Ecological Indicators 130 (2021) 108027

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

Invasive Spartina alterniflora in controlled cultivation: Environmental

implications of converging future technologies
Shameer Syed, Min Xu, Zaifeng Wang, Chengfeng Yu, Bin Lian *
College of Life Sciences, College of Marine Science and Engineering, Nanjing Normal University, No. 1, Wen Yuan Road, Nanjing 210023, Jiangsu Province, China

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

Keywords: Invasive species have become a major threat to ecosystem stability, diversity and sustainability in recent times.
Invasive plant Invasive plants such as Spartina alterniflora (SA), outgrow resident flora by rapidly utilizing essential soil nu­
Biomass production trients depriving the native plant species, impacting the ecosystem diversity and stability. SA is also notorious for
Soil resilience
its effect on local biodiversity as it grows and spreads faster, ultimately dominating local biodiversity. This
CO2 sequestration
Coastal area protection
dominance and the negative effect of invasive SA is due to features such as allochemical’s secretion, positive
feedback between plant and soil, and rapid absorption of soil nutrients, exerting a competitive advantage over
native plants. Therefore, in this review, we aim to contemplate the implications and potential applications of SA
under control conditions or special circumstances as a source of biomass, soil restorer, CO2 capture agent, soil
resilience enhancer and pollution remover as most of the studies on SA have focused on controlling its impact on
the local ecosystem. On the other hand, SA’s ability to grow quickly in poor alkaline soils is useful but very little
is explored to exploit SA for biomass production in controlled cultivation in different environmental conditions.
In addition to these aspects, the CO2 sequestration capability requires greater focus, especially in sensitive salt
marshes and wetlands along the coastal areas.

1. Introduction Vilà et al., 2011; Prabakaran et al., 2019).

In the 21st century, biological invasion caused primarily by the
human introduction of non-native organisms is a serious global
ecological challenge that significantly threatens local biodiversity and
ecosystem stability. These invasive plants are biological invaders and
are known as unwanted weeds and detrimental to plant diversity and the
environment because of their rapid growth and proliferation (Hender­
son, 2001; Mashhadi and Radosevich, 2004; Liu et al., 2017). In addi­
tion, these invasive plants are one of the major threats to indigenous
biodiversity and contribute directly or indirectly to habitat loss of
approximately 42% of indigenous and endangered species (Pimentel
et al., 2005). However, the intriguingly rare characteristics of invasive
species such as resilience, aggressive growth can be utilized under
control conditions for alleviating and remediating multiple facets of
human activity-induced soil pollution and degradation, along with the
fight against environmental predicaments such as CO2 emissions, etc.
Additionally, steerable invasive plants have been used to restore
degraded ecosystems such as wetlands, salt marshes and enhance their
resilience to environmental fluctuations (Pyšek and Richardson, 2010;
Spartina alterniflora (SA) a perennial grass, native to North America’s
Atlantic coast habituated to low tidal wetlands or coastal marshes
(Zhang et al., 2017). SA is also a prolific invader, capable of spreading in
many ecological niches through seeds, rhizomes, or vegetative frag­
ments (Duns, 2020; Meng et al., 2020). Furthermore, SA is one of the
100 most destructive invasive species (increasing in area at a rate of 40%
annually), Weber et al. (2008) and Xu et al. (2012) recorded a 55%
annual spread rate in Chinese coastal areas, making it a threat to both
native biodiversity and ecosystem functions if left unchecked. The
deliberate introduction of SA to different coastal marshlands in Europe,
Asia, and Australia was to utilize its potential to dispel tidal wave en­
ergy, trap and stabilize sediments creating a vertical accretion of soil
(Wan et al., 2009; Meng et al., 2020). Furthermore, SA introduction in
Chinese coastal marshes from the late 1970s was exclusively for coastal
erosion prevention (Zhang et al., 2017). Despite the negative impacts of
the SA invasion, multiple benefits are reported, including ecosystem
restoration, soil functionality improvement and serves as a carbon stock
and sink in wetlands and salt marshes (Yang et al., 2016a,b). In this
review, we highlight SA’s potential applications such as

* Corresponding author at: Nanjing Normal University, No.1 Wen Yuan Road, Qi Xia District, Nanjing, China.
E-mail address: bin2368@vip.163.com (B. Lian).

