Marine Environmental Research 172 (2021) 105506

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Marine Environmental Research

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

Plant invasion affects vegetation structure and sediment nitrogen stocks in

subtropical mangroves
Jéssica Airisse Guimarães Sampaio a, Carla Roberta Gonçalves Reis b, Marília Cunha-Lignon c,
Gabriela Bielefeld Nardoto a, b, Luiz Felippe Salemi a, *
a
Núcleo de Estudos e Pesquisas Ambientais e Limnológicas – Programa de Pós-Graduação Em Ciências Ambientais, Área Universitária 1, Vila Nossa Senhora de Fátima,
Campus de Planaltina, Universidade de Brasília, 73340-710, Planaltina, Distrito Federal, Brazil
b
Programa de Pós-Graduação Em Ecologia, Instituto de Ciências Biológicas, Campus Darcy Ribeiro, Universidade de Brasília, 70910-900, Brasília, Distrito Federal,
Brazil
c
Campus Experimental de Registro, Universidade Estadual Paulista, 11900-000, Registro, São Paulo, Brazil

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

Keywords: Plant invasion can primarily affect the structure and functioning of terrestrial and aquatic ecosystems. Although
Aboveground biomass there is evidence that plant invasion can modify organic matter dynamics in mangroves, it is uncertain whether
Nutrient cycling and to which extent these changes can affect carbon (C) and nitrogen (N) dynamics in the sediment-plant system.
Exotic species
Here, we measured: (i) the structure of native vegetation and C and N in the sediment-plant system in subtropical
Nitrogen Pollution
Tree Mortality
mangroves subjected to aquatic macrophytes invasion in southeastern Brazil. We answered the following
questions: i) Do invaded mangroves differ in aboveground biomass compared to non-invaded mangroves?; ii) Are
there C4 macrophytes in these sites? iii) What are the C and N stocks in sediment of invaded mangroves? We
quantified C and N concentrations and the isotopic signature of such elements (δ13C and δ15N) in the sediment-
plant system, the C and N stocks in the sediment (0–20 cm depth), and mangrove aboveground biomass.
Mangrove aboveground biomass was lower at invaded compared to non-invaded sites reflecting the species
displacement in invaded sites. The sediment at invaded mangroves did not significantly contribute to C4 sources
because of the large predominance of both mangrove and invasive C3 plants. While sediment C stocks were
similar among study sites (~47 Mg ha− 1), N stocks were lower at invaded (2.7 Mg ha− 1) comparing to non-
invaded (3.2 Mg ha− 1) mangroves. The lower N stocks at invaded sites can reflect the higher leaf N concen­
trations and lower C:N ratios of invasive plants compared to mangroves. Thus, the effects of macrophytes in­
vasion in subtropical mangroves are more apparent for vegetation structure and N stocks. C stocks alteration is
expected the be detectable in the future.

1. Introduction
Mangroves are highly productive forest ecosystems (Alongi, 2012;
Donato et al., 2011) that occur at the interface of terrestrial and brackish
water environments in tropical, subtropical, and warm temperate re­
gions worldwide (Schaeffer-Novelli et al., 2000; Spalding et al., 2010).
Mangroves play critical roles in biogeochemical cycling at the
terrestrial-aquatic interface in coastal areas and help dictate carbon (C)
and nitrogen (N) storage and transfer at a global scale (Alongi, 2014,
2018; Donato et al., 2011; Reis et al., 2017; Davidson et al., 2018). For
instance, although mangroves comprise only 1% of Earth’s terrestrial
surface, these ecosystems fix 11 Mg C ha− 1 y− 1 in aboveground biomass
and contribute 10–15% of C storage in soil globally (Alongi, 2014). The
high C storage in mangroves is a consequence of their high productivity
and C allocation in belowground biomass associated with tidal inputs of
organic matter and low decomposition rates due to periodic flooding
and low oxygen status in sediment (Bouillon et al., 2008; Kristensen
et al., 2008; Rovai et al., 2021). However, mangrove C stocks are also
highly responsive to environmental change, which have the potential to
substantially affect mangrove’s biogeochemical roles in coastal areas
and their climate feedbacks (Hutchison et al., 2014).
Biological invasion is one of the main drivers of global environ­
mental change with harmful consequences to biodiversity and
ecosystem structure and functioning (Biswas et al., 2018; Liao et al.,

