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Biological Conservation 296 (2024) 110655
0006-3207/© 2024 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
Navigating complex geopolitical landscapes: Challenges in conserving the
endangered Arabian wolf
Gavin T. Bonsen
a
,
*
, Arian D. Wallach
a
,
1
, Dror Ben-Ami
b
, Oded Keynan
b
,
c
, Anton Khalilieh
d
,
Yara Dahdal
d
, Daniel Ramp
a
a
Centre for Compassionate Conservation, TD School, University of Technology Sydney, New South Wales 2007, Australia
b
Compassionate Conservation Middle East, Steinhardt Museum of Natural History, Tel Aviv University, Israel
c
Dead Sea and Arava Science Centre, Central Arava Branch, Hazeva, Israel
d
Nature Palestine, Ramallah, Palestine
ARTICLE INFO
Keywords:
Middle East
Predator conservation
Desert ecology
Human-wildlife coexistence
Conservation conicts
ABSTRACT
Grey wolf (Canis lupus) populations are beginning to increase globally after centuries of decline. While protective
legislations and policy implementations have been driving this increase, evidence suggests that these work
because of a general rise in public acceptance of wolves. As people have become more knowledgeable of the
important ecological roles played by wolves, protection has gained increasing community support, with human-
wolf coexistence now being achieved in some areas. However, this is not universal, and some subspecies remain
endangered. This is the case for the little-known Arabian wolf (Canis lupus arabs), which inhabits arid regions of
the southern Levant and Arabian Peninsula: a geopolitically diverse region crossing multiple jurisdictions with
disparate cultures, legislations, and attitudes towards wildlife and conservation. Here we review global efforts
within the wolf conservation sphere to explore potential opportunities and challenges in the Middle East,
acknowledging and accounting for the unique geopolitical complexity of this region. We bring together what is
known about this wolf's taxonomic status, distribution, ecology, and conservation across eleven countries in
which it resides, collating decades of legislation, governmental, and non-governmental conservation efforts, and
summarising literature pertinent to the Arabian wolf. Cross-border collaborations remain challenging in the
Middle East, but we identify practical and culturally-based solutions that may improve coexistence and con-
servation goals, building upon areas where coexistence already occurs. We write this review in the hope that it
will highlight the synergies, opportunities, and obstacles that require more serious deliberation and collaboration
to conserve one of the region's remaining apex predators.
1. Introduction
Large predators like the grey wolf (Canis lupus) have experienced
major decline and range contraction over the last few centuries, pri-
marily because of persecution resulting from efforts to protect livestock
(Ripple et al., 2014). The loss of large predators has wide-ranging and
cascading ecological effects, contributing to what some describe as
‘trophic downgrading’ (e.g., Estes et al., 2011). In response to this,
conservationists have focussed on increasing legislative protection, and
prioritising ecological restoration (i.e., rewilding) and reintroduction
programs. The result of these programs is that global wolf populations
are beginning to increase as protective legislation and policy imple-
mentations have proven successful in supporting recovery (Boitani,
2003; Mech, 2017), while public support for tolerance of predators like
wolves has increased (Treves and Bruskotter, 2014). Yet protection of
wolves is not universal across their wide range as conict remains and
conservation actions are lacking in some areas. This is particularly the
case in Asia, where wolves do not get the same attention as their Eu-
ropean and North American counterparts (Chakrabarti et al., 2023).
Seven subspecies of grey wolf have been described across Asia
(Nowak, 2003), and though there is some conjecture over their status,
they are thought to be extirpated or in decline across much of this range
* Corresponding author.
E-mail addresses: gbonsen@outlook.com.au (G.T. Bonsen), a.wallach@qut.edu.au (A.D. Wallach), drorbe@post.tau.ac.il (D. Ben-Ami), oded@adssc.org
(O. Keynan), Daniel.Ramp@uts.edu.au (D. Ramp).
1
Present address: School of Biological and Environmental Science, Faculty of Science, Queensland University of Technology, Brisbane, QLD, Australia.
Contents lists available at ScienceDirect
Biological Conservation
journal homepage: www.elsevier.com/locate/biocon
https://doi.org/10.1016/j.biocon.2024.110655
Received 29 July 2023; Received in revised form 26 April 2024; Accepted 25 May 2024
Biological Conservation 296 (2024) 110655
2
(Boitani, 2003; Chakrabarti et al., 2023). At the south of their distri-
bution in the Middle East, two subspecies meet – the ‘Indian wolf’
(traditionally described as C. lupus pallipes but see a recent genomic
study from Hennelly et al., 2021) and the Arabian wolf (C. lupus arabs).
The Indian wolf has a broad distribution, encompassing much of
southern and western Asia from India to Turkey, and favours Mediter-
ranean or temperate climates. The Arabian wolf, on the other hand,
inhabits arid environments of the southern Levant and Arabian Penin-
sula. We focus this review on the ecology and conservation of this
unique desert-adapted subspecies threatened with extinction (Mallon
and Budd, 2011).
The increase in wolf populations in other parts of the world brings
hope. However, the circumstances for the Arabian wolf differ in two
important ways: it resides in ecosystems that are far less productive than
the temperate ecosystems of Europe and North America (Barocas et al.,
2018), and the geopolitical mosaic and distinct forms of human land use
in the Middle East present unique challenges. Hence, to align with the
UN's push to promote transboundary cooperation in biodiversity con-
servation (United Nations General Assembly, 2021), policy initiatives
targeting the conservation of the Arabian wolf require careful and
culturally appropriate consideration. Today, Arabian wolves occur
within multiple jurisdictions shaped by disparate cultures that manifest
various attitudinal contexts towards predators. Despite this, apart from a
single recent study (Bonsen et al., 2022), almost all previous research on
relationships between Arabian wolves and people has been limited to
individual jurisdictions. Here, we synthesise available information
across the entire distribution of the Arabian wolf to highlight current
knowledge and identify challenges, pitfalls, and gaps in knowledge, with
the aim to improve awareness and stimulate robust conservation ini-
tiatives for this endangered wolf.
To begin this process, we draw upon conservation experiences of
wolves in other regions to understand the history, including the drivers,
mechanisms, failures, and successes, of wolf conservation. We
acknowledge that each region will give rise to its own unique opportu-
nities and insights, but we start by examining global wolf conservation
efforts, with a focus on social and political factors that have facilitated
wolf recovery. We then review what is currently known about the
Arabian wolf, including its taxonomic status, ecology, distribution, and
how people's values and attitudes towards wolves differ across the
Middle East. We end by articulating how cross-jurisdictional planning
and collaboration are vital to ensure the ongoing conservation of this
keystone and iconic subspecies of grey wolf.
