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Review article
Grasslands are globally threatened and their biodiversity, including grassland birds, is
declining markedly. To inform grassland bird conservation globally, we systematically
reviewed threats and conservation actions for grassland birds, extracting data from 528
papers. Across the 10 primary grassland regions of the globe, agriculture was the most
frequently or joint most frequently reported threat in nine regions (reported as a threat
in 73% of publications); hunting was the most frequently reported threat in the remain-
ing region. Natural system modifications (reported as a threat in 32% of publications)
and climate change and severe weather (24%) were less frequently reported threats com-
pared with agriculture. The types of threat from agriculture varied regionally, but the
most pervasive were livestock farming and ranching (reported in 58% of publications
where agriculture was a primary threat) and non-timber cropping (43%). Most agricul-
tural threats relate to intensification, but agricultural abandonment, typically the cessa-
tion of grazing, sometimes accompanied by tree planting/succession, poses an emerging
threat to some grassland birds (reported in 32% of publications where agriculture was a
primary threat). The most frequent conservation actions implemented to date include
land/water management and protection, and species-specific management actions.
Authors of reviewed publications in almost all regions recommend more land/water
management, followed by calls for further land/water protection. The parlous state of
grassland birds globally suggests that existing conservation actions for grasslands are inad-
equate. Furthermore, our review suggests that these should be primarily targeted at
*Corresponding author.
Email: david.douglas@rspb.org.uk
Twitter: @davidjtdouglas
Around 75% of the global land surface has been threatened (Azpiroz et al. 2012). Despite wide-
significantly altered by human activities, and most spread declines and highly threatened avifaunas,
indicators demonstrate declines in ecosystem integ- grassland birds have, at least in some regions, been
rity and biodiversity (IPBES 2019). Globally, one poorly studied compared with forest birds (Kamp
of the most threatened habitats is grasslands, found et al. 2011, Douglas et al. 2014, Maphisa
on every continent except Antarctica and esti- et al. 2017). It is nevertheless widely recognized
mated to cover 26% of global land, second only to that in the Americas, grassland birds are facing
forests in area (Panunzi 2008). Natural and urgent conservation needs (Rosenberg et al. 2019)
anthropogenically modified grasslands supply and in Africa, research is needed to understand
essential ecosystem services including water provi- how moist high-altitude grasslands can be managed
sion, carbon sequestration and biodiversity (Neary to help conserve birds (Maphisa et al. 2017).
& Leonard 2020). Designing conservation science agendas for
Grasslands are, however, highly threatened, due important grassland areas requires an understand-
largely to habitat loss and inadequate protection ing of the ecological impacts of grassland manage-
(Hoekstra et al. 2005). These threats to grasslands ment (Ahlering et al. 2020). However, threats and
include changes in agricultural intensity (Squires required conservation actions may vary regionally;
et al. 2018); in southern Africa, for example, 23% for example, grasslands are under constant threat
of grassland biomes are now under cultivation, from drought in some regions (Gibson & New-
60% have been irreversibly transformed, only man 2019) but at greater risk from land use
around 2–4.5% are formally protected with most change in others (Kamp et al. 2011). Some regions
of the remainder used for livestock grazing, with might also have more effective conservation pro-
only around 3.6% in a natural or intact state (Fair- grammes than others. This reinforces the need for
banks et al. 2000, Maphisa et al. 2017, Skowno a comparison of regional threats and conservation
et al. 2019). Similarly, vast areas of other major actions to help define the scope and scale of grass-
grasslands such as the Eurasian steppe (Kamp land bird management challenges globally.
et al. 2011) and North American prairies (Comer We used a systematic literature review and syn-
& Hoagland 2020) have been converted for culti- thesis to quantify the key threats and conservation
vation and/or domestic livestock grazing. Further actions for grassland birds across the globe’s major
threats include habitat loss and degradation to grassland regions. Using these summaries, we
infrastructure development for mining, energy gen- explored regional commonalities and differences to
eration and urbanization, fragmentation, shrub and better identify the needs of grassland birds
tree encroachment, and climate change (Far- worldwide.
ley 2007, Daniel & Koper 2019, Gibson &
Newman 2019).
