Introduction
Alien plants can have detrimental impacts
on biodiversity, including the decline and local extinction of indigenous
species (Vilà et al. 2010; Pyšek et al. 2012a; Pérez et al. 2022). Assessing their potential impacts is important to decision making
and the application of successful management strategies (van Kleunen et al. 2018; Pyšek et al. 2020). The Poaceae family constitutes a high portion of the alien flora,
for example in Japan (372 out of 1553), China (205 out of 14710), the Czech
Republic (152 out of 1378), Italy (163 out of 1200), Belgium (118 out of 2500)
(Pyšek et al. 2012b; Galasso et al. 2018; Lin and Ma 2022; Verloove 2023) and Iran (34 out of 311) (Sohrabi et al. 2023b). Generally,
grasses possess traits that facilitate long-distance dispersal, establishment,
colonization, and transformation of novel environments, and resilience to
disturbance and drought (Linder
et al. 2017; Bastida et al. 2021; Leal et al. 2022).
Alien grasses with high seed production, long seed viability, fast seed
germination, vegetative reproduction and low palatability have high potential
to invade disturbed sites (Medeiros
and Focht 2007). They have spread over large areas
of natural ecosystems displacing native species owing to their aggressiveness
and highly competitive ability (Damasceno and Fidelis 2020).
Furthermore, alien grasses are an important threat to biodiversity conservation
due to their resilience to global warming (some are C4 species) and potential
for increasing fire risk (D’Antonio and Vitousek 1992).
One way to quantify the potential
environmental impacts of alien plants is a standard approach for ranking alien
species based on the damage they may cause in recipient areas (Nentwig et al. 2016; Blackburn et al.
2014; Vilà
et al. 2019; Kumschick et al. 2020).The IUCN
Environmental Impact Classification of Alien Taxa (EICAT-IUCN) is a scheme for
classifying alien taxa in terms of the magnitude of their environmental impacts
and the mechanism involved to prevent or limit their negative consequences (www.iucn.org).
EICAT-IUCN proposes a global assessment where the category assigned to a
species is based on the highest impact ever recorded or its highest current
impact, observed anywhere (IUCN
2020a and b; Kumschick et al. 2024).
Grasses have also large economic impacts on
agricultural lands (Milton 2004; Goggin et al. 2012; Follak and Essl
2012). For example, the rigid ryegrass (Lolium
rigidum) infestation in southeastern Australia costs an estimated $93
million (AUD) per annum in grain loss and considerably more in control costs (Bajwa et al. 2021). Johnson grass (Sorghum halepense) is classified as an
important weed in over 53 countries occurring in 30 different crops (Follak and Essl 2012; Peerzada et al.
2017), causing substantial crop yield loss (Klein and Smith 2021). There are 311 alien plants in Iran and
the number is increasing every year (Sohrabi et al. 2023b; Sohrabi and
Pagad 2024). A large proportion of alien plants in
Iran are grasses (Sohrabi et al. 2023b), but information
on their potential impact on both native species and crops is lacking. For
example, Rottboellia cochinchinensis is present in the Khozestan
province, (Dinarvnd
and Ale-Bakhit 2013; Sohrabi et al. 2023a). This species is
known worldwide for invading crops and disturbed habitats in tropical and
subtropical regions (Funez et al.
2016). However, we do not know its potential impact
in Iran and how it compares to that of other alien grasses present in the
country. Overall, there is a lack on the impact of grass invasions in Iran. To
contribute to filling this knowledge gap, the aim of this paper is to
categorize alien grasses in Iran based on their environmental impacts by using
the EICAT-IUCN classification, and furthermore, to assess potential impacts on
agricultural production. We assumed that the life cycle, photosynthetic
pathway, invasion status and distribution of the grass species in Iran´s
ecological zones contribute to the severity of the impacts. We also discuss
management options for alien grasses for both natural ecosystems and arable
lands.
Methods
The selection of plants
Thirty-four alien grasses present in Iran
were classified as casual, naturalized, and invasive based on their stage in
the invasion process (Richardson
et al. 2000; Sohrabi et al. 2023b). The potential
environmental impacts of these species were assessed following the EICAT-IUCN
guidelines, which consider 12 mechanisms of impact (IUCN 2020b; Volery et al. 2020). The severity of the
impacts was categorized as minimal concern (MC), minor (MN), moderate (MO),
major (MR), or massive (MV).