2. Methods
We reviewed the scientic literature, governmental and non-
governmental reports, and policy implementations to outline key pat-
terns and motivations behind successful wolf protection efforts in
Europe and North America. We searched Google Scholar using combi-
nations of keywords related to key factors associated with wolf conser-
vation (e.g., human attitudes towards wolves; tolerance of wolves; wolf
protection/conservation in Europe/North America; trophic cascades
driven by wolves), and searched Google for any regional reports and
policy implementations. We then brought together the literature, where
available in English, related to wolf conservation and protection within
the Middle East. We acknowledge, of course, that knowledges occur
other than those encountered in English literature; hence, our work
summarises existing knowledge available in English academic literature
and can be used to stimulate discourse and engagement with other
knowledges.
To put this into context for the Arabian wolf, we also searched Google
and Google Scholar for any scientic and grey (e.g., web articles, news
articles) literature pertaining to wolves in the Middle East, using both
scientic (Canis lupus [arabs/pallipes]) and common (English: wolf,
Arabian wolf, Indian wolf; Arabic: ﺫﺉﺏ , “the’eb”; Hebrew: זאב , “ze’ev”)
names, and browsed books and conservation reports containing
information about wolves and other predators in the Middle East. To
focus this review on the Arabian wolf and discern between other sub-
species such as the Indian wolf, we collated available genetic and
morphological studies of both Arabian and Indian wolves and compared
these records to arid zone borders in the countries these wolves were
recorded. From this, we hypothesised a potential distribution of the
Arabian wolf, as distinct from the Indian wolf.
We then compiled an in-depth overview of governmental and non-
governmental conservation efforts and legislation relating to wildlife
protection across the range of the Arabian wolf. Information was again
gathered from scientic and grey literature, governmental and non-
governmental reports, and websites. Some websites were either no
longer active or had not been updated for several years. In the case of
legal frameworks, individual legislations were veried against ofcial
documents or their translated versions. The FAOLEX database on the
Food and Agriculture Organisation of the United Nations (FAO) website
(http://www.fao.org/faolex/en/) was used as the primary repository for
sourcing legislation as it was the most comprehensive. However, for
some nations the full suite of legislation did not feature in this database,
so additional legislations were found in governmental reports.
3. Lessons from global conservation efforts
3.1. Ecology and importance of wolves globally
Emerging as arguably the most important body of research to inspire
appreciation of wolves and conservation initiatives to protect them over
the last few decades, their ecological roles as apex predators and trophic
regulators has increasingly been recognised as vital for the functioning
and regulation of many ecosystems (Ripple and Beschta, 2011, 2004;
Terborgh and Estes, 2010; Wallach et al., 2015b). As large-bodied apex
predators, wolves can have cascading effects on ecosystems, either
through direct predation or by indirectly inuencing biota in lower
trophic levels. These top-down forces affect the abundance and modify
the behaviour of lower-trophic animals (Ripple and Beschta, 2011;
Wallach et al., 2015b), in turn inuencing a broad range of biotic and
abiotic processes (Ripple and Beschta, 2004). The suppression of this
function through the removal of wolves results in a series of cascading
effects that can ultimately lead to species loss, functional shifts in
community complexity, and declines in vegetation productivity (Ripple
and Beschta, 2011). The most high-prole example of this comes from
Yellowstone National Park in North America, where ecologies trans-
formed following the reintroduction of previously extirpated wolves
(Ripple and Beschta, 2011, 2004).
Trophic cascades are, however, more complex than these simple,
albeit signicant, three-level linear chains of species interactions.
Rather, they can involve multifaceted networks of species with various
types and strengths of interactions (Wallach et al., 2016). For example,
vegetation recovery in Yellowstone promoted the return of several
species (Ripple and Beschta, 2011), and ultimately increased the
complexity of interactions occurring there (Smith et al., 2003; Beschta
et al., 2023). Wolves also interact with and shape the distribution and
behaviour of other predators, through intraguild predation (Polis et al.,
1989), interspecic competition (Berger and Gese, 2007), and predation
risk effects (Suraci et al., 2016). One implication of this is that they
suppress populations of medium-sized predators (i.e., mesopredators;
Ripple et al., 2013), and the cascading effects of ‘mesopredator release’
have been documented in places where wolves have been suppressed –
as evidenced by continental range expansions of coyotes (Canis latrans)
in North America and jackals (Canis aureus) in Europe with widespread
wolf removal (Krofel et al., 2017; Newsome et al., 2017).
3.2. Conict, persecution, and mitigation
Wolves have experienced persecution and extirpations during the
last several centuries across Eurasia and North America. Various
G.T. Bonsen et al.
Biological Conservation 296 (2024) 110655
3
countries in Central and Western Europe eradicated wolves in the 16th
and 17th centuries, and by the early 20th century, wolves were nearly
extinct across most of Europe. This pattern was repeated in North
America (Boitani, 2003). While the International Union for the Con-
servation of Nature (IUCN) now lists the grey wolf as least concern
(Boitani et al., 2018), several populations and unique subspecies remain
imperilled. Included in these are the three most southern subspecies, all
listed as endangered: the Arabian wolf (Mallon and Budd, 2011) and
Indian wolf (Sharma et al., 2019) of Eurasia, and the Mexican wolf (C. l.
baileyi) of North America (Paquet et al., 2001).
Along with loss of habitat and declining prey species, pervasive le-
thal control and exclusion of wolves from agricultural landscapes have
exacerbated this demise. For millennia, predators like wolves have
conicted with people and their interests (Fritts et al., 2003). At the turn
of the 21st century, livestock depredation was the most frequently re-
ported reason for conict between humans and wolves (Sillero-Zubiri
and Laurenson, 2001). Targeting of wolves and other predators can also
be due to other interests related to aesthetics (Childes, 1988), fear of
threatening encounters (Linnell et al., 2003), conservation concerns
(Cohen et al., 2013), or simply because of perceptions that they ‘don't
belong’ (Van Dooren, 2011; Wallach et al., 2015a). As a result of such
conicts, large predators continue to experience substantial population
decline and range contraction (Ripple et al., 2014).
