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\begin{document}

\title{Weed management for the land-scape scale restoration of global temperate
grasslands: a review.}



\author[1]{Talia Humphries}%
\author[1]{Singarayer Florentine}%
\author[1]{Kim Dowling}%
\author[1]{Chris Turville}%
\author[2]{Steve Sinclair}%
\affil[1]{Federation University Australia}%
\affil[2]{Victoria Department of Environment Land Water and Planning}%


\vspace{-1em}



  
  \date{March 15, 2025}


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\selectlanguage{english}
\begin{abstract}
Globally, temperate grasslands have been significantly altered and
subsequently degraded as a result of increased human population,
urbanisation, and agriculture. Weeds now dominate most of these
ecosystems, resulting in the loss of ecosystem services, reduced
carrying capacity for farmers, and loss of habitat for native animals.
This paper reviews the literature of temperate grassland restoration
efforts from across the globe, and observes what techniques and
combinations have been used successfully and unsuccessfully to reduce
weed dominance and promote native recruitment and establishment. The
findings of this review clarify that weed management should be ongoing
in all projects, while optimal revegetation methods and grazing regimes
are specific to site location and study scope. There is a need for an
increase in long-term monitoring of restoration projects in order to
make assumptions with greater confidence.%
\end{abstract}%



\sloppy


\textbf{Introduction}

Temperate grasslands once covered almost 9 million
km\textsuperscript{2}, which is equivalent to about 8\% of the earth's
surface (IUCN, 2013). They include the Prairies of North America, the
Pampas of South America, the South African Veldts, the Tussock
grasslands of Australia and New Zealand, and the Steppes of Eurasia
(Table 1). These biomes are often species rich (Faber-Langendoen \&
Josse, 2010), providing natural habitat for many plants, animals and
soil biota. In addition, these grasslands offer invaluable ecosystem
services such as high-quality forage for herbivores (Boval \& Dixon,
2012), harbour pollinators for crops and native species (Bendel \emph{et
al} ., 2019), provide significant levels of carbon sequestration
(Eze\emph{et al} ., 2018), and are places for many recreational and
cultural activities (Gomez-Limon \& de Lucio, 1995). They also afford
many other environment stabilizing services, such as soil erosion
control and mitigation of flood waters (Sankaran \& Anderson, 2009).

Given the significance and contribution of this ecosystem, they're
currently one of the most altered ecosystems in the world
(Suttie\emph{et al} ., 2005), warranting immediate action to restore
these beneficial services. Estimates suggest that 70\% of these
ecosystems were altered or degraded before 1950, and a further 14\% by
1990 (Hassan\emph{et al} ., 2005). This decline in ecosystem health is
directly attributed to rapid population growth and subsequent urban
expansion (Williams \emph{et al} ., 2005), as well as the concomitant
conversion of these fertile ecological systems into sites for
agriculture, particularly for cropping systems and livestock grazing
(Martin \emph{et al} ., 2005; Prober \emph{et al} ., 2005; Bartolome
\emph{et al} ., 2009; Sankaran \& Anderson, 2009). While increased
protection of the remaining intact system is critical, it is not enough
to ensure the future resilience and functionality of these systems.
Therefore, this paper reviews restoration methods, both active and
passive, that reduce invasive plant biomass within global degraded
temperate grasslands to promote the return of natives and subsequent
ecosystem functionality at a landscape scale.

\textbf{Ecological Restoration Methods}

Ecosystem degradation is the movement of a high functioning and healthy
ecosystem into an altered state whereby ecosystem functions are reduced
or lost as a result of a single or multiple disturbance event, usually
due to human actions (Suding \& Hobbs, 2009). Healthy ecosystems are
often resilient enough to withstand moderate disturbances, and these
events can be important for maintain biodiversity and healthy
ecosystems. When the intensity and/or frequency of these disturbance
events change, an ecosystem can undergo hysteresis and pass through
irreversible degradation thresholds (Suding \& Hobbs, 2009). An example
of this was observed in the grasslands of California where the combined
factors of weed invasion by Mediterranean species and altered grazing
regimes from the introduction of livestock as well as multiple severe
drought seasons resulted in the sever degradation of this ecosystem,
pushing it into an alternative stable state (Bartolome \emph{et al} .,
2009). Different levels of degradation can influence the scale of
restoration required, depending on if the biotic factors (weed
invasion), abiotic factors (drought or altered fire regimes) or a
combination of both are affected.