We obtained information on the
environmental and agricultural impacts of the study of alien grasses from
primary scientific literature related to their global alien range. Data on
their distribution in Iran, habitat type, introduction pathways, and
photosynthetic pathways were sourced from scientific journal articles, reports,
books, book chapters (see references in Appendix), and online databases such
as WIKTROP
(Weed Identification and Knowledge in the Tropical and
Mediterranean areas). The distribution of all alien grasses in Iran was
assigned to one of the four ecological zones of the country (Fig. 1):
(a) Hyrcanian zone: located in the north, with a temperate climate, it extends
along the southern coast of the Caspian Sea and the northern part of the
country; (b) Khalij-o-Omani zone: located in the south, covering an area of
2 130 000 ha, it is characterized by a sub-equatorial climate (Heshmati 2012);
(c) Zagross zone: situated in the west, this mountainous region covers
approximately 4 749 000 ha, characterized by a semi-arid climate and
temperate winters; and (d) Iran-o-Touranian zone: dominated by the central
Iranian plateau, this zone covers 1 648 000 km² and features a wide
variety of climates, surrounded by mountain ranges.
Statistical analysis
A contingency table was used to analyze
associations between EICAT-IUCN impact scores (harmful (MV, MR and Mo) and
non-harmful (MN, MC and DD)) and variables such as species status (casual and
naturalized invasive), life cycle (annual and perennial), and photosynthetic
pathway (C3 and C4). Fisher's exact test was applied to identify significant
relationships at 0.05 level. Statistical analyses were performed using R 4.3.0
(R Core Team 2022), and the packages ggplot2 (Wickham 2016) and ggpubr (Kassambara 2025) were used for data visualization.
Results
General characteristics of alien
grasses
In Iran, there are 22 naturalized, 11
casuals, and one invasive alien grass species. Most of
these grasses utilize the C4 photosynthetic pathway (27 species). The C4 pathway
is strongly associated with successful naturalization in new areas. Among the
22 naturalized grasses, only Five species utilize the C3 pathway. By longevity,
there is a similar number of perennial species (18) and annual species (17).
Notably, only two of the 17 annual species use the C3 photosynthetic pathway. The Poaceae family, due to their seed size,
is easily transported through different introduction
pathways (Table 1). Approximately 19 species originate from tropical and temperate
Asia, while seven are native to Africa and North America, respectively. In
Iran, contamination (e.g. seed contamination) was identified as the primary
pathway (56%) for the introduction of alien grasses. In most cases, the
introduction in Iran has been accidental (14%), with a few exceptions involving
deliberate releases, such as Bambusa vulgaris and Chrysopogon
zizanioides.
Impact assessment based on
EICAT-IUCN
Based on EICAT-IUCN framework, the most
common impact scores for alien grasses in Iran were Minor (MN) and Major (MR);
with 13 and seven species assigned to these categories, respectively. Seven
species were classified as Data Deficient (DD). Among the 22 naturalized
species, ten were rated MN, while five were categorized as MR (Table 1; Fig. 2). We
identified the species Rottboellia cochinchinensis, Paspalum
distichum, and Microstegium vimineum, as having Massive (MV)
impacts. Of these, only R. cochinchinensis is invasive in Iran, while
the others are naturalized. Additionally, Cortaderia jubata and Phyllostachys
reticulata, which are casual grasses, exhibited major impacts (Fig. 2).
Competition (23.5%) and transmission of disease (20%) were the most frequently
reported mechanisms of impact for alien grasses. Structural impacts on the
ecosystems, hybridization, and toxicity/poisoning were also common (Table 1).
Fisher's exact test revealed that there is no evidence of an association
between EICAT-IUCN scores and invasive status (p-value: 1, odds
ratio=1.03), while the association between EICAT-IUCN scores and life cycle is
considered to be statistically significant (p-value= 0.0.032, odds
ratio=0.036). Various EICAT-IUCN scores had shown insufficient evidence to
reject the null hypothesis and that there is no evidence of an association with
photosynthetic pathways (p-value = 0.74, odds ratio=0.65), although all
three grasses with MV score were C4 species.
Distribution of alien grasses in
Iran
The proportions of alien grasses in Iran's
ecological zones were 61.7% in the Hircanian, 41.1% in the Zagros, 26.4% in the
Iran-o-Turonian, and 8.8% in the Kh-O-Omanian zone. The Hircanian ecological
zone had the highest proportion of grasses with MN (47.6%) and MR (28.5%)
impact scores. In contrast, 50% of the grasses in the Zagros zone had a Minor
(MN) impact score, whereas no grasses with this score were recorded in the
Kh-O-Omanian zone. The proportion of grasses classified as Data Deficient (DD)
was higher in the Iran-o-Turonian zone than in others ecological zones (Fig. 1).