METHODS
Cumulatively, these threats have resulted in
numerous impacts on grassland-dependent wildlife,
Definitions
including birds. Globally, grassland birds are
decreasing precipitately. In North America and We defined ‘grassland’ following Dixon
Europe, they are declining more than birds of any et al. (2014): not permanent wetland; at least 10%
other biome (Burns et al. 2021, North American vascular vegetation cover; graminoids with > 25%
Bird Conservation Initiative 2022). In Asia, grass- cover or at least exceeding other herbaceous and
lands are second only to forests for the number of shrub cover; dominant over other growth forms;
globally threatened bird species supported (Birdlife shrubs have < 25% cover, trees have < 10% can-
International 2003) and in southeast South Amer- opy cover. We therefore included grasslands along
ica more than a quarter of bird species that regu- a moisture gradient from arid to wet and included
larly use grasslands are globally threatened or near- highveld, meadows, pampas, prairies, steppe and
Figure 1. Study sites of publications in the review. Smaller dots represent studies for which exact coordinates were provided by the
authors or that could be identified on maps from the site description. Larger dots are the centroids of larger study areas (often span-
ning several 100 km). A total of 664 study sites and regions from 449 reviewed papers are mapped. For 84 publications with
national, continental or global focus, no locations are displayed. The shaded grey areas are defined as primary (naturally occurring)
grasslands by Dixon et al. (2014); dots outside these areas largely represent studies from secondary grasslands (created from
anthropic activities). Our definition of grassland is therefore: not permanent wetland; at least 10% vascular vegetation cover; grami-
noids with > 25% cover or at least exceeding other herbaceous and shrub cover; dominant over other growth forms; shrubs have
< 25% cover, trees have < 10% canopy cover. This includes grasslands along a moisture gradient from arid to wet and included
highveld, meadows, pampas, prairies, steppe and upland wet mesic grassland habitats, excluding habitats with a marked tree or
shrub component such as forest-steppe, chaparral or savanna. Map uses a Winkel tripel projection.
allowed for multiple common terms for grasslands that were recommended, including any past, exist-
per region. We used Boolean search operators and ing or recommended action described in publica-
wildcards (*) to account for variation in the suf- tions, even if not the focus of the study. We
fixes of words included in the search string. In gen- present a tallied summary of threats and actions
eral, the Web of Science search string followed the globally, and a narrative summary of relevant liter-
format ((country OR region) AND (steppe* OR ature per region. The ‘tallying’ approach adopted
grassland* OR prairie* OR pasture* OR meadow* for summing threats and actions provided a repeat-
OR pampas* OR highveld*) AND (avian* OR able and logistically feasible means for summariz-
aves* OR bird* OR ornith*) AND (conservation* ing the large and diverse volume of literature
OR management*)) (Table S2). reviewed, and identifying the most frequently
cited threats and actions, and we consider this
appropriate. A full meta-analysis was beyond the
Publication review
available peoplepower. As a result, our approach
We screened and subsequently reviewed up to a does not consider potential biases such as variation
maximum of the 200 most recent publications in the scale or intensity of threats per publication,
returned per region, with the oldest publications and accordingly the number of species impacted
reviewed varying regionally from 1992 to 2018 by threats/actions. Where a threat to grassland
(Table S3). In well-studied regions this approach birds arises due to impacts on the underlying habi-
meant that publications were all recent, whereas tats, which probably applies to numerous species,
in less studied regions they comprised newer and e.g. agriculture, this is less of an issue. But care
older publications. Given the available personnel, should be taken in the interpretation of threats
it was not feasible to process more than 200 publi- such as hunting, where only a select number of
cations per region. Although threats might increase species might be subject to hunting. Similarly, for
or diminish in importance over time, this approach regions returning small sample sizes of relevant
does mean that, particularly in well-studied publications (Caribbean and to a lesser extent
regions, we are confident that we have reviewed Middle East), we urge caution in the interpretation
studies addressing the most current, pressing of threats.
threats. We first screened publication abstracts for
eligibility, retaining papers focusing on ecology,
RESULTS
conservation, or management of grassland birds in
grassland habitats, within the focal region, exclud- No relevant publications were found for Oceania
ing publications unavailable in English or inaccessi- (excluding Australasia). Across the remaining 10
ble online (which was rare). Publications meeting regions, data were extracted from 528 publications
the above inclusion criteria were reviewed in full (Table S3).