Throughout the world, grasslands are currently subjected to multiple
degrading pressures. The most common of these pressures include; habitat
fragmentation, which has been observed in Australia (Prober \emph{et al}
., 2005), New Zealand (Standish \emph{et al} ., 2009) and Europe (Kiehl,
2010), altered grazing pressures in the USA (Martin \emph{et al} .,
2005; Bartolome \emph{et al} ., 2009), Africa ( Sankaran \& Anderson,
2009) and Australia (Prober \emph{et al} ., 2005), desertification and
bush encroachment have been observed in Africa (Sankaran \& Anderson,
2009), and climate change has been linked to the gradual decline in
grasslands throughout China (Zha \& Gao, 2011). These degrading
pressures often act to promote the invasion of exotic plants, which in
turn create positive feedback loops that maintain the degraded altered
state. The alternative state theory explains how internal disturbances
and external shocks lead to positive feedback loops which promote a
stable degraded state (Chisholm \emph{et al} ., 2015). In this state,
the degrading factors have altered the environment to promote their own
development, as observed in south east Australian grasslands where
annual exotic grasses outcompete the perennial native grass, Themeda by
developing new positive feedback loops that increases soil nitrogen, and
unless the available soil nitrogen levels are reduced, the invasive
species will maintain its competitive edge (Prober \emph{et al} .,
2009). Further, cross-facilitation of invasive plant feedback loops has
been identified by observing \emph{Agropyron cristatum} in northern USA,
which alters native soil biota and thus, soil dynamics (Jordan \emph{et
al} ., 2008). This reduces the competitiveness of the native vegetation,
promoting niche availability for the invasive\emph{Bromus inermis}
(Jordan \emph{et al} ., 2008). It is in this aggressive context that
ecological restoration needs to reverse and prevent further degradation,
and then assist in the recovery of an ecosystem. An important element in
this task is to more clearly understand how human behaviour influences
different aspects of an ecosystem (Schroder, 2009). Restoration models
have been evolving for several decades (Suding \& Hobbs, 2009), and have
proven to be successful for prioritising large- and small-scale
restoration projects.

The restoration of weed-dominated grasslands has received extensive both
popular and academic interests. Native tufted perennial grasses have
been described as keystone species as they resist weed invasion and
maintain ecosystem processes (Stromberg \emph{et al} ., 2009;
Prober\emph{et al} ., 2005). Human induced disturbance has resulted to
the decline of these native grasses and thus non-native perennial
grasses have established, which significantly reduces the carrying
capacity and biodiversity. This has been observed in grasslands
dominated by\emph{Nassella trichotoma} throughout south-eastern
Australia (Campbell \& Nicol, 1999; Jacobs \& Everitt, 2012), South
Africa (Joubert, 1984) and New Zealand (Lameroux \emph{et al} ., 2011;
Lusk \emph{et al} ., 2017). Annual weeds are also highly problematic,
and outcompete native perennial grasses in their early life stages, and
this is most prevalent after disturbance (Musil \emph{et al} ., 2005;
Bartolome \emph{et al} ., 2009; James \emph{et al} ., 2011). Because
invasive plants are generally one of the main drivers for holding these
ecosystems in degraded states, the main focus of restoration efforts is
on reducing the dominant weed population and promoting competition from
native species. Passive and active restoration techniques have been used
in diverse combinations to achieve this outcome at varying levels of
success throughout different temperate grasslands (Table 2).

\textbf{Passive restoration}

Passive ecological restoration involves removing human induced degrading
pressures from a site with minimal remediation. In many cases, it is
presumed that non-target species will expand without human intervention,
however many passive restorations have observed weeds decline with
sufficient native recruitment (Sinkins \& Otfinowski, 2012;
Valko\emph{et al} ., 2017). Notable vegetation shifts often occur within
10 years of rest (Smallbone, 2014), however, some may take several
decades to reach similar species richness as the remnant sites, and even
then, the composition of vegetation can significantly differ (van de
Merwe \& van Rooyen, 2011).