One conservation approach used globally as an attempt to combat
such losses is land sparing (i.e., the setting aside of land for conserva-
tion). During the last 100 years, more than 200,000 terrestrial protected
areas such as national parks and nature reserves have been established
worldwide, covering around 17 % of the Earth's terrestrial land surface
(UNEP-WCMC and IUCN, 2023). However, this strategy of ‘separating
people from nature’ is not sufcient to protect wide-ranging species
from hunting and persecution, particularly those whose home ranges
extend beyond protected area boundaries and into agricultural land-
scapes, as is often the case with large predators (Johansson et al., 2016).
Protected areas are also not ‘pristine’ environments outside of human
inuence. Management often involves pervasive and intrusive measures
that disturb the ebb and ow of ecosystems: lethal control and harvest of
unwanted or abundant species can cause cascading ecosystem effects
(Colman et al., 2014); tourism and recreational activities bring pollu-
tion, environmental degradation, and wildlife disturbance (Pickering
et al., 2003); and roads cut through protected areas, causing vehicular
collisions with wildlife (Ramp and Ben-Ami, 2006; Roger et al., 2012),
habitat fragmentation (Roger et al., 2011), and edge effects (Ben-Ami
and Ramp, 2006). As such, other forms of human-caused wildlife mor-
tality may not be altered by protection, and in some cases can be higher
than non-protected areas (Hill et al., 2020). Recognition of the short-
comings of protected areas has led to broader social initiatives linked to
coexistence and land sharing (Fischer et al., 2014).
For wolves, the frontline of coexistence is often within agricultural
landscapes. Predator-friendly farming is an all-encompassing term for
non-lethal strategies that farmers can use to protect livestock from
depredation (Johnson and Wallach, 2016), and it has been implemented
to ease tensions between farmers and wolves (Bogezi et al., 2019). In
both modern agricultural and traditional pastoralist systems of livestock
production, the move by some to non-lethal predator control has
enabled human-predator coexistence in intensive livestock growing re-
gions (Ohrens et al., 2019; Stone et al., 2017). Non-lethal methods often
combine improved livestock husbandry practices with tools that act as
deterrents to prevent encounters between livestock and predators
(McManus et al., 2015). Livestock guardian dogs have been used for
millennia (Gehring et al., 2010) and continue to be an effective deterrent
against predators (van Eeden et al., 2018), while technological de-
terrents such as adry, livestock collars, and ashing lights also show
promise in reductions in livestock depredations (McManus et al., 2015;
Miller et al., 2016; Ohrens et al., 2019). Compensation schemes offered
by governmental agencies or non-governmental organisations (NGOs)
have also relieved agriculturalists from potential hardships associated
with predator-related livestock loss (Naughton-Treves et al., 2003).
Such strategies promoting human-predator coexistence have been
gaining traction, particularly in regions with targeted public education
and conservation programs aimed towards increasing acceptance of
predators (Wallach et al., 2015a).
3.3. Human attitudes and social research
As awareness and knowledge of the ecological roles of wolves have
increased globally, so have intentions and efforts to conserve them.
North America and Europe are prime examples, where wolf populations
have made an incredible comeback thanks to efforts over the last 25
(Jimenez et al., 2017; Mech, 2017) and 40 (Chapron et al., 2014; Randi,
2011) years, respectively. Wolf populations are now recovering in seven
states in the US (Mech, 2017), ending more than 60 years of extirpation
(Jimenez et al., 2017). While protected areas played a large part in this
success story, similar recoveries have been documented in Europe where
recovery has been less reliant on protected areas (Boitani, 2003; Mech,
2017). Across much of Central and Western Europe and Scandinavia,
wolves have recovered since the 1970s (Chapron et al., 2014), particu-
larly in refugia where they were able to maintain resilient populations
(Salvatori and Linnell, 2005). The recovery of European wolf pop-
ulations can largely be attributed to the development of transboundary
policy initiatives (Epstein et al., 2016; Linnell and Boitani, 2012), where
coordinated legislation is enforced across the continent through inter-
national collaboration (Linnell et al., 2008). However, the derivation of
these policies and their ability to protect wolves is fundamentally a
result of a positive shift in public sentiment towards wolves (Bencin
et al., 2016; Kansky and Knight, 2014; Treves and Bruskotter, 2014).
Conservationists and wolf advocates have long utilised social
research to assist with large carnivore conservation efforts (Bruskotter
and Wilson, 2014; Carter et al., 2012). Human attitudes towards pred-
ators, which range from acceptance to intolerance, are primary de-
terminants of the success of long-term conservation of wolves
(Bruskotter et al., 2014). Attitudes towards wildlife are shaped by social
and cultural factors (Arbieu et al., 2019; Sagie et al., 2013), such as
educational (Holsman et al., 2014; Manfredo et al., 2003; Williams et al.,
2002) or socio-economic (Rode et al., 2021) differences, and vary across
spatial (Bencin et al., 2016; Karlsson and Sj¨
ostr¨
om, 2007) and temporal
(Fern´
andez-Llamazares et al., 2020; George et al., 2016) scales. For
example, positive attitudes towards wolves prevail in people that live
further from wolf territories (Ericsson and Heberlein, 2003; Karlsson
and Sj¨
ostr¨
om, 2007) and have access to higher levels of education
(Manfredo et al., 2003; Naughton-Treves et al., 2003; Williams et al.,
2002). Conversely, people that live near wolves and have had direct
experiences with wolves (e.g., through livestock loss or threatening
encounters) tend to hold more negative attitudes towards wolf recovery
(Ericsson and Heberlein, 2003; Karlsson and Sj¨
ostr¨
om, 2007).
Overall, public attitudes towards large carnivores shifted signi-
cantly during the latter half of the 20th century (Kellert et al., 1996). A
driving factor in this shift is the transition of wildlife value orientations
from dominion to mutualism (Manfredo et al., 2009); where dominion
holds that wildlife exist for human use, mutualism emphasises the
interconnectedness of wildlife and humans, holding that wildlife are
deserving of care and compassion (Ramp and Bekoff, 2015). While such
values have a long history in certain spiritual domains (Yeshey et al.,
2023), these changes have been particularly prevalent in Western post-
industrialised cultures after WWII (Inglehart and Baker, 2000), and
generally in social groups with higher levels of education (Williams
et al., 2002). Although general education has contributed to such
change, carnivore education focussed on tolerance and ecological roles
has been important for increasing support for predator conservation
(Arbieu et al., 2019; Bruskotter and Wilson, 2014). It remains common,
however, for predators such as wolves to be viewed with widespread
hostility in regions where knowledge of their ecological importance, low
threat to human life, and alternative, non-lethal, methods of livestock
G.T. Bonsen et al.
