereby contributing to decisions related to forest and water
management, which are key components to WES maintenance in
human-modified landscapes.
2. Methodology
2.1. Study area
Our study was conducted in the Brazilian Atlantic Forest, a tropical
biodiversity hotspot (Myers et al., 2000), with only 12–16% of its ori-
ginal cover remaining (Ribeiro et al., 2009; SOS Mata Atlântica, 2017).
We selected eight actively restored riparian forests, located in the
eastern part of São Paulo state, Brazil (Fig. A.1; Table A.1), totaling
10 ha of land under restoration. The restorations were implemented by
a program for large-scale forest restoration – Projeto de Recuperação de
Matas Ciliares (PRMC, 2005) that aimed to restore forests and recover
ecosystems services in relevant regions to safeguard the water supply of
São Paulo state, with a population of about 46 million people (IBGE,
2010) which has recently faced a water crisis (Coutinho et al., 2015).
The plantations were started in 2007 with a seedling planting density of
1666 individuals per hectare and at least 80 native species, as suggested
by the law (SMA – 8/2008).
The original vegetation of the region is Seasonal Semideciduous
Forest – SSF (PRMC, 2005). Sites are characterized by a granite-gneiss
lithology and comprise two mountain-ranges with crystalline parent
material. Restoration sizes ranged from 0.3 to 4.4 ha, with elevation
from 640 to 930 m above sea level, and annual precipitation from
1231.4 to 1662.1 mm with an average variation of 198.8 ± 20 mm
between the wettest and driest months DAEE, 2011). The rainy season
occurs from October to March. Early agricultural use of these lands
began during the 19th century (Dean, 1997), while cattle grazing
continues to be practiced on high slopes and under heavy precipitation
(Machado et al., 2014). Previous land-use at the sites included the
cultivation of exotic grasses for grazing (Brachiaria sp., Panicum max-
imum, and Melinis sp.).
2.2. Study design
2.2.1. Rainfall interception
During a 1-year period, we measured monthly rainfall interception
(Imm) in restored forests using 80 interceptometers distributed below
the forest canopy (10/site) to collect the water that goes through the
forest structure and would reach the ground (throughfall – Thmm). We
used 24 pluviometers in open fields (3/site) close to the restoration sites
to collect incidents of total precipitation (Pmm). Interceptometers were
distributed in two plots (10 × 10 m) randomly allocated along four
transects (4 × 50 m) previously established to obtain forest attributes
2
Ecological Indicators 109 (2020) 105779
(Section 2.2.2) in each site. Each plot was subdivided into a 2 × 2 m
grid resulting in 20 possible positions of which five were randomly
selected as interceptometer positions (see Section 2 of the
Supplementary material for detailed information). Data were collected
monthly for a year. The amount of water stored in each collector was
quantified with a 5 ml precision plastic measuring cylinder. We did not
measure stemflow given the low amount of water involved in this