Successful passive restoration involves careful grazing management and
the ability for target species to recruit and establish. Rapid recovery
of a degraded Hungarian alkaline grassland was observed (within one
year) in sites directly adjacent a natural grassland, and within six
years, all sites where restored regardless of proximity to the remnant
site (Valko \emph{et al} ., 2017). In this example, the dispersal of
native plant propagules was promoted by livestock roaming between
natural and degraded sites (Valkko \emph{et al} ., 2017). Grazing
animals were also observed to provide an important service in
maintaining species richness for highly productive Themeda grassland in
south-east Australia (Schultz \emph{et al} ., 2011). Grasslands often
require disturbances such as grazing or fire to maintain species
richness and grazing animals remove excess phytomass in order to
generate niche space for rarer species.

While grazing plays an important role in maintaining highly productive
grasslands, those suffering extensive degradation or of lower
productivity often benefit from the complete removal of grazing
livestock. Grazing exclusion is a cost-effective tool for passive
restoration, particularly if native species are well represented. A
long-term (20 and 30 years) grazing exclusion zone was developed in the
steppe grasslands of China, which observed an increase in perennial
grass cover, as well as higher density bud banks of these favourable
grasses when compared to the grazing sites (Zhao \emph{et al} ., 2019).
The long-term (40 years) removal of cattle from northern fescue prairies
in Canada was effective for reducing some invasive plants, but not
others, including \emph{Poa pratensis} , which in some areas occupied up
to 90\% of the canopy (Sinkins \& Otfinowski, 2012). Generally,
grassland species have short-lived seedbanks and if desirable species
are rare, or the site is isolated from remnant patches, the seedbank
will continue to diminish (Bossuyt \& Hermy, 2003). Further, the
recruitment of native species in degraded temperate grasslands is rare,
as seedlings often fail to survive as a result of competitive weed
interactions (Morgan, 2001; Lenz \& Facelli, 2005). This demonstrates
that isolated sites dominated by aggressive weeds may not be suitable
for passive restoration. While passive restoration has proven successful
under specific conditions, in sites where invasive plants have dominated
for several decades and biotic and abiotic thresholds have been crossed,
active intervention will be required.

\textbf{Active restoration}

Active ecological restoration involves the integration of management
techniques, such as revegetation, herbicide application, or mechanical
soil disturbance to take an ecosystem from a degraded state to one that
is functional, self-sustaining and resilient. In weed dominant systems,
restoration efforts that focus to remove invasive plants and promote
dense, native competition are often the most successful. In order to
actively restore a degraded landscape, understanding the sites history
can be critical. The history of a site can identify the factors that
moved it into a degraded state, and whether these changes occurred
rapidly or continuously over an extended interval (Prober \& Thiele,
2005). Further, the historic vegetation cover can act as a restoration
target, and guide managers on revegetation assemblages (Prober \&
Thiele, 2005). Active restoration of weed dominated temperate grasslands
should consider; i) the removal of the weeds biomass, ii) manipulation
of the soil to return it to remnant condition, iii) revegetation of
native propagules, and ix) site specific grazing management.

\emph{Targeting weed biomass}

The removal of dense weed biomass is critical for reducing competition
for naturally recruiting native's species, or those added via
revegetation efforts. Weeds are often fast growing and form dense
canopies, which reduces light to the soil and can thus restricts the
germination and subsequent growth of native seeds or seedlings. Further,
many annual grassland weeds have higher nutrient requirements than
native perennial grasses, thus creating a highly competitive environment
for natives to establish. The most commonly used methods for reducing
weed biomass include hand removal, herbicide application, and fire.