Figure 1. The distribution of 34 alien grasses in Iran along with their
EICAT-IUCN scores. The graphs display the number of species classified under
each EICAT category (DD - Data deficient, MC - Minimal concern, MN – Minor, MO
– Moderate, MR – Major and MV – Massive).
Figura 1. Distribución
de 34 gramíneas invasoras en Irán junto con sus puntuaciones EICAT-UICN. Los
gráficos muestran el número de especies clasificadas en cada categoría EICAT
(DD: datos insuficientes; MC: preocupación mínima; MN: menor; MO: moderada; MR:
mayor y MV: masiva).
Table 1. List of 34 alien grass in Iran and their main characteristics and
environmental impacts.
Tabla 1. Lista
de 34 especies de gramíneas invasoras en Irán y sus principales características
e impactos ambientales.
Figure 2. Invasion status and EICAT-IUCN impact score of 34 alien
grasses in Iran. EICAT-IUCN ranking included data deficiency (DD), minimal
(MC), minor (MN), moderate (MO), major (MR) or massive (MV) impact. Two cases
are subspecies: Triticum turgidum subsp. durum and Diplachne
fusca subsp. Uninervia.
Figura 2. Estado de invasión y puntuación de
impacto EICAT-UICN de 34 gramíneas invasoras en Irán. La clasificación
EICAT-UICN incluyó datos insuficientes (DD), impacto mínimo (MC), menor (MN),
moderado (MO), mayor (MR) o masivo (MV). Dos casos son subespecies: Triticum
turgidum subsp. durum y Diplachne fusca subsp. Uninervia.
Impact
of alien grasses on agricultural areas
Twenty-three of the 34 alien grass species
identified in Iran are invading agricultural areas (Table 2). The majority of these species were observed in
rice fields, comprising 14 species. Six species were observed in maize and
sugarcane fields, while only five species were recorded in orchards and
vineyards. The primary mechanisms of impact include competition with crops and
serving as hosts for pests. Furthermore, some species exhibited herbicide
resistance, allelopathic potential, and toxicity to animals (Table 2). We recorded thirteen alien weedy
grass species in northern Iran, six in the southern region, with the remaining
species distributed across central and eastern Iran (Table 2).
Table 2. Alien weedy grasses in Iran invading agricultural areas and their
characteristics. See full Literature in Appendix.
Tabla 2. Las gramíneas invasoras de zonas agrícolas en Irán y sus
características. Véase la bibliografía completa en el Apéndice.
Discussion
General characteristics of alien
grasses
Alien grasses contribute 13%, 9%, and 7.6%
to the total number of naturalized, casual, and invasive alien plants in Iran,
respectively (Sohrabi et al.
2023b). The proportion of naturalized grasses in
Iran is comparable to findings from other studies in China (Lin and Ma 2022)
and worldwide (Pertierra et
al. 2023). The faster
growth rates and larger sizes of C4 species enhance their establishment success
compared to C3 species (Jia et al.
2016). Spartina alterniflora invasion, a
perennial C4 grass in China, has been linked to its rapid growth and ability to
stabilize tidal flats (Cheng et
al. 2006). The worse invasive alien grasses have C4
photosynthetic pathway (GISD 2013). For example, Cenchrus spinifex is one of the foremost
invasive C4 grass in Hungary (Botta-Dukát and Balogh 2008). Numerous
studies have highlighted the successful establishments of the perennial grasses
from temperate-tropical regions due to their higher competitive ability in warm
climates (Hacker and
Dethier 2006; Lopes et al. 2023). In our study, most
C3 grasses were perennial. The rapid spread of these species, whether by seed
or vegetative propagation; facilitates their invasion into disturbed areas,
such as the Cerrado in Brazil (Zenni
et al. 2020). In addition, Seed contamination as the primary pathway of new introduction is
consistent with findings in Europe (Poschlod et al. 2009).
Impact assessment based on
EICAT-IUCN
Among different mechanisms competition is
well documented, while other mechanisms, such as hybridization and biofouling,
are more challenging to assess due to indirect evidence and complex data
interpretation (Foxcroft et al.