(2–101 per region; see Table S3 and Results). We
used hierarchical categories and definitions from
Summary of threats
Salafsky et al. (2008), with additional categories of
our own, characterizing 15 primary threats (Fig. 2, ‘Agriculture and aquaculture’ was the threat most
Table S4), and eight primary conservation actions frequently reported to impact birds in eight
(Fig. 3, Table S5). Primary threats and actions regions and tied with natural system modifications
were also subclassified into sub-threats from Sal- for most frequently reported threat in the Carib-
afsky et al. (2008) and our own additional catego- bean (Fig. 2). In the Middle East, biological
ries (Fig. 2, Tables S4 and S5). Any statistically resource use (hunting and collecting) was the most
supported threats described in text, figures or frequently reported threat. Globally, agriculture
tables were assigned to the categories described and aquaculture far exceeded other threats in
above. We summed the number of publications terms of the overall number of publications report-
citing each primary threat per region, and ‘Agricul- ing it as a threat (73%), followed by natural sys-
ture and aquaculture‘ sub-threats, the latter tem modifications (32%), and climate change and
because this primary threat far exceeded other pri- severe weather (24%; Fig. 2).
mary threats in terms of number of publications The frequency of reporting of different agricul-
reporting it. We separated conservation actions tural sub-threats varied regionally. ‘Livestock farm-
into those in place (‘existing’ actions) and those ing and ranching’ was most frequent or joint most
Figure 2. Major threats to grassland birds (following Salafsky et al. 2008) and their proportions in respective regions (a–j) expressed
as the percentage of studies in which they were mentioned. ‘Agriculture and aquaculture’ was the most frequently, or joint most fre-
quently, reported primary threat to grassland birds in nine out of 10 regions globally and second most frequent in the other region.
No data were available for Oceania. Sample sizes are number of studies from the total sample assessed that addressed threats to
grassland birds.
frequent in three regions (Australasia, Central and fallow, also poses risks to some grassland birds as it
South America, Fig. 3; 45% of publications within was most frequently or joint most frequently
‘agriculture and aquaculture’). Non-timber crop- reported in three regions (Caribbean, Central Asia
ping was most frequent or joint most frequent in and North America, Fig. 3). It should be noted
three regions (Asia, Caribbean and Europe, Fig. 3; that the sample size of papers addressing threats in
42% of publications within agriculture and aqua- the Caribbean was small (n = 2), which could
culture). Agricultural abandonment, typically the result in a threat or sub-threat being assigned a dis-
cessation of domestic grazing or cultivated land left proportionately large weighting.
Figure 3. The relative importance of sub-threats (following Salafsky et al. 2008) to grassland birds within the primary threat agricul-
ture and aquaculture varies by regions.
Figure 4. The frequency of existing (grey bars) and recommended (black bars) conservation actions (a–j) for grassland birds varies
regionally. Sample sizes are number of studies from the total sample assessed that addressed conservation actions for grassland
birds.
habitats need to be conserved is such that large et al. 2012). Low rainfall may also compound the
landscape mosaics of exclosed or lightly grazed impacts of overgrazing on the Plains-wanderer
grasslands and low-intensity farmland, including (Baker-Gabb et al. 2016).
community conservation reserves, represent a Land protection through nature reserves and
workable compromise, vital for the survival of national parks is the most common conservation
three species of bustard, for example (Collar action (Johnstone et al. 2015). Recommended
et al. 2017, Varun & Dutta 2020). actions include patch grazing and burning under
Other recommended conservation measures in appropriate regimens to increase habitat heteroge-
this region include regulation of powerlines and neity, exclosures and culling of herbivores to limit
fencing; control of hunting, poaching, dog preda- overgrazing, and payments to landowners to incen-
tion and inappropriate burning of grassland; sus- tivize conservation (Baker-Gabb et al. 2016).
tained campaigns of public awareness and Well-known species such as the Plains-wanderer
engagement; and strong government and interna- could make effective local flagships for promoting
tional support (Collar et al. 2017). grassland conservation (Johnstone et al. 2015).