process (1–3% of Pmm) (Shinzato et al., 2011). Thus, to calculate Imm we
directly subtracted the amount of Thmm from Pmm. We collected 1248
samples from 104 collectors during the period of study.
We used Eqs. (1)–(3) to calculate the monthly average of Pmm, Thmm,
and Imm, respectively, at each site. We also calculated the ratio of Imm
related to Pmm (I(%), Eq. (4)).
3 Vtot
( ∑i = 1 )
Pmm = A
10
n (1)
10 Vi
( ∑i = 1 )
Thmm = A
10
n (2)
Imm = Pmm − Thmm (3)
I
I% = ⎛ mm ⎞ ∗ 100
⎜ ⎟
P
⎝ mm ⎠ (4)
where Pmm is monthly observed precipitation in the open field (mm);
Thmm is monthly observed throughfall in the interceptometers (mm);
Imm is monthly observed rainfall interception (mm); n is the number of
collectors (pluviometers and interceptometers) used at each site to
collect total rainfall (n = 3) and throughfall (n = 10); Vtot is the total
amount of monthly rainfall collected in each pluviometer (ml); Vi is the
total amount of monthly throughfall collected in each interceptometer
(ml); A is the catchment area of each collector (cm2); and I(%) is the
monthly ratio of Imm related to Pmm.
2.2.2. Forest attributes
We measured four parameters of the plant community closely in-
volved with the interception function: (i) Density (individuals/ha), (ii)
Basal Area (BA – m2/ha), (iii) Species richness, and (iv) Ratio of de-
ciduous trees. These data were obtained by surveys conducted in four
4 × 50 m transects randomly established within each site, where trees
with diameter at breast height (1.30 m height) ≥ 5 cm were identified
and the diameter at breast height recorded. When required, leaf/floral
samples of the plants were collected for further identification at the
Escola Superior de Agricultura Luiz de Queiroz (ESA herbarium; USP).
We also collected data related to site condition: (v) Canopy continuity,
(vi) Vertical stratification, (vii) Cattle presence, and (viii) Grass cover.
A full description of the attributes and site condition evaluations is
available in the Supplementary material (Table A.2).
2.3. Data analysis
All data were analyzed n R 3.2.4 (R Core Team, 2016). ANOVA was
used to test for differences in interception data among sites and to select
the model that best fit Imm and Pmm. Outliers were removed. We used the
Student’s t-test to compare average I(%) between rainy and dry seasons.
We performed a self-organized maps (SOM) analysis, an artificial neural
network based on the topological distribution of the data of each
variable (Fig. A.3), allowing variables to cluster according to the si-
milarity of this distribution (Kohonen, 2013; Kohonen and Hari, 1999).
SOM is a similarity analysis, based on a non-parametric and recursive
regression process, applicable to discrete or continuous data (Kohonen,
1998) and without assuming any relations among variables. To perform
the analysis, the values of monthly I(%) and Pmm and the four vegetation
parameters were normalized by the standard score method – Z-score
normalization. Parameters related to site condition were measured as
binary values (Table 1; Table A.2). More information about SOM is
F.R. Gardon, et al. Ecological Indicators 109 (2020) 105779