Biological Conservation 296 (2024) 110655
4
protection are lacking (Seddon and Khoja, 2003). It is often these regions
where people generally have strong adherence to social norms, which
can exacerbate cultural resistance to the acceptance of novel beliefs and
practices.
4. State of knowledge of the Arabian wolf
4.1. Taxonomy of Canis lupus arabs
Taxonomic debate about Middle Eastern wolves is evidenced by
different perspectives in the literature (Ak and Alkon, 1983; Cun-
ningham and Wronski, 2010; Mukherjee et al., 2009; Reichmann and
Saltz, 2005; von Jaffa, 2013; Wronski and Macasero, 2008). However,
most generally accept that two subspecies of wolf occur in the Middle
East. The Indian wolf occurs in temperate regions characterised by
Mediterranean or semi-arid climates (Ferguson, 2002; Khosravi et al.,
2013), while the Arabian wolf inhabits the deserts of the southern
Levant and Arabian Peninsula (Bray et al., 2014; Cohen et al., 2013;
Hefner and Geffen, 1999; Nowak, 2003). Though the two subspecies
share similarities – for example, both being small relative to those in
northern regions (Ferguson, 2002; Nowak, 2003) – Arabian wolves are
noticeably smaller than Indian wolves, with the latter being up to 1.5
times larger than the former (Ferguson, 2002). Clear genetic distinctions
have also been documented between the two subspecies, with the
wolves of Arabia being more closely related to Eurasian wolves (C. l.
lupus) than Indian wolves (Bray et al., 2014).
Despite clear differences in climatic conditions between the distri-
butions of Arabian and Indian wolves, it is unclear where and how
spatial divergence occurs. Limited genetic and morphological evidence
available suggests that primarily Indian wolves occur in Iran (Khosravi
et al., 2012, 2013) and Arabian wolves in Arabia (Bray et al., 2014).
Furthermore, Khosravi et al. (2012) state that wolves are absent from
the central deserts of Iran (i.e., Dasht-e Kavir and Dasht-e Lut); however,
they occur in semi-arid environments (Tourani et al., 2014). With cur-
rent evidence as it stands, it seems plausible that the range of the
Arabian wolf extends more than 3 million km
2
throughout the arid re-
gion southward from the southern deserts of Iraq and possibly Syria,
encompassing the southern Levantine and Arabian deserts (Fig. 1). Some
degree of overlap between the two subspecies may occur in the northern
part of the Arabian wolf's range; however, no empirical data shows
where or if this is the case. Further genetic testing across a broader range
is required to truly understand the divergence between Arabian and
Indian wolves.
4.2. Ecology and importance
Is it necessary to differentiate between the two subspecies? When it
comes to setting conservation goals, it may be. The Arabian wolf is the
only wolf specically adapted to arid environments (Fig. 2). While other
wolves venture into deserts – the Gobi Desert in eastern Asia is home to
populations of the Mongolian wolf (C. l. chana; Kaczensky et al., 2008)
and the Mexican wolf previously occurred in North American deserts
(Hendricks et al., 2016) – none of these subspecies are exclusive to arid
regions, unlike the Arabian wolf. There are certainly similarities be-
tween Arabian and Mexican wolves: the Arabian wolf is the smallest
subspecies in the Old World, while the Mexican wolf is the smallest in
Fig. 1. Regions that are likely to encompass distributions of the two grey wolf subspecies that occur in the Middle East. The Indian wolf (C. l. pallipes) inhabits
temperate climates from India to Turkey while the Arabian wolf (C. l. arabs) resides in the deserts of the southern Levant and Arabian Peninsula (photo credit: N
Dragi´
c and D. Kwan).
G.T. Bonsen et al.
Biological Conservation 296 (2024) 110655
5
the New World (Nowak, 1995). Sadly, the Mexican wolf is also the most
endangered wolf subspecies, and it has been at the centre of conserva-
tion efforts in North America for the last 40 years (U.S. Fish and Wildlife
Service, 2022, 1982). Efforts have included ongoing reintroduction
programs and stringent enforcement of protective legislation (Hendricks
et al., 2016), however, populations of Mexican wolves continue to be
precarious because of a lack of tolerance (Paquet et al., 2001).
There is no doubt that the Arabian wolf is of considerable ecological
and functional importance to the desert ecosystems of the Middle East
(Bonsen et al., 2022). Since the widespread eradication of other large
carnivores throughout its range, the Arabian wolf remains the last sur-
viving apex predator with a wide distribution. Until relatively recently,
the wolf shared this role with two large felids: the Asiatic cheetah
(Acinonyx jubatus venaticus) and the Arabian leopard (Panthera pardus
nimr). Today, with the cheetah extinct throughout the Arabian wolf's
range (Farhadinia et al., 2017) and the leopard likely conned to a few
protected areas (Spalton et al., 2006), the sole remaining large carnivore
to ll a similar niche is the striped hyaena (Hyaena hyaena); albeit not as
apex predator. While the Arabian wolf periodically hunts gazelle
(Gazella spp.), ibex (Capra nubiana), and young onagers (Equus hemi-
onus), striped hyaena are not successful hunters of ungulate prey (Kruuk,
1976). However, both share a largely omnivorous diet. Each actively
hunts small mammals, birds, and reptiles, and scavenges from agricul-
tural crops, carrion, and garbage (Ak and Alkon, 1983; Kruuk, 1976;
Qarqaz et al., 2004; Shalmon, 1986). In regions where wild prey den-
sities are low and free-range livestock farming common, domestic un-
gulates form a considerable part of each species' diet (Qarqaz et al.,
2004; Shalmon, 1986).
The difference between the two lies in their methods of acquiring
prey. The striped hyaena is primarily a scavenger, and mostly feeds on
carcasses of livestock that have died from prior causes (Tourani et al.,
2012). Apex predators like wolves, on the other hand, are known to prey
on livestock (Yom-Tov, 2003), especially where natural prey resources
are diminished or when domestic animals are not adequately secured
(Khorozyan et al., 2015). Throughout most of the Arabian wolf's range,
wild ungulate abundance is low because Middle Eastern arid and hyper-
arid environments couple low productivity with strong hunting pressure
from people, pushing many of the region's ungulate species to extensive
decline or extinction (Mallon and Kingswood, 2001). Local and regional
efforts to revive ungulate populations are largely conned to protected
areas (Amr et al., 2004; Barichievy et al., 2018; Mallon and Kingswood,
2001), leaving few suitable alternatives for large prey, other than live-
stock, outside of protected areas.