Horn 2016). Many grasslands in the region arise may be ecological traps for species including the
through agricultural activity (Mendenhall Black Lark Melanocorypha yeltoniensis because of
et al. 2016), including pastoral grasslands in forest higher nest predation (Lameris et al. 2016).
clearings, which fluctuate spatially and temporally Moreover, recent recultivation of abandoned
with agricultural activity (Holl et al. 2000, Ospina cropland might lead to new declines (Kamp
et al. 2012, Roels et al. 2019). Furthermore, some et al. 2011).
grassland bird species are more abundant in modi- Grassland birds have also been strongly
fied compared with natural landscapes, making it affected by historical fluctuations in livestock
challenging to develop conservation strategies management. Traditional semi-nomadic grazing
(Roels et al. 2019). Other threats to grasslands and generated a landscape of varying grazing intensi-
their birds include commercial forestry, forest res- ties until the 1980s (Robinson et al. 2003), pro-
toration or succession, and hunting (Arguedas- ducing habitat for a diversity of grassland species.
Negrini 2001, Martin & Buchholz 2018, Roels However, livestock numbers collapsed after the
et al. 2019). Forestry, energy generation and agri- break-up of the Soviet Union, and vast areas are
culture can lead to secondary threats such as resi- no longer grazed (Dara et al. 2020). The resulting
dential and commercial development and roads biomass accumulation is associated with increased
(Piaskowski et al. 2005). Only one publication fire frequency, triggering a transition from diverse
indicated that climate change is a threat to grass- steppe to uniform swards dominated by grasses
land birds, and land use change is considered a (Freitag et al. 2020). This has resulted in changed
more immediate threat (La Sorte et al. 2017). abundances of steppe mammals (Koshkina
Land/water management was the most common et al. 2023) and has probably also affected steppe
existing conservation action, followed by law and bird communities. Remaining livestock are now
policy, land/water protection and species manage- concentrated around human settlements, leading
ment. Recommended actions were for land/water to localized overgrazing. Several steppe birds
management, law and policy, and land/water pro- avoid these areas, although some, such as the
tection, but livelihood, economic and other incen- globally threatened Sociable Lapwing Vanellus gre-
tives were also considered important (Holl garius, have benefitted (Kamp et al. 2011). Other
et al. 2000). Recommended nature conservation threats, such as mining or powerlines, are unlikely
actions tended to focus on slowing deforestation to match agriculture in severity or extent (Kamp
and including alternative sources of income for et al. 2016).
local communities (Holl et al. 2000), although the Kazakhstan has an ambitious and growing pro-
former may not of course be directly beneficial for tected area programme (Baumann et al. 2020),
grassland birds. External capacity building and while also managing individual species through
education were a complementary action suggested action plans (e.g. Sheldon et al. 2012). Intensifica-
in Belize (Piaskowski et al. 2005). tion of existing cropland with minimum agro-
chemical use is recommended, instead of
recultivating abandoned fields; that is, land sparing
Central Asia (44 studies)
rather than sharing (Kamp et al. 2011). Other rec-
The steppes of this region are still among the ommendations include restocking abandoned pas-
largest Palaearctic grasslands, but vast areas were tures with free-ranging livestock (Kamp
converted to cropland before 1900, with another et al. 2015). Populations of wild ungulates, such as
46 million hectares during the Soviet era 1953– Saiga Antelope Saiga tatarica, should be restored
60 (Wesche et al. 2016), resulting in steppe bird to re-establish natural grazing levels (Kamp
declines and the probable extinction of the et al. 2016). Although some issues such as the
Slender-billed Curlew Numenius tenuirostris impacts of climate change and human activity on
(Buchanan et al. 2018). Conversely, the collapse grassland productivity have been studied on Cen-
of the Soviet Union in 1991 led to the abandon- tral Asian grasslands (Chen et al. 2020), most
ment of 15 million hectares of cropland in grasslands outside Kazakhstan are poorly studied
Kazakhstan alone (Kamp et al. 2011). Popula- for their bird communities, and threats to birds in
tions of many steppe birds consequently recov- the dry lowland grasslands of Turkmenistan and
ered where abandoned croplands reverted to Uzbekistan, and high-altitude pastures of Kyrgyz-
steppe (Kamp et al. 2011) but abandoned fields stan and Tajikistan, require assessment.