https://doi.org/10.1016/j.ecolind.2021.108027
Received 24 May 2021; Received in revised form 17 July 2021; Accepted 22 July 2021
Available online 27 July 2021
1470-160X/© 2021 The Author(s). Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license
(http://creativecommons.org/licenses/by-nc-nd/4.0/).
S. Syed et al. Ecological Indicators 130 (2021) 108027

phytoremediation, erosion control, production of biomass in depleted
and contaminated soils with restricted nutrient availability for biofuel
substrates, thus improving soil functional matrices. Thus, intrigue cur­
rent researchers to study such invasive plants beyond their assumed
implications but under controlled environments.
2. Methods
2.1. Search strategy and inclusion criteria
To identify research articles, reviews and relevant databases appli­
cable to this study, the search strategy was mainly keyword-based search
in literature search engines like google scholar (https://scholar.google.
com/), web of sciences (https://www.webofknowledge.com/),
PubMed (https://pubmed.ncbi.nlm.nih.gov/) and science direct
(https://www.sciencedirect.com/). The specific keywords used are
“Spartina alterniflora”, “Invasive species”, “growth and spread”,
“ecological impact”, “biodiversity”, “potential applications”, “heavy
metal remediation”, “soil pollution remediation”, “soil carbon”,
“biomass from Spartina”, “ecological restoration”, “CO2 sequestration”,
“soil resilience”, “ecological implications”, “benthic macrofauna
habitat”, “control methods”, “controlled cultivation” etc and a combi­
nation thereof was also used to select research as well as review articles
focusing on Spartina alterniflora or its attributes. And the articles pub­
lished during the period from the early 1970s (wide use of SA had
started) to the year 2020 were considered. Furthermore, the articles
focusing on the application potential were given more focus as the aim
of the current review is to improve the application range of Spartina
alterniflora beyond its traditional invasive attributes.

Table 1
Distribution of Spartina alterniflora in different regions of the world
Continent/Country/ distribution Origin Invasiveness Remarks Reference
Region

Asia
China Present, Introduced Invasive Used to stabilize banks prior to the discovery of Li et al., 2020; Meng et al., 2020; Ding et al., 2008.
Widespread invasiveness
-Jiangsu Widespread Introduced Invasive
-Shanghai Widespread Introduced Invasive
-Zhejiang Widespread Introduced Invasive
India Present introduced invasive Shah and Badrinath, 1985

Europe
France Present Introduced Invasive Weber, 2017;
United Kingdom Present Introduced Invasive Maskell and Raybould, 2001

North America
Canada present Invasive Gillespie et al., 2007; Kilbride et al., 1995
-British Columbia Present, Introduced Invasive
Widespread
-Nova Scotia Present, Native Invasive
Widespread
United states Present
-California Present Introduced Invasive Kolton et al., 2020; Robertson et al., 2017;
-Connecticut present Native Invasive Lockwood et al., 2013; Więski and Pennings,
-Florida Present Native Invasive 2014.
-Georgia Present Native Invasive
-Louisiana Present Native Invasive
-Maine Present Native Invasive
-Maryland Present Native Invasive
-Massachusetts Present Native Invasive
-Mississippi Present Native Invasive
-New Hampshire Present Native Invasive
-North Carolina Present Native Invasive
-Oregon Present, Introduced Invasive
Widespread
-Rhode Island Present Native Invasive
-Texas Present Native Invasive
-Virginia Present Native Invasive
-Washington Present Introduced Invasive

Oceania
Australia Present Spartina introduced to Australia in late 1920s. Shimeta et al., 2016.
-South Australia Present, Introduced Invasive In southern Australia, introduced and used to
Widespread stabilise mud banks but, now considered invasive.
-Tasmania Present, Introduced Invasive
Widespread
-Victoria Present, Introduced Invasive
Widespread
New Zealand Present, Introduced Invasive Rai, 2015; Shaw and Gosling, 1997
Widespread

South America
Brazil Present Perillo et al., 2018; Costa et al., 2003
-Rio Grande do sul Present Introduced Invasive
Uruguay Present Introduced Invasive