For over a thousand years, Middle Eastern deserts have been farmed
by Bedouins: nomadic or semi-nomadic tribal pastoralists whose shep-
herding methods often require them to travel for days on end. Across
much of the region's deserts, Bedouin pastoralism remains the primary
source of agriculture. However, in the latter half of the 21st century,
technological advances in groundwater acquisition and farming prac-
tices saw a rapid increase of agricultural areas in arid regions. These
transformed ‘oases’ provide an abundance of fruits, vegetables, and
water accessible to wolves, reducing their reliance on the ungulate prey
populations (Shalmon, 1986). In such areas, and those where livestock
are well protected, livestock depredation by wolves can be rare (Nem-
tzov and King, 2001). This is the case in Israel's hyper-arid Arava Valley,
where shepherding is minimal, and crop farming is the predominant
form of agriculture (Lewin et al., 2021). Here, wolves have developed
such an afnity for human habitation that several GPS-tracked in-
dividuals were observed to seldom venture more than ve kilometres
from human infrastructure (Barocas et al., 2018). It is possible that their
reduced tendency to hunt large prey may partially explain why current
pack sizes are generally smaller than those reported in Europe and North
America (Hefner and Geffen, 1999), with solitary or pair sightings of
wolves common.
4.3. Distribution and abundance
Arabian wolf numbers vary considerably throughout its wide
Fig. 2. The Arabian wolf commonly uses wadis (dry riverbeds) and springs in Middle Eastern deserts: (a) an Arabian wolf during winter in the Arava Valley of Israel
(credit: N. Dragi´
c), (b) a wadi in Sheizaf Nature Reserve, Israel, and (c) Ein Salamani, a spring in Dana Biosphere Reserve, Jordan.
G.T. Bonsen et al.
Biological Conservation 296 (2024) 110655
6
distribution (Fig. 3). Evidence suggests the Negev Desert of Israel, which
includes the Arava Valley, has the most stable population of Arabian
wolves. The population here was estimated at 90 to 150 individuals a
decade ago (Cohen et al., 2013); thought to be a stable number for an
arid to hyper-arid region with an area of less than 15,000 km
2
. West of
Israel, the wolf's range extends into the Sinai Peninsula of Egypt. While
we were unable to nd published data regarding population estimates or
how far across Sinai wolves occur, they were recently recorded in St
Katherine Protectorate in the south (Gecchele et al., 2017), and they
continue to cross the border from Israel (Barocas et al., 2018).
In the eastern part of its range, the most recently published estimated
population size across the three largest countries on the Arabian
Peninsula (i.e., Saudi Arabia, Yemen, and Oman) was 500–600 in-
dividuals, but declining; however, this was nearly twenty years ago
(Sillero-Zubiri et al., 2004). Wolves appear to be common in remote and
protected areas of Yemen (Khorozyan et al., 2014). In Oman, wolves
have been recorded throughout the country, but again seem to be
conned to remote and protected areas (Mazzolli et al., 2017; Spalton,
2002a). Records are common throughout Saudi Arabia; however, most
of these records derive from killed individuals (Alou and Amr, 2018;
Zafar-ul Islam et al., 2019). The Arabian wolf is now thought to be
locally extinct in Qatar (Mallon and Budd, 2011), the United Arab
Emirates (UAE; Cunningham, 2004), and Kuwait (Mallon and Budd,
2011).
Between the Negev/Arava and Arabian Peninsula populations lies
Jordan, where wolf numbers are unknown, but they are believed to be
rare (N. Hamidan pers. comm.). Past surveys conrmed wolf presence in
north-eastern Badia (Bunaian et al., 2001), and wolves were recently
recorded during camera-trapping and passive tracking surveys in Dana
Biosphere Reserve and Fifa Nature Reserve in Jordan's Wadi Araba re-
gion (Bonsen and Khalilieh, 2021). Jordan provides an important
stepping-stone between the Negev population and the dwindling pop-
ulation of the Arabian Peninsula. However, the most critical jurisdiction
for Arabian wolf conservation is likely to be Saudi Arabia due to its large
area and geographic location. The country stretches 2.15 million km
2
and lies between the populations of the southern Levant (i.e., the Sinai
Peninsula, Israel/Palestine, and Jordan) and the southern and eastern
extremities of Arabia (i.e., Yemen and Oman).
5. Geopolitical diversity
5.1. Conservation of the Arabian wolf
The decline of large carnivores in the Middle East mirrors the global
extirpations and range contractions of large carnivores (Ripple et al.,
2014). Of 20 carnivores known to occur in Arabia, eight have been listed
as threatened or near threatened (Mallon and Budd, 2011): one being
regionally extinct, another critically endangered, and two endangered.
Although the global status for C. lupus is least concern (Boitani et al.,
2018), status reviews for the Arabian wolf over the last few decades list
it as vulnerable (Boitani, 2003), endangered (Mallon and Budd, 2011),
or critically endangered (Benson, 2009). This at least gives it the
advantage of being stringently protected in Israel; aside from the
authorised killing of almost 40 wolves to protect the critically
Fig. 3. Previously estimated population sizes of the Arabian wolf throughout its range (Cohen et al., 2013; Sillero-Zubiri et al., 2004), and its location within the
Middle East (inset).
G.T. Bonsen et al.
Biological Conservation 296 (2024) 110655
7
endangered acacia gazelle (Gazella gazella acaciae) in the mid 2000s, no
legal killing of Arabian wolves is sanctioned (Cohen et al., 2013). While
the subspecies is currently legally ‘protected’ across its range, the reality
of this protection is not homogeneous. In areas of human-wolf conict,
this protection can be rather lax.
Small population sizes and low densities make the Arabian wolf
particularly susceptible to stochastic events (e.g., disturbance, disease)
and changes in protection status. Without concerted and coordinated
conservation efforts, the Arabian wolf might end up following the path
already traversed by the region's large felids. The critically endangered
Arabian leopard (Stein, 2020), which was sympatric to the Arabian wolf,
is now likely conned to small pockets of protected areas in Saudi
Arabia, Yemen, and Oman (Khorozyan et al., 2014; Spalton, 2002b;
Spalton et al., 2006). The leopard's demise is recent, with extirpations in
Israel, the Palestinian Territories, and Jordan likely to have occurred
within the last 20 to 30 years (Farhadinia et al., 2019). The Asiatic
cheetah, on the other hand, has been restricted to central Iran since the
1980s (Farhadinia et al., 2017). Remaining leopard populations are
being closely monitored in the three countries they are known to remain,
with little hope of widespread recovery without systemic changes in
tolerance and accommodation for predators (Farhadinia et al., 2019).