2019). For example, detecting hybridization impacts
requires molecular research, which is costly and more time-consuming compared
to evaluating allelopathy or competition (Gioria and Osborne 2014; Kalisz et al. 2021). Phenolic acids are the main identified allelochemicals (67%) in
the Poaceae family (Favaretto
et al. 2018). In addition to irritating trichomes
on the leaf sheaths of R. cochinchinensis, its phytotoxic effects
on adjacent plants specie cause reduced growth of seedlings (Meksawat and Pornprom 2010). Changes in disturbance regimes and natural succession have been
reported for grasses like Chrysopogon zizanioides, due to their root
system in upper horizons of the soil (Eab et al. 2015; Freschet and Roumet 2017; Badhon et al. 2021). The transmission of disease is another significant mechanism, and
the importance of grasses species in transmitting endophytic fungi is well
known (Lapierre and
Signoret 2004; Yuan et al. 2010). For example, Bipolaris
gigantea, a foliar fungal pathogen in alien Microstegium
populations, had significant impact on biomass responses of three native North
American grass species (Kendig et
al. 2021); and Xylella fastidiosa, a
widespread bacterial pathogen of olives, almond and citrus, can be transmitted
indirectly (xylem-feeding insects) by alien grasses (Najberek et al. 2022).
Distribution of alien grasses in
Iran
The highest number of naturalized and
invasive plants in the Hircanian ecological zone is attributed to its favorable
environmental conditions and high population density (Sohrabi et al. 2023a). Biodiversity hotspots with high environmental and economic value
are at the greatest risk of biological invasions and should therefore be
prioritized for invasive species management (Li et al. 2016; Yang et al. 2023).
The Caspian forests, located in the Hircanan zone and recognized as a UNESCO
World National Heritage site (UNESCO
2019), are a critical area in Iran that requires
attention as a potential hotspot for biological invasion. Rottboellia
cochinchinensis was found in the Zagros zone and it is the only invasive
grass species recorded in Iran. With a high seed production potential of over
3000 seeds per plant, immediate action is essential to curb its rapid spread (Dinarvnd and Ale-Bakhit 2013). Understanding the ecological zones most vulnerable to invasion is
crucial for predicting the threats posed by alien plant invasions (Sohrabi and Gherekhloo 2015). Effective management strategies can be better formulated and
implemented by combining distribution data with environmental impact scores (Panda et al. 2017; Sohrabi et al.
2023a). The agricultural lands and ruderal areas
are frequently cited as the primary habitats invaded by alien plants (Gaertner et al. 2017; Sohrabi et al.
2023b; Potgieter et al. 2024). However, it is
worth highlighting that many cultivated alien grasses have significantly
contributed to agricultural production and economic development (Randriamampianina et al. 2024). For instance, Sorghum bicolor and Cortaderia jubata
have been introduced to Iran as crops and ornamental plants, respectively.
Globally, Echinochloa oryzoides and Setaria italica are among the
most commonly encountered weedy grass species. Their frequent occurrence can be
attributed to their remarkable adaptability to various environmental stressors,
such as flooding, drought, and salinity (Mohammadvand et al. 2012; Lapuimakuni et al. 2018; Kaya-Altop et al.
2019; Nisa and Jadid 2021). Among the most
damaging species in agricultural areas are P. repens, P. distichum and L. chinensis, which pose a significant threat to crop production, potentially
causing yield losses of up to 60% (Hossain et al. 2001; Kojima et al. 2005; Hayyat et al. 2023). Paspalum distichum infestation has
been reported in perennial crops throughout Spain, where it threatens olive and
vineyard production (Costa 1997). The introduction of P. distichum for pasture purposes has
increased its potential to invade, particularly arid regions, and its
impact extends to river flora and the integrity of river systems in Portuguese
floodplains (Bernez et al. 2005; Driscoll et al.
2014). Similarly, the negative impact of P.
repens has been extensively documented in agricultural areas and native
plant communities in the USA, where its management in flood control systems for
Florida’s citrus groves alone costs $2 million annually (Cuda et al. 2007;
Langeland and Burks 2022). Leptochloa chinensis is a serious weed in the
direct-seeded rice crop, causing substantial yield reductions at high densities
due to its competitive ability, which is attributed to its C4 photosynthesis
pathway and increased biomass production rate (Benvenuti et al. 2003; Sage et al. 2012;
Deng et al. 2021). In total, there are 25 predominant weedy grass species found
across various agricultural lands in Iran, of which seven species, accounting
for 28%, are alien (Mozafarian
2021; Sohrabi et al. 2023b, 2024).
Therefore, a better understanding of the portion of alien grasses in the weed
flora and their impact can benefit and improve the ecological sustainability of
crop production systems.