Africa
South Africa Present Introduced Invasive Adams et al., 2016

2
S. Syed et al.
3. Implications of SA: environmental impact, biodiversity and
habitat loss.
The spread of SA was deliberate in regions such as China and acci­
dental in others like England as presented in Table 1. The invasion by SA
has major implications including altered wetland fertility, soil water
quality, hydrodynamics, sedimentary processes, soil composition and
nutrient accumulation (Yang et al., 2013, 2016c). The intensity of SA’s
impact fluctuates correlating location and habitat conditions. For
example, the invasive SA in the early stages has played a beneficial role
in protecting against wind and tidal erosion and improving the soil
stability of reservoirs (Currin, 2019). However, SA is currently regarded
as a weed in China, affecting the biodiversity of tidal areas, while in
some regions of the USA and Europe, it has caused great damage to
maricultural activities (Zhang et al., 2012; Li et al., 2020). Furthermore,
SA invasion sometimes decreases habitat structure and biodiversity but
also provide foraging area ultimately promoting some species, depend­
ing on the hydro-sediment dynamic conditions (Bruno, 2000; Chung,
2006; Dickie et al., 2014; Feng et al., 2017). Loss of unvegetated tidal
flats and the creation of stagnated surfaces are the most striking facets of
SA invasion, where the former hinder bird numbers (through habitat
reduction) and later boosts fisheries development (Meng et al., 2020).
SA invasion gradually but significantly modifies the coastal
ecosystem through its establishment timeline, resulting in a decreased
diversity and functionality of benthic macro-invertebrate communities
along SA invaded habitat. The initial establishment of SA possibly pro­
moted the macrofaunal biodiversity but subsequently decreased sub­
stantially (Zhao et al., 2014). SA’s ecological effects primarily depend on
the species’ traits making up the macrofauna of habitat and vary
seasonally which in turn varied markedly in different habitats (Quan
et al., 2016). The total organic carbon (TOC), total nitrogen (N) along
with the ratio of carbon to nitrogen (C/N) correlating with SA coverage,
stem density, and height are interesting to observe. Environmental
indices, benthic macrofauna and nutrient dispersion and availability
exhibited remarkable progression along with SA invasion chronology.
Additionally, SA invasion modifies habitat governing properties (vege­
tation development and nutrient accretion) impacting benthic macro­
faunal species sustainability in coastal marshlands (Ge et al., 2020).
The SA triggered alterations in the ecosystem affect the biological as
well as physicochemical characteristics affecting the successive food
chain and thereby interdependent species diversity, subsequently
shrinking, or eradicating such habitats (Gan et al., 2009; Ge et al., 2020).
SA triggers ecosystem changes in coastal and estuarine wetlands by
changing species speciation within the ecosystem by habitat modifica­
tions, competition and altered predator–prey interactions (Gan et al.,
2009; Wang et al., 2016b). A substantial shift in the food web is observed
in coastal ecosystems under aggressive propagation of SA, as it adversely
affects the diversity and density of local flora and fauna. Consequently,
impacting local bird communities, as seen in the case of shorebirds from
North America to Australia, where the shorebird numbers decreased
with the invasion of SA, as it reduces shorebird feeding grounds (Strong
and Ayres, 2013; Ju et al., 2017). The explosive progression of SA in
coastal marshlands resulted in a major habitat loss for the red-crowned
crane (Grus japonensis), alarmingly habitat shrank to just 22.36 km2 in
2015 from the 52.07 as recorded in 2000. Additionally, SA invasion
limits the movement of sediments forming an impenetrable barrier for
benthic organisms a food source of red-crowned cranes subsequently
influencing their density and habitat as reported by Wang et al., (2019).
4. Applications of Spartina alterniflora (SA)
In many regions of the world, SA is employed intentionally for the
restoration of coastal areas prone to degradation, tidal erosion, pollution
mostly due to human activities (Gao et al., 2014; Yang and Guo, 2018a).
SA is an ideal plant species, adapting synergistically (Fig. S1), to grow at
a wide range of salinity, pH and tolerant to many pollutants (heavy
3
Ecological Indicators 130 (2021) 108027
metals, hydrocarbons) and sea-water inundation for applications like
biomass production, phytoremediation, increasing soil resilience and
restoring carbon sinks in the coastal marshes and their sustenance
(Travis and Grace, 2010; Zuo et al., 2012).