marked threat to grasslands in the Middle East, stocking rates, which can increase erosion and in
primarily through desertification, leading to a some cases invasion by exotic plants, can result in
reduction in biodiversity, agricultural productivity grassland degradation. The reported threats and
and ecosystem services (Mirzabaev et al. 2019). actions show some geographical variation. The
Increasing road infrastructure across Iran, Turkey Great Plains of Canada and the USA represent the
and Syria is also fragmenting the land and facilitat- largest zone – upland songbirds here are strongly
ing human access and associated disturbance to affected by habitat loss to agriculture, and this area
once remote areas (Bowden 2015). is also experiencing significant renewable and non-
Existing actions for grassland birds include inter- renewable energy development impacts (Daniel &
national management planning for threatened spe- Koper 2019). In the eastern grasslands, woody
cies such the Northern Bald Ibis (Bowden 2015), encroachment is a threat to grassland birds and
Sociable Lapwing (Sheldon et al. 2012) and Saker can follow agricultural abandonment (Monroe
Falcon (Kovacs et al. 2014). The designation of et al. 2019). The western zone includes extensive
protected areas has been initiated, but manage- sagebrush-steppe habitat; threats include wildfire,
ment of these is typically under-resourced and invasion by non-native grasses, shrub encroach-
poorly regulated (McGrady 2018). Important Bird ment, expansion of agriculture and urban areas, oil
Areas have been identified (Evans 1994), but this and gas drilling, and mining (Runge et al. 2019).
programme requires a comprehensive update in The southern zone includes shrubs and desert hab-
this region. itat and is of critical importance for overwintering
Recommended actions include a greater empha- grassland birds, but their wintering ecology is rela-
sis on sustainable livestock management, perhaps tively poorly studied (Macıas-Duarte et al. 2017).
including the introduction of rotational grazing sys- The southern zone is experiencing rapid habitat
tems using traditional livestock. The establishment loss and populations of grassland birds that over-
of more protected areas, improved management of winter in the Chihuahuan Desert are considered
existing ones and improved hunting legislation are to be declining more rapidly than other North
also considered necessary (Brochet et al. 2019). American grassland birds (Macıas-Duarte
The development of more international and et al. 2017). Relatively few publications reported
national species and habitat action plans with the climate change as a threat, although some high-
necessary financial and human resources to imple- elevation or restricted-range populations may be
ment them would make a substantial contribution susceptible to climate change (Pierce et al. 2019).
to grassland bird conservation across the Middle Land/water management, particularly livestock
East. grazing at appropriate densities (Campomizzi
et al. 2019), was the most common existing action
in North America, followed by land/water protec-
North America (101 studies)
tion, and law and policy. Direct species manage-
North American grasslands occupy large areas of ment was less common. The wider Conservation
the continental interior and grassland types vary Reserve Program of the USA, established to recon-
regionally with environmental gradients in temper- vert croplands to perennial grasses, has reduced
ature, rainfall and elevation (Comer & Hoag- soil erosion, improved water quality, and restored
land 2020). North American prairies evolved in and protected wildlife habitats (Harryman
the presence of natural disturbances such as et al. 2019), and many bird species have conse-
lightning-driven fire and grazing by native ungu- quently benefited (Reiley et al. 2019, Shew
lates, which are important for maintaining spatio- et al. 2019).
temporal heterogeneity, but the consequences of Grassland bird conservation in much of this
altering these has been increasingly recognized region will depend on managing disturbance regi-
(Campomizzi et al. 2019). Agriculture, mainly the mens, as well as protection for extant grasslands.