* Corresponding author.
E-mail address: lfsalemi@gmail.com (L.F. Salemi).

https://doi.org/10.1016/j.marenvres.2021.105506
Received 4 June 2021; Received in revised form 13 October 2021; Accepted 15 October 2021
Available online 16 October 2021
0141-1136/© 2021 Elsevier Ltd. All rights reserved.
J.A. Guimarães Sampaio et al.
2008; Steffen et al., 2015; Vitousek et al., 1996; Zhang et al., 2012).
Several studies have reported a growing number of invasive species in
mangrove ecosystems (e.g. Biswas et al., 2018, 2012, 2007; Ren et al.,
2014; Röderstein et al., 2013; Wang et al., 2018; Zhang et al., 2010).
Invasive plants can attain notably large populations and become domi­
nant in invaded ecosystems, thus affecting the performance and survival
of native species, and at the ecosystem level, it can affect the structure of
native vegetation (Vila et al., 2011; Vilà and Weiner, 2004). For
instance, native vegetation biomass reduced about 50% in experimental
settings due to the co-occurrence of invasive species (Vilà and Weiner
2004). Moreover, beyond impacting the native community structure and
aboveground biomass, invasive plants often exhibit nutrient re­
quirements and organic matter quality different from the native species
(Liao et al., 2007, 2008; Lee et al., 2017). The input of organic matter
from invasive species in the soil can modify decomposition rates and
ultimately the ecosystem C and N cycling and stocks (Asthon et al., 2005;
Liao et al., 2007, 2008; Ehrenfeld, 2010; Vila et al., 2011; Lee et al.,
2017; Su et al., 2020). However, the direction and magnitude of such
changes are still a challenge. A few studies have shown an increase of
sediment C stocks in mangroves and saltmarshes following plant inva­
sion (Liao et al., 2007; Chen et al., 2015; Wang et al., 2018). On the other
hand, some studies have reported a reduction (Mao et al., 2011) or no
effect of plant invasion on C stocks in mangroves (Lu et al., 2014;
Davidson et al., 2018). Therefore, despite the critical biogeochemical
roles of mangroves in coastal areas and the increase of plant invasion in
these systems, more is needed to understand interactions between plant
invasion and C and N dynamics in these mangroves.
The leaf and litter quality differences between native and invasive
species can explain the direction, magnitude, and mechanisms of alter­
ations in the fluxes and stocks of C and N in soil (Lee et al., 2017). In this
way, the C:N stoichiometry in plants and soil represents a critical tool to
investigate the effects of plant invasion in ecosystems. Knowledge of the
consequences of plant invasion can be further refined by combining
stoichiometric and isotopic approaches. The natural abundance of C
(δ13C) and N (δ15N) isotopes in the plant-soil system has been shown to
be central in explaining changes in soil organic matter decomposition
(Craine et al., 2015; Feng et al., 2017) and in the stocks of these elements
in soil due to environmental change (Zhang et al., 2010; Feng et al.,
2017). The δ13C indicates the degree of fractionation against the heavier
isotope 13C during photosynthesis (Ehleringer et al., 2000). Plants with
different photosynthetic pathways (e.g., C3 vs. C4) exhibit different δ13C
signatures. Thus, the δ13C offers a way to assess the magnitude of plant
invasion impacts in the ecosystem. The δ15N integrates the isotopic
signature of N sources and the fractionation against 15N in trans­
formation reactions in the sediment of mangroves and N losses to the
atmosphere (Reis et al., 2019).
Here, we combined stoichiometric and isotopic approaches to
investigate the effects of plant invasion on C and N stocks in sediment,
C4 vs. C3 contributions to soil organic matter in a subtropical mangrove
located in southeastern Brazil. Since 2010, mangroves at the north
section of this region have been increasingly invaded by aquatic mac­
rophytes (Cunha-Lignon et al., 2011, 2015; Cunha-Lignon and Men­
ghini, 2016; Prado et al., 2019; Sampaio et al., 2021). Several aquatic
macrophytes are recognized as some of the most aggressive invasive
species, given their exceedingly high productivity, ecological tolerance,
and dispersal ability (Pieterse and Murfhy, 1993; Chambers et al., 2008).
Because macrophytes might present C4 metabolism (Mortillaro et al.,
2016) and also high N uptake and content (Preiner et al., 2020), we
selected invaded and non-invaded mangroves to answer the following