Table 1
Annual interception (% of annual Pmm), forest attributes and site condition evaluated in each site.
Site Annual interception Tree density/ BA (m2/ Species Deciduous trees (%) Canopy continuity Vertical stratification Cattle presence Grass cover
(%) ha ha) richness

1 26.4 1438 25.9 24 71.0 0 0 1 1

2 21.4 125 0.4 6 71 1 1 0 0
3 21.8 625 7.3 17 81.0 1 0 0 1
4 23.5 975 23.7 24 57.6 0 0 1 1
5 22.6 588 15.6 21 61 0 0 1 1
6 14.9 813 7.4 21 31.6 1 1 0 0
7 22.8 513 4.9 11 40.5 1 0 0 0
8 15.6 675 4.5 21 43.4 1 1 0 0

available in the Supplementary material (Section 3).

3. Results

3.1. Precipitation and interception patterns

Restoration planting showed an average I(%) of 21.4 ± 3.9%

(minimum 14%, maximum 27%) of the annual Pmm, an amount
equivalent to 396 mm returning to the atmosphere. We found sig-
nificant differences in annual Imm among sites (ANOVA – p < 0.05).
About 65% of Imm collected monthly was concentrated at low values,
ranging from 3.1 to 30 mm. The model that best fit the data (Imm–Pmm)
was a linear model (ANOVA – p < 0.05, R2 = 0.8254, Residual
Standard Error = 12.1) (Fig. 1).
We found no statistical difference in average I(%) between the rainy
and dry seasons (t-test – p > 0.05). In the rainy season (October to
March) an average of 22.4% of Pmm was intercepted by the plant
community, corresponding to a ratio of 77% of annual Imm. The dry
season (April to September) showed an average interception of 18.3%,
corresponding to 23% of the amount of water annually intercepted
(Fig. 2).
3.2. Forest attributes
We identified 460 individuals belonging to 72 species and 22 fa-
milies. Only 14 species accounted for 65% of the total plants identified
(Table A.3). The families Anacardiaceae, Euphorbiaceae, Fabaceae,
Malvaceae, and Verbenaceae were the most frequent, representing 78%
of the plants identified. Data related to forest attributes and site con-
dition were disparate and presented different sets of conditions
(Table 1).
Fig. 1. TIF. (1.5 column). Models built with Precipitation (mm) and Interception
(mm), and Precipitation (mm) and Interception (%) values monthly collected. The
light shaded line and letters “w” (data collected in the wet season) and “d” (data
collected in the dry season) represent the linear model Imm-Pmm; The dark
shaded line and black circles represent the linear model I(%)-Pmm.
Fig. 2. TIF. (1.5 column). Average and standard deviation of Pmm, Imm, and
Ratio of interception (%) related to Pmm monthly observed among sites.
3.3. Effects of forest attributes on rainfall interception
SOM analysis revealed that BA and species richness are forest at-
tributes most positively correlated to each other and to I(%), observed
by the similarity of the maps (Fig. 3). Tree density was also correlated
with these variables and I(%) but at a lower intensity. A negative cor-
relation of Pmm and deciduous trees with I(%) could be observed. Canopy
continuity and vertical stratification were negatively correlated with
cattle presence and grass cover. Positive correlations were observed for
canopy continuity and vertical stratification and between cattle pre-
sence and grass cover.
4. Discussion
After a decade, restoration planting can intercept on average 21% of
total Pmm, a value similar to that of pristine and advanced stage tropical
forests in Brazil, 22.6% and 20.6%, respectively (Cuartas et al., 2007;
Alves et al., 2007). This highlights the potential of actively restored
forest in recovering the interception function in the short term. How-
ever, there was high variability of I(%) among sites (14% to 27%) and
values of individual sites were similar to those of secondary forests at
initial, intermediary, or even advanced stages of regeneration (Ghimire
et al., 2017; Lorenzon et al., 2013). We observed that monthly I(%) did
not change significantly in response to increased monthly Pmm (Fig. 1).
The amount of water intercepted by forests that returns to the at-
mosphere contributes to the maintenance of the water cycle at both the
regional and global scales (Oliveira et al., 2008). The control of erosion
processes may also result from interception reestablishment owing to a
decrease in the kinetic energy of water drips that reaches the soil,
promoting soil conservation (Geißler et al., 2013; Goebes et al., 2015).
These positive effects of interception are important regulating eco-
system services provided by restoration (Alamgir et al., 2016; Locatelli
et al., 2017; Viglizzo et al., 2016).
Increasing forest cover can also diminish the amount of water
available, as the plant community, aside from rainfall interception, also
uses groundwater for biomass accumulation (photosynthesis process)