4.1. Soil pollution remediation
SA exhibit a rigorously dynamic and intricate elemental utilization
pattern in established coastal marshlands. SA possesses resistance to
different stresses like heavy metal pollution, salinity, pH and organic
pollutants like PAHs and hydrocarbons (Lian et al., 2018; Prabakaran
et al., 2019). SA also possesses extraordinary heavy metal assimilation
potential via root uptake when grown in heavy metal containing soils.
Subsequently, decreasing heavy metal content in the proximate habitat
and remediating heavy metal contamination (Chen et al., 2017). Salla
et al., (2011) reported 65–140% higher content of Copper in SA leaves
and shoots than control. Pan et al. (2016), reported the Zn absorption
rate up to 500 µg⋅g− 1 in leaves of SA. Lian et al. (2018) found that SA
possesses a significant accumulation of Cr, Pb, Cu, Zn and Mn with
annual accumulation rates of 19.1, 84.2, 63.7, 442.6 and 774.7 mg m− 2,
respectively. More recently, Negrin et al., (2019) showed heavy metal
absorption was varied for different metals and Pb and Ni were more
efficiently absorbed than Zn, which too mostly in roots and rhizoids. In
SA, heavy metal absorption is throughout the plant body and involves
metal translocation from sediments via roots to above ground parts
where the heavy metal accumulates (Fig. 1). Additionally, after the
heavy metal absorption, the SA biomass can be burned in controlled
conditions to recover the absorbed heavy metals having commercial
worth (Chen et al., 2017). Consequently, SA becomes the biological
protector of marshlands from heavy metal contamination (Anjum et al.,
2014). The cropping and harvesting of the SA at appropriate times to
remediate contaminated soils will also overcome the release of the
heavy metals from the decomposition of the plant biomass (below and
aboveground plant parts) (Chai et al., 2014). Additionally, the SA is also
effective in remediating organic contaminants too. Lin and Mendelssohn
(2008) observed that SA has a high tolerance to crude oil (640 mg/g of
crude oil in the soil) and enhanced oil degradation both in surface and
subsurface contamination. Similarly, Hong et al., (2015) reported
enhanced degradation of PAHs through the plant-rhizosphere microbe
interaction unique to the SA. The phytoremediation of oil spills is
commonly attributed to the rapid growth of SA owing to higher
photosynthetic output and nutrient assimilation, influencing the rhizo­
sphere microflora which in turn degrade the organic contaminants
reducing its stress on the plant growth (Smith, 2013).
4.2. Soil resilience and functionality enhancing
Soil degradation is a complex phenomenon involving various factors
and measured by quantifying topsoil loss, organic matter, salinization,
acidification and deterioration of soil structure, the global as well as
local indices of soil health (Gonneea et al., 2019). Coastal ecosystems are
vital in maintaining ecological functions like sustaining coastal food
chains, preventing saltwater seepage into inland, and provide coastline
protection (Wang et al., 2016a). Ryan et al., (2007) and Yang et al.,
(2017) have reported coastal ecosystem revitalization approaches
employed by introducing nonnative or invasive species to increase
organic matter and sediment deposition preventing soil erosion. Wan
et al., (2009), Novy et al., (2010), Yuan et al., (2015) and Bilkovic and
Mitchell (2017), have established SA’s potential to synergistically affect
coastal ecosystem services and soil functions like improving the organic
carbon, soil organic carbon and soil carbon stock (Fig. 1). Feng et al.,
(2017), reported the increase of soil carbon stocks (surface soil, 0–10
cm) in SA invaded mangrove mudflats suggestive of its carbon seques­
tration ability. According to Wang et al., (2016b), the introduction of SA
into degraded salt marshes, mangroves, wetlands enhanced the soil
compaction, reduced soil erosion by tidal waves, stress of saltwater and
S. Syed et al.
Fig. 1. S. alterniflora: effects and benefits in controlled cultivation and natural/non-native habitats.
enhanced the soil productivity gradually as SA’s invasion is character­
ized by greater carbon accretion, improved nutrient percolation and
cycling. Nonetheless, SA is not considered the first option for resilience
restoration or enhancement because its adaptability can be a threat as
detailed in the management section below, and it should be considered
for cultivation in controlled environments or habitats (where less
adaptive species are stressed) where continuous monitoring is available
or possible, such as sludge treatment plants, abandoned mine drains,
and so on where remediation of the contaminants is of paramount