Management of alien grasses
Certain interventions can reduce the
propagule pressure and the likelihood of establishment and spread of newly
introduced alien grasses (Perrings
et al. 2005; Simberloff 2009; Siddiqui et al. 2022). Strategies such as limiting contamination vectors, monitoring
pathways for target grasses, controlling contaminants, and adopting long-term
approaches have been suggested as the most effective methods for managing
invasive grasses in specific areas (Pyšek et al. 2011; Gentili et al. 2021; Rudell et al. 2023). However, we highlight that the most accurate management options
will depend on the infested habitat (natural areas or agricultural lands) and
the life cycle of the species (annual or perennial). In natural areas, the most
effective control methods often involve an integrated approach that include
burning, mowing, hand clearing, and herbicides application (Marushia and Allen 2011). In agricultural areas, effective control typically involves the
use of certified seeds, proper land preparation, stale seedbeds to promote weed
germination prior to seeding, careful crop and water management, herbicides,
and crop rotation (Milton 2004; Zand et al. 2017; Khasraw et al. 2023; Gherekhloo et
al. 2020; Siddiqui et al. 2022).
The application of herbicides is a widely
used strategy for managing invasive plant species, particularly in agricultural
systems. However, the continuous and intensive use of these chemicals has
resulted in the emergence of herbicide-resistant weed populations, posing
significant challenges to crop productivity and sustainability (Hassanpour-bourkheili et al.
2024; Heap
2024). To achieve complete control of P. repens
in naturally infested fields often requires multiple applications of methyl
sulfanilylcarbamate (a dihydropteroate synthase inhibitor), as plowing-induced
rhizome fragmentation disperses the weed across various soil layers,
complicating eradication (Hossain
2001). In paddy fields in Rasht, Iran,
bensulfuron-methyl, an acetolactate synthase inhibitor, has demonstrated higher
efficacy in controlling Echinochloa oryzoides (Pouramir and Yaghoubi 2020). In USA rice fields, Leptochloa chinensis and P.
distichum have evolved herbicide resistance due to the repeated application
of a single herbicide or herbicides group (Chauhan et al. 2012). To manage P.
distichum and mitigate the risk of glyphosate resistance, the integration
of herbicides with different modes of action has been recommended (Alcantara et al. 2016).
As a sustainable alternative to herbicides,
biological control methods have been suggested, including the use of grazing
animals such as rabbits, goats, sheep, and ducks, as well as cover crops and
intercropping. These approaches have been explored for managing invasive
species like Paspalum repens, P. distichum, and Rottboellia
exaltata in turmeric fields (Prabhakaran Nair 2013).
Relying on a single control tactic may be
insufficient for effectively managing invasive plant species, particularly
given the high reproductive potential and dispersal ability of certain species.
In this context, Integrated Weed Management (IWM) presents a promising strategy
for controlling and mitigating the impact of invasive species in both
agricultural and natural ecosystems (Gonzalez-Andujar 2023).
Conclusion
This study, based on the EICAT-IUCN
assessment, highlights the significant impact potential of several alien grass
species in Iran, including Rottboellia cochinchinensis, Paspalum
distichum and Microstegium vimineum. These species primarily affect
ecosystems through competition, disease transmission, and structural
alterations. The prioritization of management efforts should be guided by these
findings, particularly for P. distichum and Panicum repens which
pose substantial threats to biodiversity and agricultural productivity. The
highest number and impacts of alien grasses in Iran were found in the Hircanian
zone, followed by the Iran-o-Turanian zone.
Notably, a correlation between EICAT-IUCN
score and life cycle underscores the critical role of perennial and species in
driving environmental impact. Given the prevailing warming conditions, it
is crucial to focus on the introduction and management of alien C4 perennial
grasses in Iran. Effective management of strategies will depend on the life
cycle of the species and the nature of the invaded habitat, though the most
successful approaches typically involve a combination of methods.
Data Availability
More information
is available in Appendix and raw data will be presented upon reasonable
request to the corresponding author.
Financing, required permits, potential
conflicts of interest and acknowledgments
Gorgan University of Agricultural Sciences
and Natural Resources (GUASNR), Iran supported this research (project no.
GAU-03-508-20). We would like to thank Sabrina Kumschick for her valuable
suggestions. The authors are grateful to anonymous reviewers for their valuable
comments that greatly enhanced this work.
The authors
declare that they have no conflicts of interest.
Authors' contribution
SS, JG, JLG-A, MV, designed the study; SS
collected the data; SS analyzed the data; SS led the writing of the manuscript,
and all authors contributed critically to the drafts and gave final approval
for publication.
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Appendix / Anexo
Literature
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