Cheetahs are unlikely to return without the aid of reintroduction pro-
grams (Durant et al., 2017). For canids like wolves, the situation is
potentially less dire as they seem better adapted to human environments
(Barocas et al., 2018).
5.2. Scale of jurisdictional crossover
The Arabian wolf possibly navigates one of the most complicated and
fraught geopolitical landscapes on Earth. Its distribution crosses a socio-
politically complex region with diverse historical and contemporary
cultural heritage. In the current political mosaic, the Arabian wolf's
range covers eleven recognised states, intertwined with swathes of
disputed land. Of course, each nation has its own set of legislations
related to wildlife protection and levels of enforcement. For wildlife
with large home ranges like the Arabian wolf, traversing this complex
geopolitical landscape can be precarious and troublesome as actions and
behaviours in one location may not cause conict with people, but in
another may result in them being shot (Hefner and Geffen, 1999). A
review of the protection status and conservation efforts across different
jurisdictions led to insights that may be used to shape further discussion
(please see Table S1 in Supplementary Material for sources of informa-
tion for this section).
Interest in wildlife conservation has proliferated throughout the
Middle East, with governmental and non-governmental effort
continuing to be established. The UN's Convention on Biological Di-
versity, launched during the 1990s, spurred many nations to become
active participants. By the turn of the century, most nations across the
region had ratied the convention and subsequently produced a Na-
tional Biodiversity Strategy and Action Plan (NBSAP), with all eleven
nations now having concrete plans to conserve biodiversity. These
involve dening pitfalls within the current legal and institutional
frameworks and developing solutions to improve strategies for biodi-
versity conservation. It has also led to many governments forming
focused ministries or agencies to act as authorities for environmental
affairs.
Although protective legislation now exists throughout the region,
awareness and enforcement vary considerably across and within state
boundaries. An overview of the legal frameworks, governance, and
conservation efforts within each country highlights key differences and
similarities, pertinent to the conservation of the Arabian wolf (Table 1).
Currently, all countries have legislation that prohibits or regulates
hunting. It is universally illegal to hunt or kill protected species and to
hunt or kill wildlife inside protected areas. However, legal frameworks
vary between countries. For example, some countries have had consis-
tent legislation relating to wildlife protection in place for decades, while
the legal frameworks in others are still in their nascent stages. Some
countries have had endless repeals and amendments, whereas others
Table 1
Overview summary of Arabian wolf status, protection, and conservation effort in the eleven countries likely to be within its distribution. PA =protected area. For
further details, see Table S1.
Country Enforcement of protective
legislation
Conservation effort PA as
% land area
Arabian wolf status
(Density; trend;
persecution)
Egypt (Sinai Peninsula) Some inside PAs
a
Active network of NGOs across Egypt working towards hunting
reduction.
13.14 Rare
a
; unknown;
widespread
a
Iraq Negligible Signed Convention on Biological Diversity (2009) and produced
National Biodiversity Strategy and Action Plan (2015). Has active
network of conservation NGOs.
1.53 Unknown; unknown;
widespread
Israel / Palestinian
Territories (West Bank
and Gaza)
Israel: Strong inside and
outside PAs
Palestinian Territories:
Strong in some areas
Israel: Strong governmental and NGO support for wildlife
protection and conservation.
Palestinian Territories: Network of NGOs but many are inactive
due to cuts in overseas funding.
Israel: 24.49
Palestinian
Territories: 8.36
Israel: Common; stable;
negligible
Palestinian Territories:
Unknown; unknown;
unknown
Jordan Strong inside PAs, some
outside
Strong semi-governmental support for wildlife protection and
conservation, and a large network of NGOs.
3.09 Occurs in PAs; declining
a
;
widespread
Kuwait Strong inside PAs Well-established network of NGOs with an increase in prioritising
biodiversity conservation.
17.10 Extinct; NA; NA
Oman Strong inside and outside
PAs
a
, but poaching
common
Strong governmental and NGO support for wildlife protection and
conservation.
2.57 Occurs in PAs; declining
a
;
unknown
Qatar Some inside PAs An increase in governmental and NGO effort. 13.23 Extinct; NA; NA
Saudi Arabia Some inside PAs Several governmental and non-governmental pushes towards
wildlife protection and conservation, but difculty in moving from
policy to practice.
4.76 Common; declining;
widespread
Syria Negligible A small network of NGOs exists, but civil and political tensions
have caused conservation efforts to cease.
0.69 Unknown; unknown;
unknown
United Arab Emirates
(UAE)
Strong inside PAs, some
outside
Strong governmental and NGO support for wildlife protection and
conservation.
17.95 Extinct; NA; NA
Yemen Negligible Predator protection efforts increasing (e.g., focused on the Arabian
leopard), but civil and political tensions hinder sufcient
expansion.
0.77 Occurs in PAs; declining;
unknown
a
Considerable level of uncertainty due to either a lack of, or conicting, information.
G.T. Bonsen et al.
Biological Conservation 296 (2024) 110655
8
have had few regulations added to their long-standing laws (see Table S1
for a more detailed review of legislation and conservation effort).
Inconsistencies in legal frameworks create confusion within policies;
legislations that are present in some countries seem to exist solely on
paper, but not in reality. Adding to this confusion is the fact that many
governments continue to lack solid institutional structure attributed to
biodiversity conservation or environmental matters. Such muddled
institutionalisation often results in a lack of trained staff, funding, and
infrastructure. Most nations acknowledge such shortcomings in their
NBSAP's and aspire to improve (see Table S1 sources). However, even
with institutional and strategic improvements being made, shortfalls
remain in moving from policy to practice. For example, considerable
effort went into implementing Saudi Arabia's conservation strategies
and action plans. The relevant governmental authority engaged in dra-
matic transformation, prioritising the structured support for their pro-
tected area network and protected species lists. Yet, actions assigning
protective status, enforcing laws, and acting upon violations have been
noted as unsatisfactory (Barichievy et al., 2018). Although wolves
persist within Saudi Arabia's designated protected areas (Abuzinada,
2003; Seddon et al., 1997; Wronski and Macasero, 2008), it is unclear
how much actual protection they are afforded (Cunningham and
Wronski, 2010; Seddon, 2000). What is known is that Arabian wolves
are subjected to heavy persecution outside protected areas (Alou and
Amr, 2018; Cunningham et al., 2009; Cunningham and Wronski, 2010;
Wronski and Macasero, 2008; Zafar-ul Islam et al., 2019).