significance.
4.3. CO2 sequestration and carbon sink development
By assimilating atmospheric C and conducting different ecosystem
services such as nutrient cycling, coastal wetlands play a critical role in
global climate mitigation and function as a habitat for various organisms
(Nellemann et al., 2009; Adhikari and Hartemink, 2016). Nevertheless,
human activities such as urban expansion and land reclamation from the
sea for commercial and recreational purposes have severely accelerated
the degradation of coastal wetlands and subsequent environmental
changes. Furthermore, deprivation of coastal marshlands fuels putre­
faction of plant organic debris resulting in significant CO2 emissions into
the atmosphere, reversing the C sequestration (Chmura et al., 2003;
Drake et al., 2015). Restoration projects with native species to revitalize
ecosystems can be a challenge, as the native species are prone to local
ecological restrains and detrimentally lack aggressive colonization and
growth traits (Matthews and Spyreas, 2010). Therefore, to increase
coastal resilience further, the introduction of SA is proposed in some
special situations (Luo et al., 2015; Currin, 2019). And Davis et al.,
(2015) reported substantial CO2 sequestration (58 to 283 g C m− 2 yr− 1)
in Californian shorelines, USA (SA habitat) in the last three decades.
Yang and Guo (2018b) also found a similar but significant increase in
soil organic carbon content in the topsoil (1 m) with SA introduction in
Jiangsu coastal marshes (China). Similarly, Yuan et al., (2015) found SA
populated marshes have remarkably higher CO2 assimilation potential
(3.16 Mg C ha− 1 yr− 1) a two to eight-fold increase in comparison to
similar but native species populated marshes. The invasive SA has
several ecophysiological properties, like, extended seasonal growth,
superior photosynthetic rate, higher root growth, and low internal Ni­
trogen contents compared with native species, enhance CO2 fixation into
plant biomass and subsequently into soils (Zhang et al., 2017). Many
reports (Zhou et al., 2015; Zhang et al., 2018) have substantiated the
4
Ecological Indicators 130 (2021) 108027
restoration of coastal wetlands, along with an increase in CO2 seques­
tration and soil organic carbon in the topsoil (0–30 cm) because of SA
introduction/or invasion (Artigas et al., 2015). SA invaded ecosystems
exhibit considerably increased C and N deposition than with other
native flora. However, these reports are of the intentional introduction
of SA and its effects on soil characteristics; based on the most recent
assessment, it is believed that more extensive studies are required before
it can be widely used.
4.4. Biomass for multiple purposes
Biofuel from plant-based lignocellulosic biomass serves as an envi­
ronmentally friendly alternative, possibly replacing fossil fuel-based
gasoline, as it is a renewable source of plant biomass produced
through photosynthesis (Li and Qiu, 2012). Hence, utilization of halo­
phytes such as SA having more than 1.0 photosynthesis to respiration
ratio and an average net primary production of 1–1.5 kg dry mass m− 2 is
an ideal choice. Additionally, halophytes grow in saline soils and do not
compete with agricultural soils used for food crop production for
humans and animals (Hongguang et al., 2007; Li and Qiu, 2012). Yang
et al., (2017), found that SA generates a significant quantity of lingo-
cellulose in their biomass. Qin et al., 2016 reported, SA (in jointing,
heading, and flowering stages) was an excellent fodder supplement for
dairy cows (at up to 25% of the diet) as it has higher lignocellulose
content and did not alter the rumen fermentation. Furthermore, SA can
be cultivated with sewage water consequently removing the contami­
nants (heavy metals, organic compounds and hydrocarbons) (Liu and
Tian, 2002) and also as a green fertilizer (whole biomass) to improve
organic carbon in soils (Wang et al., 1996). The SA is a huge biomass
resource and biological conversion (i.e., anaerobic digestion) into
methanogenic fuel (biogas) serves both green energy goals (Shiguan
et al., 2008; Lu and Zhang, 2013; Hao et al., 2018). The biomass from SA
can be burnt to be utilized as a carbon component in organic fertilizer
formulations or used as such in agricultural soils as green manure to
increase soil carbon content.
5. Management of SA-recommendations and risks
SA, as described earlier has tremendous application potential in
multiple fields yet, it is considered harmful or invasive owing to its
negative effects. There has been great development both in the usage
and processing of SA, including traditional (biomass, biochar, etc.) and
S. Syed et al. Ecological Indicators 130 (2021) 108027