expansion of row cropping and livestock grazing, is Recommended actions focus on land/water man-
the most frequently reported threat across these agement, followed by land/water protection. Pre-
grasslands, followed by energy production and nat- scribed burning, plus grazing by native or domestic
ural systems modification (Ogden et al. 2019). ungulates at appropriate densities could increase
Both cessation of livestock grazing, which can habitat heterogeneity (Campomizzi et al. 2019).
result in decreased plant diversity, and heavy Minimizing road expansion, restricting noise
Central America, the threat from reforestation require better enforcement, especially on private
reflects natural succession and encroachment due lands (Azpiroz et al. 2012).
to agricultural abandonment, other management Species within grassland bird communities have
effects or climate change (O’Connor et al. 2014, varying ecological needs (Maphisa et al. 2017,
Stanton et al. 2018a), whereas in Europe, Asia, Elliott & Johnson 2018, Budka et al. 2019), often
Australasia and South America it mainly results making management challenging. Our results gen-
from plantations (Mortelliti & Lindenmayer 2015, erally suggest that management should aim to pro-
Phifer et al. 2017, Douglas et al. 2020, World Eco- vide a mosaic of habitat types with different grass
nomic Forum 2022). height and cover (Maphisa et al. 2019), to benefit
Among conservation actions, authors of publica- the greatest range of species; this can be facilitated
tions make particularly strong calls for land/water by ensuring the largest areas possible are under
management and land/water protection. The par- appropriate management, to capture natural vari-
lous state of grassland birds globally suggests that ability in habitat types. Even so, the potential for
existing conservation actions for grassland birds are trade-offs exists, both in terms of how species will
globally inadequate. Below, we expand on these respond to grassland threat (responses may not all
recommendations and make further suggestions for be negative) and conservation actions (responses
conservation action and research. may not all be positive).
A recurring recommendation for land/water The large extents of grasslands, the relative scar-
management in our sample of studies is to restrict city and/or limited distribution of some grassland
livestock grazing to sustainable densities. These bird species, and species-specific variation in corre-
densities are, however, rarely, if ever, stated (see lates of occurrence, mean that spatial models of
Future action for grassland bird conservation). grassland and bird species distributions and threats
Another recurring management recommendation may be required to effectively direct conservation
is to use fire appropriately. Prescribed fire is often efforts (Niemuth et al. 2017). Further, the exten-
at frequencies far above natural (lightning-driven) sive scale of grasslands and global extent of this
cycles (Manry & Knight 1986), leading to habitat conservation issue mean that co-ordinated conser-
degradation, proliferation of exotic grasses and vation delivery, supported by the right policies,
greater future fire risk. Again, appropriate fire regi- will be needed at local, regional, national and
mens are rarely stated (see Future action for grass- international scales, covering single species or
land bird conservation). multi-species assemblages, and multiple stages of
In terms of land/water protection, the reported the annual cycle for migratory species (Vickery
area of grasslands under protection is consistently et al. 1999, McCracken 2005, Azpiroz et al. 2012,
low across regions. It is estimated that only about Lark 2020), and sometimes in grasslands in differ-
3% of grasslands in North America ent hemispheres for individual species (e.g. Alfaro
(McCracken 2005), 2–4.5% in southern Africa et al. 2019). In some regions, lack of achievement
(Maphisa et al. 2017) and 1% in southeastern may be related to lack of co-ordination in promot-
South America (Azpiroz et al. 2012) are pro- ing conservation, and public and political support
tected. In areas such as the alpine grasslands of the (Azpiroz et al. 2012).
Qinghai-Tibetan Plateau in Asia, protected area The global threat of agriculture to grassland
coverage of biodiversity hotspots (26% of hot- birds indicates the need for solutions within the
spots) is considered inadequate (Su et al. 2019). agricultural industry itself, and at a global scale.