Grubbing, or hand weeding, is a restoration technique that completely
removes unwanted plants (Tikka \emph{et al} ., 2001). While highly
effective, this method is also very labour intensive, and is usually
only appropriate for smaller scale projects (Gibson-Roy \emph{et al} .,
2007). That said, every three years, community efforts have successfully
removed 34\% of the invasive perennial grass, \emph{N.
trichotoma}throughout Canterbury, New Zealand, which has contained the
population from further expansion (Bourdot \& Saville, 2019). Grubbing
is the best solution for sites where weeds are newly emerging and easy
to remove, or where only a few individuals have established, such as
roadsides. Grubbing can prove a critical tool for post restoration
management by quickly removing reinvading weeds. Grubbing is one of the
most effective methods to reduce competition for space, light and soil
nutrients as the whole plant is instantly removed.

Herbicide application is often an economically viable and effective
solution for reducing weed competition. Herbicide works most effectively
when integrated with other treatments, as seen by Johnson \emph{et al} .
(2018) who observed spot-spraying weeds with glyphosate significantly
improved the establishment of native forbs seeds when combined with
fencing, and the removal of leaf litter. Aerially spraying clopyralid
(at a rate of 37.4 L/ha) was successful in reducing woody weed
encroachment and enhancing plant diversity when combined with prescribed
burning (Ansley \& Casellano, 2006). Waller \emph{et al} . (2016) also
observed significantly improved native establishment when herbicide was
combined with fire, tillage and rodent exclusions. In a degraded grazing
exclusion zone, Huddleson \& Young (2005) identified herbicide
application on its own was effective for not only reduced annual weed
competition by 40\%, but increased native establishment ten-fold.

In some cases, herbicide application was ineffective at improving native
establishment (Cole \emph{et al} ., 2005; Conrad \& Tischew, 2011).
Spot-spraying Snapshot (a pre-emergent herbicide containing trifluralin
and isoxaban) at 2.5kg per 100m\textsuperscript{2} significantly reduced
the emergence of native forbs compared to the controls in South Africa
(Musil \emph{et al} ., 2005), however was effective for controlling
invasive annual grasses. In New Zealand, boom-spraying flupropanate at
1.49kg a.i/ha reduced native pasture grass by 89\% (Lusk \emph{et al} .,
2017). In these cases, using herbicides selectively can enhance
restoration outcomes. Selective herbicides are used to kill the unwanted
weeds, while the desirable species remain unharmed, and this can be
attributed to; plant chemistry, physical growth parameters and plant
physiology (Sutton, 1967). It is important to note that the constant use
of herbicides within an ecosystem can promote the emergence of herbicide
resistant populations, thus reducing its long-term effectiveness.
Resistance to arguably the world's most important herbicide, glyphosate,
has already been observed in several weeds (Powles 2008), including
Conyza spp. (Feng \emph{et al} ., 2004; Urbano \emph{et al} ., 2007) and
Lolium spp. (Baerson \emph{et al} ., 2002; Yanniccari \emph{et al} .,
2017). It is considered important, therefore, that herbicides should be
used selectively and in combination with other control methods in order
to secure their effectiveness for the long term.

Fire is one of the most effective tools for restoring temperate
grasslands that are dominated by weeds. Historically, grasslands are
ecosystems that are accustomed to frequent fire events, and altered fire
regimes in Australia (Stuwe \& Parson, 1977), New Zealand (Mark, 2007;
Standish \emph{et al} ., 2009), the United States (Foster \& Gross 1998;
Stromberg 2007), and South Africa (Sankaran \& Anderson, 2009) have been
linked to the modification of these landscapes (Archer \emph{et al} .,
1988; Knicker, 2007). Fire quickly creates available space for heat
resistant seeds to germinate and grow relatively free of competition
(Meyer \& Schiffman, 1999), and a number of studies have observed that
fire significantly reduces weed species and promotes native recruitment
(Huddleson \& Young, 2005; Prober \emph{et al} ., 2005; Bryant \emph{et
al} ., 2017). Lipoma \emph{et al} . (2018) identified fire to
significantly reduce the viable number of seeds in the soil compared to
pre-burnt conditions, and as most weeds often have dense seedbanks, this
can be beneficial in reducing at least the surface seedbanks of some
species (Peltzer \& Douglass, 2019). In contrast, some species,
particularly broadleaf weeds such as \emph{Echium plantagineum} , are
promoted by fire (Prober \emph{et al} ., 2004). Heat tolerance in seeds
has been linked to seed shape, with more rounded seeds demonstrating
higher resistance than thinner seeds in European temperate grasslands
(Ruprecht \emph{et al} ., 2015). This suggests that follow up weed
management of burnt sites is critical for the successful establishment
of native species. In Australia, a summer wildfire was observed to kill
90\% of the standing native spear-grass (Austrastipa spp.), which is
considered relatively fire tolerant (Sinclair \emph{et al} ., 2014).
Fire also offers soil manipulation services as carbon and nitrogen
volatize at 180 and 200\textsuperscript{O}C respectively (DiTomaso
\emph{et al} ., 2006), therefore hot fires can remove soil nutrients
that advantage annual weeds and further inhibit their re-establishment
(Knicker, 2007). Strategically burning when problematic weeds are
actively growing can effectively prevent seed set for that season
(Prober \emph{et al} ., 2005). The complexity of fire effects suggests
that post management plans should be specific for the site in order to
promote the establishment of a healthy native grassland community (Musil
\emph{et al} ., 2005; DiTomaso \emph{et al} ., 2006).