Fig. 2. S. alterniflora biomass based products and the processes involved/applied in their production.

sophisticated processing (biofuel). Many reports of SA’s potential

application and deliberate introduction originate from areas with low Table 2
Spartina alterniflora invasions and effects.*
bioresources (less fertile soils, low forest cover, unfavorable environ­
mental conditions) (Duns, 2020). It is of consensus SA cultivation is Negative effects Positive effects
beneficial and even economically feasible if taken up in areas/soils Habitat loss (for macro marine fauna) Coastal soil protection
lacking native plant cover and with significant alkalinity and salinity Biodiversity loss Rejuvenation of wetlands/marshlands
(Wan et al., 2009; Duns, 2020). Using sewage water or sludge to culti­ Native plant diversity and density Reduces coastal erosion (via soil
depletion (due to habitat stabilization)
vate SA in nurseries is another effective approach (Liu and Tian, 2002), modification)
which tackles pollution removal and biomass generation at once. Lu Eradicates native plant species (Niche Generates biomass for fuel/energy/fodder
et al., (2020) suggested utilizing the SA biomass in a seven-step colonisation)
ecological engineering system as a biomass feedstock. To resolve CO2 sequestration (improving soil carbon
sink)
certain environmental issues (like heavy metal leaching, degradation of
Functions as a bioremediation agent (for
sludge, shoreline erosion), the SA can be effectively utilized in closed or heavy metal and organic pollutants)
controlled environments (sewage/sludge treating plants where the *
Chen et al., 2014; Duns, 2020.
sludge itself can be used as cultivation soil, abandoned mine drains,
shores of artificial islands) due to its rare and exploitable traits.
Many of the investigations also note how invasive SA is, thus many affected by tidal waves hence, effective as a natural barrier for coastal
control measures are investigated. The most common practice is burning line protection, also serving as wave impact disperser. SA is also resilient
the SA stands which becomes a good fertilizer to the soil, but this leads to to the presence of contaminants both organic (polyaromatic hydrocar­
potential air pollution. Xie et al., (2019) have reviewed different me­ bons, hydrocarbons) and inorganic (heavy metals) thus effective in
chanical and chemical control methods to manage SA spread and in­ phytoremediation projects of heavy metal and organic polluted soils.
vasion, wherein tilling/mowing is the most effective mechanical SA’s resilience is handy in restoring degraded soils of coastal areas due
approach to control SA in and around water bodies like riverbanks. In to human activities like dredging for transport or fossil fuel explorations.
the case of intentional cultivation of SA, biomass should be regularly Thus, SA is a plant species with potential multiple applications, of
harvested to manage and contain the growth effectively. Herbicides such course, we need to strictly monitor and develop its downstream pro­
as Haloxyfop-R-methyl and Imazapyr were also found to be effective (in cessing and utilization technology.
the case of relatively sporadic SA patches). Along with the above
methods, biological approaches like adding much faster-growing Declaration of Competing Interest
mangrove plants (Sonneratia apetala) have been reported (Chen et al.,
2014). Despite the positive and negative effects (Table 2) of SA, the The authors declare that they have no known competing financial
positive effects in many areas outweigh the negative ones when utilized interests or personal relationships that could have appeared to influence
under control. the work reported in this paper.

6. Conclusion
Acknowledgements
SA is a saltmarsh perennial grass having multiple advantages like
This work was jointly supported by the Jiangsu Provincial Marine
rapid growth, propagation, resistance to high salinity and wide pH
Science and Technology Innovation Special Project (grant number
range, resource acquisition and utilization efficiency when compared to
HY2019-3) and the Introduction Plan for High-Level Foreign Experts
the native plant species. SA can negatively impact the environment by
(grant number G2019021434).
disturbing the local biodiversity and gradually dominates and eliminates
the local native plant species resulting in severe changes in the
ecosystem attributes along with the soil modifications like soil organic Authors’ contributions
carbon, soil carbon stocks and soil physicochemical properties. How­
ever, not all SA traits are devastating if used constructively. The rapid SS Curated related data and prepared original draft. BL Conceptu­
growth and biomass accumulation even under lower temperatures and a alized, reviewed and edited the draft manuscript. MX, ZW and CY
wide range of pH and salinities make it an ideal choice for soil stabili­ participated in discussion and revisions. All others: wrote the
zation and erosion prevention particularly in lowlands and inland areas manuscript.

5
S. Syed et al.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.ecolind.2021.108027.
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