Author recommendations are to expand protected Incentivizing low-intensity agriculture on grass-
area coverage but also to ensure greater effective- lands is central to conservation in Europe but is
ness by targeting biodiversity hotspots and other little used elsewhere (but see South America
important areas (Su et al. 2019). Authors recom- (Azpiroz et al. 2012)). Incentivizing lower-yield
mend that this protection needs to be accompa- food production could, however, have unin-
nied by appropriate management and sufficient tended consequences: if food production drops in
operating funds, as simply delineating protected one region globally, demand will probably be
habitat is no guarantee of species’ population via- met elsewhere in another, leading to intensifica-
bility in landscapes where agricultural or livestock tion and habitat loss there (Tanentzap
production continues to predominate (With et al. 2015). One suggestion is that conservation
et al. 2008). Legal measures to protect grasslands efforts should focus on the most intensive
(‘profit-maximizing’) food producers, using direct conclusion regarding the importance of agriculture
economic incentives that ‘foster the culture of in explaining the decline of grassland birds, it is a
grassland preservation and restoration’ and problem across scientific disciplines and we
include Ecosystem Services Payments (Drum encourage the publication of non-significant results
et al. 2015). Optimizing agriculture–biodiversity by researchers as long as the studies in question
trade-offs through land sparing or sharing is had sufficient power to detect effects of manage-
likely to be landscape dependent where relevant ment. We also note that simply counting conserva-
to grasslands (Macchi et al. 2020); both sparing tion actions can be misleading, as conservation
and sharing have been found to be required in programmes can vary widely in scope. Different
the Eurasian steppe (Kamp et al. 2015) whereas discrepancies in literature availability among
land sparing yields the best outcomes for farm- regions might also have affected our results. We
land (including grassland) biodiversity in eastern could only review publications in English, and
Europe (Feniuk et al. 2019). Authors argue that publications in other languages influence conserva-
measures producing significant losses to private tion worldwide. Further, research topics change
landowners should include economic compensa- over time as cultures and knowledge change;
tion (Conservation Evidence 2020). because we tried to sample similar numbers of
Activities that garner greater public support studies among regions rather than similar periods
could have direct and indirect benefits, influencing of time in which research was conducted, changing
consumer purchasing for sustainably sourced grass- ecological ‘fashions’ might result in apparently dif-
land products and encouraging corporate responsi- ferent ecological foci in regions with differing
bility (Lark 2020), now well established for forest numbers of studies. Our study design did not
protection (e.g. https://www.rainforest-alliance. allow us to distinguish the reasons for different
org/). The high biodiversity value of grasslands research foci among regions. However, the out-
means that ecotourism could help to provide an come is still of significance to conservation; where
incentive for protecting land; in some cases flag- less research is being conducted, conservation
ship mammals rather than birds may generate the actions must depend on older publications, per-
greatest tourist interest (Maude & Reading 2010). haps affecting conservation strategies. We also
Agrochemical (pesticide) use was flagged as a argue that our overall conclusion regarding the rel-
notable concern in Europe but less so in other ative importance of agriculture is likely to be inde-
regions, although some studies have highlighted pendent of our review methods and sample sizes.
the potential role of pesticide use as a contribu- Among the regions with the most studies, such as
tory factor to grassland bird declines in North North America and Europe, it is clear that agricul-
America (Stanton et al. 2018b, Rosenberg ture, rather than climate change, has posed the
et al. 2019). In some regions, such as North most significant recent threats; further, the relative
America, grassland bird populations do not importance of agriculture is clear among regions,
appear to be strongly regulated by bottom-up regardless of sample size. It seems unarguable that
processes that might relate to reduced food sup- agriculture is a key factor in the declines of grass-
ply (Wiens & Rotenberry 1979, Martin land birds globally.
et al. 1998). However, this surely varies region-
ally, and it seems likely that invertebrate declines
FUTURE ACTION FOR GRASSLAND
have contributed to declines of insectivorous birds
BIRD CONSERVATION
in some systems including European grasslands
(Bowler et al. 2019, Tallamy & Shriver 2021). Our review suggests an immediate global need for
We also note that any large-scale review is sen- (1) an end to conversion of natural and semi-
sitive to potential biases in the literature available natural grassland to other habitats including crop-
for review, and there are several methodological ping, intensively managed grassland for livestock
issues to bear in mind when evaluating our conclu- rearing and plantation forestry; and (2) urgent
sions. Publication bias often results in the absence establishment, and effective enforcement, of larger
of non-significant results from publicly available protected areas of grassland through national and
literature, and this problem plagues any broad- international legislation.