questions: i) What are the differences between invaded and non-invaded
mangroves in terms of aboveground biomass?; ii) Is there a C4 contri­
bution to organic matter in the sediment of invaded sites? iii) What are
the C and N stocks in the sediment of invaded mangroves?
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Marine Environmental Research 172 (2021) 105506
2. Material and methods
2.1. Study area
The study was conducted in eight plots in fringe mangroves sites in
the northern portion of the Estuarine Lagunar Complex of Cananeia-
Iguape, southeastern Brazil, in 2017 (Fig. 1). The ECL-Cananeia-
Iguape (hereafter referred to as ECL) is formed by estuarine and
lagoon channels (Mahiques et al., 2013). Part of the ECL has been a
UNESCO World Heritage Site since 1999, and, in 2017, it was included
as a Ramsar site. The ECL-Cananeia-Iguape is divided into two portions.
The southern portion (the Cananéia region) comprises the most pro­
tected and conserved mangroves areas in the São Paulo State (Cunha-­
Lignon et al., 2011). The northern portion (the Iguape region) is largely
affected by the diversion of freshwater from the Ribeira de Iguape River
through the artificial canal called Valo Grande (Mahiques et al., 2009,
2013), that carries excess anthropic N from both sewage and agriculture
to the estuary and mangroves in the Iguape region. Due to the high
intake of fresh water and the consequent reduction in salinity, areas of
fringe mangroves are invaded by aquatic macrophytes (Fig. 1 Supple­
mentary) associated with reducing mangrove trees (Cunha-Lignon et al.,
2011, 2015; Cunha-Lignon and Menghini, 2016; Prado et al., 2019).
The climate in the ECL-Cananeia-Iguape is classified as “Cfa” in the
Köppen-Geiger climate classification (Alvares et al., 2013). Mean
monthly temperature ranges from 24.5 ◦ C in January to 7.8 ◦ C in July.
Mean annual rainfall is about 2300 mm, without a marked seasonality.
The Ribeira de Iguape River is one of the largest rivers in southeastern
Brazil, draining more than 23,350 km2. Tides are predominantly semi­
diurnal, with a mean amplitude from 0.82 to 1.25 m during spring tides
(Schaeffer-Novelli et al., 1990).
Mangrove vegetation in the region is composed of Rhizophora mangle
L. (Rhizophoraceae), Laguncularia racemosa (L.) C.F. Gaernt. (Com­
bretaceae) and Avicennia schaueriana Stapf & Leechm. ex Moldenke
(Acanthaceae) (families according to APG III, 2009) (Schaeffer-Novelli
et al., 1990). The studied fringe mangroves are dominated by
L. racemosa, except of one site which is dominated by Rhizophora mangle.
The dominant aquatic macrophytes species at the invaded mangrove
sites are Hymenachne amplexicaulis (Rudge) Nees (Poaceae), Bacopa
monnieri (L.) Wettst. (Plantagenaceae), Crinum salsum Ravenna (Amar­
yllidaceae), and some other four morpho species (not identified)
belonging to Poaceae and Cyperaceae.
Because of the natural variability of the sedimentary dynamics
within the estuarine system and the influence of the discharge of the
Iguape River close to the Valo Grande artificial channel, the studied sites
differ in terms of sediment properties and salinity (Table 1).
2.2. Sampling design
Four plots of both invaded and non-invaded mangroves were
selected (see Table 1). In all plots, mature leaves (green and fully
expanded) were obtained from the upper canopy of three arboreal in­
dividuals with diameter at breast height (DBH) higher than 4 cm. Mature
leaves from three individuals of every aquatic macrophyte present in
every invaded plot were also sampled. A composite litter (yellow and
dark leaves) was hand-collected from the forest floor below the sampled
trees. Three sediment samples at 0–10 and 10–20 cm depth were
sampled below the mangrove trees sampled. In addition, three undam­
aged sediment samples were collected in each plot to determine the bulk
density.
Leaf and litter samples were washed with tap water and then oven-
dried at 40 ◦ C for 48 h immediately after fieldwork. All sediment sam­
ples were kept refrigerated during transportation and until the labora­
tory analysis was carried out. In the laboratory, leaf and litter samples
were oven-dried at 55 ◦ C for 48 h and grounded into a fine powder with
liquid nitrogen. Then, they were weighed with an analytical balance
(2–3 mg) and stored in tin capsules.
J.A. Guimarães Sampaio et al. Marine Environmental Research 172 (2021) 105506