Protected areas remain a key legislative protection action for the
Arabian wolf. However, the status and meaning of these protected
spaces differ between countries. For example, while some countries have
a single managing agency, authoritative control of protected areas can
be variable within a country. Jordan's Royal Society for the Conserva-
tion of Nature actively enforces protection laws in areas under their
governance; however, they are not authorised to govern all the country's
protected areas (UNEP-WCMC, 2020). Another example of difference is
in enforcement. On Yemen's mainland, protected areas do not yet afford
strong protection for wildlife (Schlecht et al., 2014). Although areas
have been designated, they are not included in ofcial registries and lack
institutional governance (Environmental Protection Agency, 2017).
Along with the gulf countries, Yemen's mainstream conservation effort
tends to focus on marine and offshore biodiversity (e.g., Socotra Island).
Plans and proposals for extending the protected area network were
presented a decade ago (Schlecht et al., 2014); however, the country has
since been engulfed in a civil war and conservation priorities have taken
a step down.
To mitigate these differences, non-governmental conservation effort
appears to be increasing throughout the region, with domestic NGOs
now established in all countries. In countries where conservation is yet
to become a national priority, local NGOs ll the void to conduct much-
needed research and monitoring surveys. The Foundation for the Pro-
tection of the Arabian Leopard in Yemen is a non-prot dedicated to
Arabian leopard conservation. In Iraq, Nature Iraq has been working
with governmental ministries towards implementing conservation
strategies since 2003, liaising with the Iraqi government on policy
change for issues relating to wildlife protection and biodiversity
conservation.
Implementing protective legislation and designating protected areas
can only go so far towards the conservation of the Arabian wolf. Making
it illegal to kill them does not necessarily stop people from doing so (Eid
and Handal, 2018). It is more important to understand the needs and
struggles of the people that share the land with such wildlife, and
compassionately work with them towards peaceful coexistence. Reso-
lution of any conict is only possible once the situation has been
addressed from the perspectives of those most affected.
5.3. Society, culture, and human-wolf relationships
Humans and wolves have shared a vibrant coexistence across the
Middle East throughout history. Archaeological (Davis and Valla, 1978;
Dayan, 1994) and genetic (Freedman et al., 2014; Vonholdt et al., 2010)
evidence points to early domestication of wolves or a wolf-dog ancestor
in the Levant at least 12,000 years ago, suggesting that humans and
canids have been forming close relationships in the region for thousands
of years. With the advent of agriculture not long after (Zeder, 2011),
conicts between humans and wolves would undoubtedly have ensued.
From this point, dogs were introduced and selectively bred for practical
purposes such as livestock protection (Gehring et al., 2010) and hunting
(Guagnin et al., 2018), and which we have observed are still used today
by Bedouins for livestock protection.
Conict continues to dominate the narrative of coexistence between
wolves and people. Pastoralists frequently lose unsecured sheep and
goats to wolf depredation, often sparking retaliation (Cunningham et al.,
2009). On top of livestock-related persecution, age-old beliefs that
wolves endanger human lives (Seddon and Khoja, 2003), and that wolf
body parts can be used for therapeutic purposes (Alou and Eid, 2016)
persist. Moreover, widespread cultural fables depicting canids as ma-
levolent and impure have triggered large-scale eradication attempts
(Subasi, 2011). Legal and illegal hunting and persecution are common
where these beliefs prevail (Eid and Handal, 2018; Giangaspero and Al
Ghafri, 2015), and many threatened species, including Arabian wolves,
are largely conned to protected areas (Amr et al., 2004).
Conversely, Arabian wolves are accepted and even appreciated in
other regions. In Israel's Negev, particularly in crop farming landscapes
of the Arava Valley, wolves and people peacefully coexist. Farmers assist
wolves and other wildlife by intentionally providing water in this hyper-
arid landscape, where wolves are free to roam through agricultural
elds. A series of incidents in 2017, in which a few wolves came un-
comfortably close to people at a village adjoining a nature reserve,
generated outrage among the locals; but this was quickly resolved with
education programs around the importance and hazards of nature and
wildlife (Bonsen, 2018). Likewise, villagers and pastoralists coexist with
wolves in Yemen's Hawf district, the location of the country's only
mainland designated protected area (Schlecht et al., 2014), where
acceptance of wolves and other predators is reportedly higher than in
other parts of the Arabian Peninsula (Khorozyan et al., 2014).
Despite this, coexistence between people and Arabian wolves ap-
pears achievable but requires systematic, cross-jurisdictional, and co-
ordinated efforts to promote coexistence. Although livestock loss is not a
major concern in Israel's crop farming landscapes, farmers are not free
from economic costs associated with living with wildlife (Moran, 2003).
However, where conservation management and enforcement are com-
mon, they have accepted and learnt to adapt to the presence of wolves;
strategizing to minimise such costs and ultimately appreciating the
many benets of having wildlife on their doorsteps. With improvements
in cooperation and planning, coexistence is possible elsewhere.
6. Collaborative planning
The key challenges for the Arabian wolf are that a) they live in arid
environments where resources are scarce and conditions harsh; b) as
large predators, they rely on populations of wild herbivores; c) avail-
ability of livestock puts them into conict with pastoralists; d) while
laws protecting them are essential, they must also be enforced and
garner local community support; and e) their range crosses many po-
litical borders, encompassing disparate jurisdictions, each with its own
values, laws, and motivations. These challenges may in themselves be
inuenced by climate change and conict between people.
Considering this, we suggest that successful protection of Arabian
wolves must involve coordination and collaboration at both local and
regional scales. At local scales, there is good evidence that it is important
for conservation groups and authoritative bodies to form caring re-
lationships with local stakeholders to cooperatively plan solutions for
mitigating potential costs and hardships associated with coexistence
(Jiren et al., 2021; Kansky and Maassarani, 2022). Rather than forcibly
G.T. Bonsen et al.