scale literature review. Although we think it is Although agriculture is clearly an overriding
unlikely that this problem influenced our global influence on grassland birds, there is still
much to learn through further research to aid of this study and many others remind us that
development of effective conservation action and grassland birds are at significant risk from
policies. We propose that the following are anthropic activities, and require urgent research,
research priorities across most grasslands globally: monitoring and action, underpinned by effective
(1) identification of the densities of native and policies, to help mitigate the plummeting popu-
non-native grazing animals that are required to lations of many grassland species worldwide.
maintain grasslands in the optimum condition for
grassland birds; and (2) in regions where natural
We thank Candace Briggs, Grace Fedirchuk and
fire regimens have shaped grasslands, identifica- Rochelle Johnson for assistance with the research, and
tion of the prescribed fire regimens that are two anonymous reviewers who greatly improved the
required to maintain grasslands in the optimum manuscript. The research did not receive any specific
condition for grassland birds; (3) the extent of grant from funding agencies in the public, commercial
species-specific variation in the ecological require- or not-for-profit sectors.
ments of grassland birds and hence responses to
interventions. This variation may require the AUTHOR CONTRIBUTIONS
maintenance of heterogeneous grassland habitats
(Fuhlendorf et al. 2010); (4) further testing of David Douglas: Conceptualization; formal analysis;
whether land sparing or sharing optimizes grass- methodology; validation; visualization; writing –
land bird conservation, particularly outside the original draft; writing – review and editing. Jessica
Eurasian steppe and Europe, where some previ- Waldinger: Data curation; formal analysis; valida-
ous work has been undertaken; (5) understanding tion; visualization. Zoya Buckmire: Data curation.
the extent to which invertebrate and grassland Kathrynn Gibb: Data curation. Juan Medina: Data
bird declines are linked; and (6) a better under- curation. Lee Sutcliffe: Data curation. Christa
standing of the mechanistic effects within grass- Beckmann: Validation; writing – original draft;
lands that will undoubtedly accompany climate writing – review and editing. Nigel Collar: Valida-
change. Both (1) and (2) could be tested tion; writing – original draft; writing – review and
through large-scale manipulations of grazing den- editing. Raymond Jansen: Validation; writing –
sity and/or fire, and their interaction (Fuhlendorf original draft; writing – review and editing.
et al. 2010), with consideration of other inter- Johannes Kamp: Validation; visualization; writing
ventions such as restoration of grassland where it – original draft; writing – review and editing. Ian
has been converted to cropping. Where grassland Tchagra Little: Validation; writing – original draft;
restoration is implemented (e.g. Society for Eco- writing – review and editing. Rob Sheldon: Valida-
logical Restoration 2022), robust monitoring of tion; writing – original draft; writing – review and
biodiversity outcomes is essential for assessing editing. Alberto Yanosky: Validation; writing –
success. Other priorities vary regionally. In many original draft; writing – review and editing. Nicola
regions, there are vast areas encompassing many Koper: Conceptualization; methodology; resources;
countries where the status of grassland birds and supervision; validation; writing – original draft;
the threats they face are poorly known; a nota- writing – review and editing.
ble example is Central Asia outside Kazakhstan.
There is also significant disparity in the utiliza-
CONFLICT OF INTEREST
tion and understanding of the effectiveness of
STATEMENT
AES. Where AES have been widely applied in
Europe, further research is needed to better The authors have no conflict of interest to declare.
understand their efficacy (Kelly et al. 2021a,
2021b). Outside Europe there have been numer-
FUNDING
ous pilot projects and interest in introducing
AES, and the extensive experience already None.
obtained from European application of these sys-
tems could provide learning opportunities for
ETHICAL NOTE
designing effective and large-scale programmes
elsewhere. Ultimately, we note that the results None.
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landscapes, South Africa. Avian Res. 8: 11. rent study.
Varun, K. & Dutta, S. 2020. Understanding land-use Table S5. Conservation actions for grassland
response of bird communities in an arid ecosystem of India. birds used in current study.
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