Fig. 1. Location of the four fringe mangroves studied sites. Two plots were located in each of the sites (●) totaling four plots on both non-invaded and
invaded mangroves.

Isotope ratios are reported in per mil (‰), where δ 13C is reported
Table 1
relative to the Vienna Pee Dee Belemnite (VPDB; 13C:12C ratio =
Physical-chemical properties of the sediment of the studied plots both in non-
0.01118) standard and δ15N is reported relative to atmospheric air (AIR;
invaded and invaded fringe mangroves. 15 14
N: N ratio = 0.0036765). Internal standards (tropical soil and sug­
Mangrove arcane leaves) are routinely interspersed with target samples to correct
Non-invaded Invaded for mass effects and instrumental drift during and between runs. Long-
Sand (%) 60–74 25–87 term analytical error for the internal standards is of 0.2‰ for both δ
13
Silt (%) 11–22 10–46 C and δ 15N, 1% for organic C, and 0.02% for total N.
Clay (%) 15–18 3–29
pH H2O 5.5 ± 0.2 4.3 ± 0.1
2.3. Vegetation structure
OM (g/kg) 108 ± 2.8 122.5 ± 7.7
Available P (mg.kg− 1) 40.8 ± 3.5 38.6 ± 2.1
K+ (mmolc.kg− 1) 16.8 ± 5.4 3.1 ± 0.8 Metrics of the vegetation structure were recorded in all plots
Ca2+ (mmolc.kg− 1) 67.5 ± 0.7 77 ± 0.1 following the method available in Schaeffer-Novelli et al. (2015).
Mg (mmolc.kg− 1) 150.5 ± 31.8 62.5 ± 3.5 Within each plot, every mangrove arboreal individual over 1 m tall
Na+ (mmolc.kg− 1) 387.1 ± 36.9 35.4 13.7
had his DBH recorded with a measuring tape in diameter at DBH, and
Al3+ (mmolc.kg− 1) 12 ± 16 25 ± 0.1
CEC (mmolc.kg− 1) 699.2 ± 80.9 376.1 ± 25.2 height (H) (m) recorded using a rangefinder or telescopic pole. The
BS (mmolc.kg− 1) 621.7 ± 73.1 178.1 ± 18.1 condition of the trunk was also checked (either alive or dead). The
Salinity (PSU) 14.5 ± 5.6 2.2 ± 1.1 average value of the basal area of live and dead trunks (%) per plot was
calculated. To calculate the living basal area (BA) by the plot area (m2/
ha), the formula “BA = π (DBH) 2/4′′ was used. The following allometric
Sediment samples were oven-dried at 55 ◦ C for 48 h, then sieved in 2
equations (see Medeiros and Sampaio, 2008) were used to estimate the
mm mash. After that, every sample had fine roots and shell residues
living biomass aboveground (AGB) (Mg/ha):
removed manually and then subjected to an acid wash treatment with
10% HCl to remove carbonates (Ingram, 1971). Subsequently, every (
AGB = 0, 2752* DBH 2 *H
)0,8529
for R. mangle
sample was oven-dried again at 55 ◦ C for 48 h and ground into a fine
powder with mortar and pistil. Finally, subsamples were weighed with ( )0,8615
AGB = 0, 1214* DBH 2 *H for L.raecemosa
an analytical balance (1.5–2.0 mg) and stored in tin capsules.
C and N concentrations and C and N isotope ratios were determined ( )0,8298
by combustion using an elemental analyzer (Carlo Erba, CHN-1100) AGB = 0, 1327* DBH 2 *H for A. schaueriana
coupled to a Thermo Finnigan Delta Plus mass spectrometer at the
Laboratory of Isotope Ecology of the Centro de Energia Nuclear na
Agricultura (CENA/Universidade de São Paulo), Piracicaba, SP, Brazil.