\emph{Soil manipulation}

Altered soil nutrients and textures resulting from agriculture have
important consequences on the ability for standing vegetation to take up
water and nutrients (Sankaran \& Anderson, 2009), therefore restoring
these factors to resemble historic levels can be important for weed
suppression. Soil nutrients such as nitrogen, phosphorous and potassium,
are altered by agricultural practises, and even after agriculture has
ceased, the soil nutrient levels remain higher than historical levels
(Prober \emph{et al} ., 2005). Annual weeds become problematic in
environments with high nitrogen, where they are able to quickly dominate
over the slower-growing native perennial grasses (Huddleson \& Young,
2005). Perennials invest in developing deeper rots systems that allow
them to store and recycle nutrients, giving established perennials an
advantage over annuals in areas of low nutrient availability. Therefore,
integrating control methods that target soil nutrient levels should be
strongly considered for those grassland restoration projects in areas
that have a history of agriculture. This can be achieved with the
addition of a carbon source, such as sucrose, can increases soil
microbial activity reduces soil nutrients, and leaves them unavailable
for used by nutrient-adapted weeds. This technique has been used
successfully in Australia (Prober \emph{et al} ., 2005; Hacker \emph{et
al} ., 2011) and the United States (Blumenthal \emph{et al} ., 2003). In
one reported prairie restoration, carbon addition reduced soil nitrogen
by 86\%, which subsequently reduced weed biomass by 54\%
(Blumenthal\emph{et al} ., 2003). While carbon addition has proven to be
successful, it is a time and resource-demanding approach. Prober
\emph{et al} . (2004) used 500g of sugar for every square metre, which
was reapplied every three months, making this technique difficult to
implement at a landscape scale. Further, it is only suitable with
nitrophillic weeds (Blumenthal \emph{et al} ., 2003).

Another method for altering soil dynamics is through mechanical
disturbance techniques, such as tilling or scalping. These techniques
are effective for creating an environment that promotes the
establishment of broadcasted seeds and reduces competition from weeds
(Tikka \emph{et al} ., 2001). As many weed seeds respond positively to
disturbance events, tillage can be used to stimulate stored seedbanks
(Stromberg \emph{et al} ., 2007). Scalping is a technique where top soil
is removed from a site and subsequently treated. This is a useful
technique in highly degraded sites that are heavily infested by weeds as
it removes their seedbank as well as the elevated nutrient levels that
promote their growth and establishment (Brown \emph{et al} ., 2017).
Consequently, scalping may result in excessive waste soil, increases
erosion rates, habitat loss and disrupted mycorrhizal symbiosis, and
therefore should be implemented with caution (Gibson-Roy \emph{et al} .,
2010; Gerlach, 2015; Brown \emph{et al} ., 2017). Further, weed
reinvasion can occur on scalped sites, and Gerlach (2015) identified
weeds to occupy 70\% of the ground cover after three years of scalping
and revegetation. Scalping treatments followed by with spot-spraying has
proven to be successful within small scale (1m X1m plots) for reducing
all vegetation (Gibson-Roy \emph{et al} ., 2010).