Biological Conservation 296 (2024) 110655
9
implementing policy and legislation, in which violators (often unwary)
are automatically penalised, better outcomes can arise when authorities
take time to codevelop solutions to underlying issues. A good example
comes from recent collaborations with Bedouin pastoralists, who are
often marginalised when promoting conservation goals. It has been re-
ported that authorities have failed to act on environmental damage re-
ported by Bedouins, penalising them more than other societal groups for
environmental violations (Gilbert, 2013). In contrast, conservation
groups in Jordan recognised early aws when failing to consider pas-
toralists in the establishment of a nature reserve in Jordan, and hence
began involving Bedouins as primary stakeholders and collaborators in
developments of further conservation projects (e.g., Wadi Rum Nature
Reserve), greatly improving their success (Chatty, 2002). Co-
development of action plans with local people, including First Nations
people, is now being recognised as an important breakthrough in local
conservation efforts around the world (Campion et al., 2023).
The cultivation of efforts and achievements from conservation
groups and authoritative bodies throughout the Middle East is certainly
promising, but these are largely unilateral and conned to individual
jurisdictions. For example, the United Arab Emirates is currently one of
the most committed Arab countries to wildlife protection: local NGOs
have spearheaded international reintroduction projects and residents
are encouraged to report any environmental or wildlife-related viola-
tions (Salama, 2018). Likewise, Israel is generally conservation-minded,
with clear goals and objectives, as well as strong public adherence to
wildlife protection (Nemtzov and King, 2001). However, collaborative
regional-scale efforts across international borders, like those in Europe,
must improve if Arabian wolf populations are to recover. All European
countries have subscribed to the Bern Convention (Convention on the
Conservation of European Wildlife and Natural History, 1979), by which
all wolf populations and habitats are fully protected (Boitani, 2003).
During the 1990s, the Large Carnivore Initiative for Europe (within the
IUCN's Species Survival Commission) was established to improve coex-
istence between people and wolves throughout the region, while pri-
oritising heightened consideration of both parties (Linnell et al., 2008).
Although transboundary conservation initiatives have also been carried
out within the Middle East, these have mostly focussed on local-scale
projects across individual borders (Knight et al., 2011).
International research collaborations have increased over the last
decade or so; however, regional-scale and multi-jurisdictional policy
implementations are yet to be pushed. For example, the Middle Eastern
Biodiversity Network was established in 2006 between universities and
research institutes from Iran, Jordan, Lebanon, and Yemen in a collab-
orative effort to conserve biodiversity across the Middle East (Krupp
et al., 2009). More recently, Compassionate Conservation Middle East
(CCME) was established as a research group collaboration between
various institutes and NGOs in Israel and the Palestinian Territories.
CCME has developed wolf research and conservation projects across
Israel and Palestine with the primary objectives of improving both
human-wolf coexistence and regional collaborations – based on the
hopeful politically-focused adage ‘nature knows no borders’ (Roulin
et al., 2017). Unfortunately, transboundary cooperation can be difcult
to achieve in regions as politically tense as the Middle East.
7. Concluding remarks
The Arabian wolf is likely to be an important trophic regulator of
desert ecosystems of the southern Levant and Arabian Peninsula. To
ascertain just how important it is, there are signicant knowledge gaps
that need to be lled. Although populations are sparse across most of its
wide range, the Arabian wolf remains the most widespread apex pred-
ator inhabiting the deserts of the Middle East. Yet, little is known about
its potential inuence on these arid ecosystems. Considering its close
association with human infrastructure and reliance on anthropogenic
food resources in Israel, there is some conjecture as to how important a
role the Arabian wolf plays in shaping trophic cascades like its larger,
temperate counterparts.
That it might do so, however, is only within the context of its
acceptance by people. Demarcations of tolerance or persecution can
override the trophic position of these desert-dwelling wolves. Conser-
vation conicts are often interrelated with intra-human conicts, and
the Middle East is one of the most complex geopolitical landscapes in the
world. Any solutions to conserve the Arabian wolf must be cognisant of
this complexity and strive to ameliorate the internal conicts shaped by
the distinct attitudes and values that are ingrained among the peoples
that share this extensive landscape.
Funding
This review was put together as part of the International Wolf Project
funded by an Australian Research Council Discovery Grant
(DP180100272), the United Kingdom Wolf Conservation Trust, and
Voiceless. GTB was supported by a University of Technology Sydney
Doctoral Scholarship.
CRediT authorship contribution statement
Gavin T. Bonsen: Conceptualization, Formal analysis, Funding
acquisition, Investigation, Methodology, Visualization, Writing – orig-
inal draft. Arian D. Wallach: Conceptualization, Funding acquisition,
Investigation, Methodology, Visualization, Writing – review & editing.
Dror Ben-Ami: Conceptualization, Funding acquisition, Visualization,
Writing – review & editing. Oded Keynan: Conceptualization, Funding
acquisition, Visualization, Writing – review & editing. Anton Khalilieh:
Visualization, Writing – review & editing. Yara Dahdal: Visualization,
Writing – review & editing. Daniel Ramp: Conceptualization, Formal
analysis, Funding acquisition, Methodology, Visualization, Writing –
original draft.
Declaration of competing interest
The authors declare that they have no known competing nancial
interests or personal relationships that could have appeared to inuence
the work reported in this paper.
Data availability
No data was used for the research described in the article.
Acknowledgements
We are grateful to the many researchers, conservation workers and
advocates, local farmers, and tour guides who helped us construct this
review. In particular, we thank Itamar Yairi, Ron Tzur, Gilad Guy,
Shlomo Preiss-Bloom, Inbar Schnitzer, Nikola Dragi´
c, Djordje Markovi´
c,
Dorit and Yonatan Narisna, Yael and Gilad Alon, Adi and Yinon Offaim,
and Ofer Kubi for sharing their intimate knowledge of the wolves of the
Arava Valley and Golan Heights; Zuhair Amr, Eli Geffen, Adi Barocas,
Uri Shanas, Yael Olek, Yohay Carmel, Oded Nezer, Michael Kam, Lee
Koren, and Shani Babel for providing invaluable information regarding
desert ecology in the Middle East; the Royal Society for the Conservation
of Nature in Jordan, Israel Nature and Parks Authority, and Society for
the Protection of Nature in Israel for hosting and assisting GTB in the
Middle East and for their insights into wolf conservation across the re-
gion. We thank Dan Malkinson, Tamar Dayan, Varun Goswami, and an
anonymous reviewer for providing helpful feedback on earlier versions
of this manuscript. Photos of Arabian wolves were provided by Nikola
Dragi´
c, and the Indian wolf in Fig. 1 was modied from an image pro-
vided by David Kwan under a Creative Commons licence (CC BY-NC 2.0
DEED).
G.T. Bonsen et al.
Biological Conservation 296 (2024) 110655
10
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.biocon.2024.110655.
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