3
J.A. Guimarães Sampaio et al.
2.4. Carbon and nitrogen stocks in the sediment
All undisturbed sediment sample was oven-dried at 105 ◦ C for 72 h
and weighed to determine the sediment density using the ratio of the dry
weight of the sediment to the fixed volume of the sampled cylinder core
to calculate the bulk density. C and N stocks in the sediment were
calculated based on the C and N concentrations in the sediment, the bulk
density 2), and the thickness of the sediment layer considered (20 cm)
according to Veldkamp (1994):
C or N stock (Mg / ha) = (C or N x bulk density(kg dm − 3 x e) / 10
2.5. Statistical analysis
Normality of the data and residuals were examined through the
Shapiro-Wilk test. Homocedasticity was evaluated using the Barlett test.
The following variables followed the normal distribution: basal area,
dead trunks basal area (%), aboveground biomass, C and N stocks). In
this case, we used the T-test to compared invaded and non-invaded sites.
We used the Mann-Whitney test for the non-normal variables (foliar and
litter C:N ratio, δ13C, δ15N, N concentration). All statistical analyses were
carried out using R software (R Core Team, 2017) at p < 0,05.
3. Results
3.1. Vegetation structure
The dominance of basal area of live stems in non-invaded were
higher (from 92 to 100%) compared to invaded. In contrast, the basal
area of dead stems was lower in non-invaded areas compared to invaded
(from 20 to 51%). Thus, the basal area of dead trunks (DTBA) was
significantly higher in the invaded areas (Table 2). Consequently, the
aboveground biomass of native mangrove was significantly lower in the
invaded areas than in the non-invaded mangrove areas (Table 2).
We recorded 24 invasive species of macrophytes in July 2017 in the
invaded sites. The minority (12%) of the species had C4 photosynthetic
metabolism (Table 1 Supplementary). The dominant species were
Hymenachne amplexicaulis (Rudge) Nees (Poaceae) and other three non-
identified from the following families: Poaceae, Cyperaceae,
Amaryllidaceae.
3.2. Plants – C and N isotopes and concentration
There were no significant differences of C:N ratio between our sites.
Foliar N of the invaded sites (including native and invasive plants) was
higher than non-invaded areas (Fig. 2a and b). Foliar δ13C and δ15N were
also higher in the invaded sites compared to non-invaded (Fig. 2c and d).
In the invaded sites, macrophytes had C:N ratio lower compared to
native plants (Fig. 3a). Moreover, foliar N was higher in the invasive
macrophytes than native plants (Fig. 3b). There were no differences
between macrophytes and native plants concerning both foliar δ13C and
δ15N (Fig. 3c and d).
C:N was significantly higher in native plants in the invaded sites than
non-invaded (Fig. 4a). Foliar N was substantially lower in native plants
Table 2
Mean and standard deviation of variables related to vegetation structure (DTBA,
dead trunks basal area, BA, basal area; AGB, aboveground biomass).
Mangrove
Non-invaded Invaded
DTBA (%) 4.1 ± 3.95a 33.1 ± 14.3b
BA (m2 ha− 1) 41.9 ± 7.28a 18.6 ± 8.2b
AGB (Mg ha− 1) 320.9 ± 133.8a 92.3 ± 46.1 b
Different letters indicate significant differences between invaded and non-
invaded sites.
4
Marine Environmental Research 172 (2021) 105506
in the invaded than non-invaded (Fig. 4b). Both foliar δ13C and δ15N
were significantly higher in native plants in the invaded sites compared
to non-invaded (Fig. 4c and d).
3.2.1. Litter – C and N isotopes and concentrations
There were no significant differences in litter charactersitics such as
C:N, N, δ13C and δ15N between invaded and non-invaded sites (Fig. 2