3
F.R. Gardon, et al.
thereby reducing soil saturation (Calder, 1998; Liu et al., 2003; Tang
et al., 2007). Thus, rainfall interception can be an ecosystem disservice,
representing a loss of water that returns to the atmosphere by eva-
poration, instead of flowing into the ground and contributing to water
storage (Honda and Durigan, 2016). The dual nature of the inter-
pretation of rainfall interception is evident, and to define if this eco-
logical function leads to the loss or gain of water to the watershed
depends on the WES that is the focus of the evaluation.
Regarding seasonality, in perennial forests such as the Brazilian
Amazon “terra firme” forests, a marked difference in interception rates
between seasons can be observed, where interception (%) values in-
crease from the wet to the dry season because there is less precipitation
but the same biomass (Oliveira et al., 2011). However, we found no
difference in interception rates between seasons in the SSF, but SOM
analysis was sensitive enough to indicate a negative correlation of I(%)
and the ratio of deciduous species recorded. This could be explained by
the existence of many deciduous species in the studied sites; species
which lose their leaves in the dry season, resulting in a lower available
foliar area (Da Costa et al., 2014) capable of rainfall interception. This
supports the principle that restoration actions should rigorously select
the set of species to be planted to restore the native plant community
and recover the original rates of interception.
SOM results indicate that BA and species richness are attributes
closely related to I(%) (Fig. 3). BA is recognized as an attribute that
governs interception and can be managed to control water dynamics in
forested watersheds. An increase of 1 m2/ha in the BA of forests can
decrease the amount of water reaching the soil by 0.8% to 1%, and is
the attribute most indicated in interception modelling (Del Campo
et al., 2014; Dietz et al., 2006; Honda and Durigan, 2016; Molina and
del Campo, 2012; Suganuma and Durigan, 2015). Consistent with these
studies, we assume that managing this attribute could help the re-es-
tablishment of expected interception values and most of the ecosystem
services related to canopy cover.
Despite our average species richness (18 species) being substantially
lower than that in the reference ecosystem (i.e., SSF remnants) (Freitas
and Magalhães, 2014; Sartori et al., 2015), SOM analysis showed a
4
Ecological Indicators 109 (2020) 105779
Fig. 3. TIF. (1.5 column). Self-Organized Maps ob-
tained for each variable sampled within the sites.
Axes are the position of the neurons distributed in the
Kohonen’s layer (20 × 20 neurons). Similar colors
represent positive correlations, and inverse colors
negative correlation. (color version in the
Supplementary material – Fig. A.4).
positive relation with I(%). Studies in natural forests (Geißler et al.,
2013; Goebes et al., 2015; Liu et al., 2003; Tang et al., 2007; Wang and
Duan, 2010) have found different responses of hydrological processes to
the increase in species richness, such as decreased runoff and erosion
and improved infiltration. Heterogeneous forests (high diversity) are
systems more vertically and horizontally stratified, where species with
different characteristics (canopy, leaf dimensions, foliage, and branches
density) inhabit different sites and niches (Wang and Duan, 2010).
According to our results, it may be suggested that species richness is
important to better understand ecological processes in the water-plant
interface.
The negative relation between canopy continuity and stratification
with cattle presence and grass cover was as expected (Fig. 3). Grazing
leads to sapling damage and death, which in turn compromises canopy
closing and provides more radiation benefit for grasses that compete
with tree saplings for resources (Brancalion et al., 2009; Holl et al.,
2000; Ignácio et al., 2007; Parrotta et al., 1997; Sampaio and Guarino,
2007).
We observed that a planted community in sites without degradation
factors may present more structured vegetation and consequently a
higher interception (Table 1). For this reason, implementing adequate
monitoring actions is crucial to ensure success of restoration over time.
It is known that canopy continuity and stratification are determining
factors in the interception processes (Crockford and Richardson, 2000),
but maybe SOM did not identify relations between these parameters
and I(%) because they were measured as binary values.
SOM analysis demonstrated that better-structured forests – higher
tree density, BA, species richness, canopy continuity, and vertical
stratification present higher rainfall interception. Thus, our study cor-
roborates Crockford and Richardson (2000); the variability of inter-
ception can be a consequence of different conditions observed among
the plant community, including every attribute mentioned earlier.
Restoration is a slow and vulnerable process in which it takes a long
time for attributes such as biodiversity and carbon stocks to reach va-
lues similar to those of mature forests (Wheeler et al., 2016); however,
our results indicated that the interception function can be quickly
F.R. Gardon, et al.
recovered if monitoring actions succeeded in avoiding degradation
factors. As active restoration is an expensive investment, we urgently
need to determine if WES and plant community are being effectively
recovered by these actions, justifying the costs of implementation.
5. Conclusions
We present early ecological outcomes related to the reestablishment
of an important hydrological process by restoration forest sites. Our
study provides evidence that young actively restored forests (10 years)
can play an important role in water cycle maintenance in tropical
landscapes and through rainfall interception return to the atmosphere
an amount of water similar to that returned by pristine and advanced
secondary forests. Seasonality affects interception efficiency and the
magnitude of the recovery of the interception function depends on the
attributes of the vegetation. Additionally, while basal area has been
recognized as a determinant of rainfall interception, species richness
also influences the recovery of interception rates. However, achieving
the goals of restoration and ensuring success depends on effective
monitoring actions. Our findings demonstrate the synergistic outcomes
of restorations that extend beyond recovering a focal ecosystem service
contributing to the increment in plant diversity in degraded tropical
landscapes.
Acknowledgment
We would like to register our sincere appreciation to each land-
owner, to the Environmental Agency of the State of São Paulo and to
the School of Civil Engineering, Architecture and Urban Design of the
State University of Campinas for collaborating with the study.
This study was supported by CAPES – Coordenação de
Aperfeiçoamento de Pessoal de Nível Superior [grant number:
01P03428/2014] and PROAP – Programa de Apoio a Pós-Graduação.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://
doi.org/10.1016/j.ecolind.2019.105779.
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