\emph{Revegetation}

Establishing dense competition from desirable species is the most
effective way to reduce weeds and return natural ecosystem
functionality. In the case that natural regeneration is an unviable
option, competition can be introduced using a variety of methods
including direct seeding (Thomas \emph{et al} ., 2019; Cole \emph{et al}
., 2005), transfer of threshing material (Baasch \emph{et al} ., 2016)
or hay (Sengel \emph{et al} ., 2016), direct drilling (Bakker \emph{et
al} ., 2003) and plant plugs (Tikka \emph{et al} ., 2001). Hedberg \&
Kotowski (2010) reviewed the effectiveness of different revegetation
options for fragmented grasslands and found that direct seeding (sowing
and broadcasting) to be the most widely used and most effective for
introducing species back to semi-natural systems. However, they
specifically recommended the use of plant plugs for the establishment of
rarer species (Hedberg \& Kotowski, 2010).

The effectiveness of species richness in seed mixes has been explored
for grassland restorations, with both high and low rates demonstrating
beneficial results dependent on the projects scope (Prober \& Thiele,
2005; Wortley \emph{et al} ., 2013). The determined species mix is often
reflective of the goals of that particular restoration project; for
example, Conrad \& Tishew (2011) found a high seed mix of 35 species
achieved their goals of increasing species diversity as well as
establishing target species, whilst in another area, Huddleson \& Young
(2005) used a mix of only three native grasses to successfully
outcompete weeds. Further, it appears that high species diversity
improves the establishment of native species (Barr \emph{et al} .,
2017), long-term resilience to weed reinvasion (Carter \& Blair, 2012;
Scotton, 2016), and provide habitat for recolonization of threatened
wildlife (McDougal \& Morgan, 2005). Nemec \emph{et al} . (2013) has
demonstrated seed diversity to be a more important factor than seed rate
for achieving reasonable competition for weeds. While high seed rates
can improve the chances in successfully outcompeting weeds (Tikka
\emph{et al} ., 2001; Bakker \emph{et al} ., 2003; Barr \emph{et al} .,
2017), this approach can waste seeds as a result of higher intraspecific
competition, and the associated high costs can make it unpractical
(Sheley \emph{et al} ., 2006; Wagner \emph{et al} ., 2011). Seed mixes
low in diversity and density can promote spontaneous secondary
succession, and this can stimulate ecosystem processes more quickly
(Lengyel\emph{et al} ., 2012). We note that the failure of sown seeds to
establish can be linked to several factors, including herbivory, adverse
weather conditions, and species competition (Gibson-Roy \emph{et al} .,
2007), therefore implementing pre-sowing management that minimises these
threats is critical. Whilst it is clear that the introduction of seeds
or seedlings is often critical for the restoration of many degraded
temperate grasslands it is also clear that the best implementation
method will be dependent on the site, scale and funding available to the
project (Prober \& Thiele, 2005).

\emph{Grazing management}

Grazers play an important role in the continuous removal of leaf litter
and generating available space for new recruitment (Lengyel \emph{et al}
., 2012; T\selectlanguage{ngerman}örök \emph{et al} ., 2018), which can promote species richness
(Towne \emph{et al} ., 2005; Klaus \emph{et al} ., 2018). Germination of
the native North American prairie grass, \emph{Nassella pulchra} was
enhanced by burning and sheep grazing (Dyer, 2002). Moderate grazing
(30-50 within a 303ha enclosure) from \emph{Bos bison}significantly
improved the species richness of a Prairie grassland within its later
stages of development (approximately 10-years after revegetation)
(Wilsey \& Martin, 2015), this was also observed within tallgrass
prairies (Towne \emph{et al} ., 2005). Livestock can transport seeds of
important species over great distances via endo or ectozoochory if
remnant sites are available (Lengyel \emph{et al} ., 2012; Török\emph{et
al} ., 2018). Further, the careful management of paddock rotations for
grazing livestock has been identified to be critical in maintaining
genetic diversity for plants threatened by fragmentation (Plue \emph{et
al} ., 2019). High genetic diversity can allow for a population to react
more responsively to disturbance events through improved resilience.