Supplementary Material).
C:N ratio of invasive plants-derived litter was significantly lower
than that of native plants (Fig. 5a). There were no significant differences
in N content and litter δ13C between litter from invaded and native
plants (Fig. 5b and c). Foliar δ15N was significantly higher in invasive
plant derived litter than the one generated by native plants (Fig. 5b).
3.3. Sediment – C:N ratio, isotopes, C and N stocks
δ13C, δ15N, %N e C:N had no differences between sampled depths
(0–10 e 10-20 cm). Thus, for the sake of comparison between invaded
and non-invaded sites, we used 0–20 cm layer. C:N, N content, δ13C,
δ15N had no significant difference between sites (Fig. 1S).
C stocks had no significant difference between invaded and non-
invaded sites (Fig. 6a). N stocks, in turn, had significant differences
between invaded and non-invaded sites (Fig. 6b).
4. Discussion
The invasion of macrophytes substantially modified the structure of
native mangrove vegetation in invaded areas. The likely high mortality
in invaded sites (Cunha-Lignon et al., 2015; Prado et al., 2019) and
lower live basal area indicated a lower density of live native individuals
than non-invaded sites. This, in turn, led to lower aboveground biomass
observed here. Consequently, this may indicate a likely reduction of C
stocks in the aboveground biomass which, in turn, may compromise
important ecosystem services such as C storage (Alongi, 2012; Donato
et al., 2011).
Invasive plant species can modify native vegetation structure and
composition, impede mangrove growth and development, and hinder
natural regeneration (Biswas et al., 2018). The high number of dead
individuals observed in the invaded areas may be associated with loss of
competitive ability of native plants. Similar findings have been found
elsewhere. For instance, the total biomass of a native mangrove species
was four times lower compared to areas of native forests after
inter-planting with exotic species (Lu et al., 2014). Furthermore,
replacing a woody individuals such as mangroves with grasses or mac­
rophytes promoted the reduction of aboveground biomass, which may
significantly reduce C and N stocks in the mangroves soil-plant system
(Wang et al., 2018).
Few species had a C4 metabolism. Thus, unlike other studies that
demonstrably showed that C4 invasive plants contributed substantially
to soil organic matter (Xia et al., 2021), due to the low number of in­
dividuals of such metabolism, the same was not observed here.
Though macrophytes include a high diversity of taxons (Chambers
et al., 2008), they generally exhibit common characteristics such as high
productivity, short-life cycle, and high nutrient efficiency to capture
nutrients from both sediment and water (Byers et al., 2006; Pieterse and
Murfhy, 1993; Wetzel, 2001). Under high nutrient availability, these
plants can present a significant increase in biomass in few days
(Chambers et al., 2008). The high foliar N and δ15N in invasive plants
resulted in a high uptake by these plants, which reflects higher N
availability in invaded sites due to sewage and agricultural runoff (Reis
et al., 2019). In fact, invasive macrophytes had high foliar N and δ15N,
leading to a two-fold decrease in C:N ratio than native mangrove species
in invaded sites.
The lower C:N ratio in the litter of invasive plants compared to the
one of their native counterparts likely generated a reduction of N stocks
in sediment observed here. In addition, a recent study detected high N
J.A. Guimarães Sampaio et al. Marine Environmental Research 172 (2021) 105506