Overgrazing by livestock is one of the leading causes of grassland
degradation (Bartolome \emph{et al} ., 2009; Zha \& Gao, 2011;
Wortley\emph{et al} ., 2013). The effects of different degrees of
overgrazing were observed by Török \emph{et al} . (2018) within four
different Hungarian steppe grassland communities. They found that the
highest richness was achieved from low grazing (less than one animal per
hectare), but medium was also suitable (1-2.5 animals per hectare), and
grazing densities above this had detrimental effects of species
richness. Further, the different grassland communities responded
differently to the grazing intensities suggesting they are grassland
specific (Török \emph{et al} ., 2018). Competition dynamics between forb
and grass species were altered by livestock grazing in southern
Argentina (Díaz Barradas \emph{et al} ., 2001). Under sheep grazing, the
grasses did not produce inflorescences and forbs became taller and more
abundant compared to non-grazing tracts, where grass species dominated
(Díaz Barradas \emph{et al} ., 2001). Forb cover was also observed to
increase in Prairie grasslands when exposed to grazing from cattle and
bison (Towne \emph{et al} ., 2005). Therefore, grazing intensities
should be carefully managed, particularly during drought periods to
promote competition from native perennial grasses (Klaus \emph{et al} .,
2018), and resting paddocks from grazing when natives are emerging,
particularly if herbage is sparse, could improve their establishment and
survival (Clarke \& Davison, 2014).

\textbf{Considerations for long-term management}

\emph{The role of grassland seedbanks}

Limitations in funding and technology makes it difficult to manage
grassland restoration projects over long time scales (Freudenberger \&
Gibson-Roy, 2011; Lengyel \emph{et al} ., 2012). Without follow-up
management, weeds can re-establish either from regeneration from the
seedbank or from migration of seeds from surrounding sites
(Gibson-Roy\emph{et al} ., 2007). By knowing how long the dominant weeds
seeds remain viable within the soil seedbank, we can make
recommendations as to how long a site should be actively managed.

Weed species are often prolific seed producers, therefore, to protect a
rehabilitated grassland from being reinvaded by a dominant weed species,
an understanding of its seedbank persistence is required.
Gardener\emph{et al} ., (2003) identified that after three years of no
seed migration within a \emph{Nassella neesiana} dominated grassland,
there was still, on average, 1457 viable seeds per square metre of this
species, indicating that ongoing management of only three years would
not be effective for controlling this invasive grass. Seed longevity
studies are useful for investigating how long a species can persist at
different depth or soil types. The common trend is that seed viability
declines with shallow burial and increased duration within the seedbank.
This is seen in \emph{Andropogon gayanus} (Bebawi \emph{et al} .,
2018),\emph{R. raphanistrum} (Reeves \emph{et al} ., 1981), \emph{Conyza
canadensis} (Vargas \emph{et al} ., 2018) and \emph{Artemisia
tridentate}(Wijayratne \& Pyke, 2012). Seeds located on or just below
the soil surface are exposed to more intense fluctuations in soil
moisture and temperature compared to deeper buried seeds, and these
fluctuations can result in the shallow buried seeds drying out. A
two-year study found\emph{A} . \emph{tridentate} had only 0-11\% of
seeds remaining viable at the soil surface compared with almost half of
the seeds maintaining viability at 3cm depth (Wijayratne \& Pyke, 2012).
Seed predation by mammals, birds, or soil microbes is also enhanced at
these shallow depths (Dalling \emph{et al} ., 2011). Seeds that are
buried deeper into the soil profile, are often better protected from
these devitalizing and predatory pressures (Bebawi \emph{et al} .,
2018). The trade-off, however, is that at these depths seed dormancy is
usually prolonged, particularly for photoblastic seeds (Benvenuti
\emph{et al.} , 2005; Ahmed \emph{et al.} , 2015), making these weeds
troublesome for managing cropping systems utilizing tillage, since this
can resurface viable invasive seeds, resulting in reinvasion.