Fig. 2. Foliar C:N (a), N (b), δ13C (c) and δ15N between invaded and non-invaded sites ***, * and NS denote p ≤ 0,001, p ≤ 0,05, Non-significant differences
respectively.

Fig. 3. Foliar C:N (a), N (%) (b), δ13C (‰) (c) and δ15N foliar (‰) (d) of mangrove and invasive macrophytes in invaded sites.*** and NS denote p ≤ 0,001 and non-
significant differences, respectively.

5
J.A. Guimarães Sampaio et al. Marine Environmental Research 172 (2021) 105506

Fig. 4. Foliar C:N (a), N (b), δ13C (c) and δ15N (d) of native magrove species of invaded and non-invaded sites. *** and ** denote p ≤ 0.001 and p ≤ 0.01 respectively.

Fig. 5. Litter C:N (a), N (b), δ13C (c) and δ15N (d) of native and invasive species in invaded sites. ** and NS denote p ≤ 0.01 and non-significant differences
respectively.

6
J.A. Guimarães Sampaio et al.
Fig. 6. Carbon (a) and nitrogen (b) stocks in sediment of invaded and non-invaded sites. ** and NS denote p ≤ 0,01 and non-significant differences, respectively.
availability in sediment of our invaded sites to be one of the main factors
driving non-plant species in the region (Sampaio et al., 2021). High N
availability in sediment (Sampaio et al., 2021) along wiht a low C:N
litter derived from macrophytes may increase microbial activity which
may increase decomposition and mineralization rates (Lee et al., 2017)
with potential consequences for C and N stocks reduction over time. In
fact, Wang et al. (2018) observed a significant reduction in the N stock in
the sediment of mangrove areas after invasion. Unlike our findings and
those by Wang et al. (2018), a global analysis on the effect of plant in­
vasion on C and N cycles verified an increase in soil C and N stocks with
an increase in litter decomposition (Liao et al., 2008). On the other hand,
other meta-analysis also showed an increase in C and N stocks with a
decrease in decomposition (Vilà et al., 2011). Though different re­
sponses of decomposition have been shown by these studies, they indi­
cate an increase in C and N stocks in invades sites. Given the lack o clear
direction on the effects of invasion on C and N stocks in sediment and
that short-term responses might differ compared to long-term, more
studies are needed.
The C stocks had no differences between sites. The lower above­
ground biomass in invaded sites may reduce the input of litter (Costa
et al., 2020) in the long term which may reflect directly on C stocks. In
addition, given that effects of invasion on organic matter in similar sites
are more evident after 7–10 years (Wang et al., 2018), we hypothesize
that differences in C stocks in our sites will be more apparent in the
future. Based on the lower N stocks at invaded sites which reflected the
higher leaf N concentrations and lower C:N ratios of invasive plants
compared to mangroves, we expect a decrease in C storage in long run.
Thus, more studies are needed to monitor C in coastal wetlands like ours
to clarify mechanisms shaping long-term carbon responses to such
invasion.
There was a lack of significant differences between invaded and non-
invaded sites regarding sediment δ15N and C:N ratio. Given the clear
differences of these variables regarding invasive and native plants in
invaded sites, we speculate that as macrophytes become more dominant
over time, both δ15N and C:N ratio will change. Unlike our findings,
previous studies showed that sediment δ15N in an invaded coastal
wetland was significantly higher compared to non-invaded sites due to
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Marine Environmental Research 172 (2021) 105506
the increase of N losses through increased N mineralization and leaching
(Cheng et al., 2008; Yang et al., 2015).
A global picture of δ15N provided some interesting insights regarding
the ecosystem functioning of our sites. In other words, given that (i)
invaded sites had higher foliar δ15N compared to non-invaded, (ii) litter
from invasive plants were also higher compared to native plants in
invaded sites, and (iii) native species from invaded sites also had lower
foliar δ15N compared to native species in non-invaded sites. These dif­
ferences altogether indicate that N sources are likely changing in
invaded compared to non-invaded sites. The presence of reactive N from
both sewage and agricultural runoff likely generated an increase in N
cycling which, in turn, favored losses (i.e. gaseous and in water) which,
in turn, promoted an accumulation of the heavier isotope in soil for
plants in invaded sites. Consequences for C stocks in sediment are yet to
be seen in these sites.
Author statement
I declare the following contributions to the presente manuscript:
Jéssica Airisse Guimarães Sampaio: conceptualization, writing,
formal analysys, funding acquisition.
Carla Reis – conceptualization.
Marília Cunha-Lignon – data curation, writing, resources.
Gabriela Bielefeld Nadoto - conceptualization, writing, funding
acquisition, project administration.
Luiz Felippe Salemi - conceptualization, writing, formal analysis,
investigation.
Declaration of competing interest
The authors declare no conflict of interest.
Acknowledgment
We would like to thank the Instituto Chico Mendes de Conservação
da Biodiversidade (ICMBio-Brazil) for the authorization to conduct this
research. This work was supported by the Coordenação de
J.A. Guimarães Sampaio et al.
Aperfeiçoamento de Pessoal de Nível Superior CAPES – Brazil [001 to J.
A.G.S. and 1422671 to C.R.G.R]; the Conselho Nacional de Desenvol­
vimento Científico e Tecnológico CNPq– Brazil [445418/2014–1 to M.C-
L]; the Fundação Grupo O Boticário, edital Bio&Clima Lagamar
[BL0006_2012_1 to M.C-L.]; Fundação de Amparo à Pesquisa no Distrito
Federal (FAPDF) [0193.000344/2017];
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.marenvres.2021.105506.
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