\emph{Anticipating climate Change}

Grasslands currently store approximately 34\% of the worlds terrestrial
carbon, making these ecosystems important carbon sinks and play a
critical role in climate change mitigation (Contant, 2010). Carbon
sequestration is achieved by grassland vegetation holding organic carbon
within their roots, therefore higher sequestration is found in less
disturbed grasslands with long lived perennial grasses that develop
dense root systems (Acharya \emph{et al} ., 2012). This suggests that
long term management of grasslands will likely provide greater climate
regulation. Carbon sequestration has been observed to improve with good
management techniques, particularly the addition of nitrogen fixing
plants (De Deyn \emph{et al} ., 2011), addition of fertilizers and lime
(Acharya \emph{et al} ., 2012) and withholding excessive grazing
(Eze\emph{et al} ., 2018). Further, grasslands are essential for human
food security and provide an income for approximately 1.3 billion people
around the world (Suttie \emph{et al} ., 2005). Livestock grazing
utilizes 80\% of the total agricultural land and contributes to 40\% of
global agricultural production (Suttie \emph{et al} ., 2005). It is
predicted that demands for animal-based proteins and dairy are only
going to increase as a result of projected population growth (O'Mara,
2012), making functioning grasslands critical for providing adequate
nutritional resources.

Accelerated climate change adds a further element of complexity for
managing restoration projects into the future. It is expected that for
every 1\textsuperscript{O}C increase in air temperature, there will be a
1.5\textsuperscript{O}C increase in soil temperature (Ooi \emph{et al}
., 2011), which may also cause disruptions to the seedbanks of many
plant species. Temperature has proven to be an important environmental
factor for breaking seed dormancy and these increased temperatures could
influence these important physiological processes (Ooi, 2012;
Prosotto\emph{et al} ., 2014). Further, atmospheric CO\textsubscript{2}
has steadily risen from 325ppm recorded in 1970 to 405ppm in 2017
(Lindsay, 2018), and this is expected to approximately double by the end
of this century (IPCC, 2019). Enhanced atmospheric CO\textsubscript{2}
can result in higher saturation of CO\textsubscript{2}, potentially
reducing photorespiration in C\textsubscript{3} plants, even under a
warmer climate. This increased physiological efficiency has been
demonstrated to alter dynamics between C\textsubscript{3} and
C\textsubscript{4}plants (Dukes, 2000). As a result of these
physiological improvements, such as increased water-use efficiency
(Varga \emph{et al} ., 2015), plants can allocate more resources to
growth and fecundity and these changes have also been observed to be
more pronounce in weeds than natives or crops (\emph{Marble et al} .,
2015). Changes in extreme weather patterns is expected to increase as a
result of human induced climate change. Compared to pre-industrial data,
changes in the intensity and pattern of rainfall events are already
being noticed (Power \emph{et al} ., 2017). Changes in rainfall have
direct consequences on the intensity and frequency of fire, drought and
flood events (Ooi, 2012). As these factors play an important role in
shaping the vegetation of ecosystems and agroecosystems, new challenges
for managing native and weed competition dynamics can be expected.

\textbf{Conclusion}

Temperate grasslands are now significantly degraded throughout the world
as a result of human actions. Weeds now dominate many of these degraded
systems and act to hold them in this undesirable altered state. A number
of successful restoration techniques have been developed to reduce weed
dominance and promote native species, but it is clear that a single
technique for restoration is rarely successful for the long term. In
order to reduce dominant weeds, we must continue to research the
integration of control methods that are economical, practical and
applicable to temperate grasslands at a local, regional and global
scales. Researchers should also aim to develop long-term studies that
observe successional changes in plant dynamics as a result of various
treatments. It is critical that managers plan now for changes in weather
patterns (such as rainfall frequency and intensity) as a result
accelerated climate change. This review recognises the similarities in
successful temperate grassland restorations involve the ongoing effort
of targeting the above and below ground density of the dominant weeds.
Revegetation methods are often site- and study-specific and depend on
proximity of remanent vegetation, budget and restoration goals.

\textbf{Conflict of Interest}

The Authors declare no conflict of interest.

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\verb`Table 2.docx`  available at \url{https://authorea.com/users/316591/articles/446754-weed-management-for-the-land-scape-scale-restoration-of-global-temperate-grasslands-a-review}

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