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\begin{document}

\title{A review of current and prospective findings concerned with the
`Landscape of Fear' concept and its implications}



\author[1]{Samuel Ashby}%
\author[1]{Kathan Bandyopadhyay}%
\author[1]{Georgia Warren}%
\affil[1]{University of Reading}%


\vspace{-1em}



  
  \date{June 16, 2024}


\begingroup
\let\center\flushleft
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\selectlanguage{english}
\begin{abstract}
The capacity and extent to which prey species forage is often dependent
on the temporal and/or spatial distribution of predators. Predation risk
within a given habitat may differ according to the structure of the
landscape and ecological community. Predators may frequent selected
habitat patches and it is these areas prey are expected to avoid. Aside
from the direct removal of prey individuals through predation, the
density of prey populations may be altered as a result of a perceived
predator presence and the energetically expensive responses initiated. A
predator presence may be perceived upon the detection of sensory
environmental cues, including a predator's pheromones. The Landscape of
Fear (LOF) concept proposes the exposure to a real or perceived
predation threat may disrupt prey distribution and activity. Such an
environment may be referred to as a `landscape of fear', though the
interspecies complexes and abiotic factors affecting a predator-prey
relationship should not be omitted when quantifying the effects of
predation. Here, we summarise the initial and more recent publications
addressing the LOF theory, identifying known aspects and potential for
future research.%
\end{abstract}%



\sloppy




\section*{Introduction}

{\label{introduction}}

Predation is a strong evolutionary force acting as a selective pressure
on various traits within a prey species including, but not limited to,
size, morphology, coloration, development of sexual maturity, and
patterns of behaviour (Werner \& Anholt 1993; Maynard-Smith \& Harper
2003; Scott-Phillips 2008). Ultimately, the distribution, abundance,
community composition, size, diversity, breeding success and ecosystem
functioning of one or more species are impacted (Hall \emph{et al.}
1976; Berger \emph{et al.} 2001; Kneitel \& Chase 2004; Beauchamp
\emph{et al.} 2007; Hawlena \& Schmitz 2010; Letnic \emph{et al.} 2011).
Interactions between predators and prey can over time make communities
vulnerable to biotic and/or abiotic perturbations (Mooney \emph{et al.}
2010), including the invasion of non-native species as a consequence of
human activity (Snyder \emph{et al.} 2004). At the most general level,
predation results in the reduced survival of prey (consumptive effects),
but the (real or perceived) risk of predation can also affect prey
animals in a variety of ways (Zanette\emph{et al.} 2011), including
stinted growth, poor reproductive output and reduced foraging efficiency
(Sih 1980; Lima \& Dill 1990; Peckarsky\emph{et al.} 1993; Abrams
\emph{et al.} 1996; Peacor \& Werner 1997). Individuals most capable of
evading predators have an increased potential for producing offspring,
and therefore should achieve higher reproductive success than less
capable conspecifics; female cheetahs (\emph{Acinonyx jubatus} ) with
higher fecundity avoided habitats with indications of elevated spotted
hyaena (\emph{Crocuta crocuta} ) or lion (\emph{Panthera leo} )
densities more than females with lower fecundity (Durant 2000).

{\label{predation-is-a-strong-evolutionary-force-acting-as-a-selective-pressure-on-various-traits-within-a-prey-species-including-but-not-limited-to-size-morphology-coloration-development-of-sexual-maturity-and-patterns-of-behaviour-werner-anholt-1993-maynard-smith-harper-2003-scott-phillips-2008.-ultimately-the-distribution-abundance-community-composition-size-diversity-breeding-success-and-ecosystem-functioning-of-one-or-more-species-are-impacted-hall-et-al.-1976-berger-et-al.-2001-kneitel-chase-2004-beauchamp-et-al.-2007-hawlena-schmitz-2010-letnic-et-al.-2011.-interactions-between-predators-and-prey-can-over-time-make-communities-vulnerable-to-biotic-andor-abiotic-perturbations-mooney-et-al.-2010-including-the-invasion-of-non-native-species-as-a-consequence-of-human-activity-snyder-et-al.-2004.-at-the-most-general-level-predation-results-in-the-reduced-survival-of-prey-consumptive-effects-but-the-real-or-perceived-risk-of-predation-can-also-affect-prey-animals-in-a-variety-of-ways-zanette-et-al.-2011-including-stinted-growth-poor-reproductive-output-and-reduced-foraging-efficiency-sih-1980-lima-dill-1990-peckarsky-et-al.-1993-abrams-et-al.-1996-peacor-werner-1997.-individuals-most-capable-of-evading-predators-have-an-increased-potential-for-producing-offspring-and-therefore-should-achieve-higher-reproductive-success-than-less-capable-conspecifics-female-cheetahs-acinonyx-jubatus-with-higher-fecundity-avoided-habitats-with-indications-of-elevated-spotted-hyaena-crocuta-crocuta-or-lion-panthera-leo-densities-more-than-females-with-lower-fecundity-durant-2000.}}

In response to the severity of the risks posed by predators, prey
species have evolved a wide range of anatomical, physiological,
neurobiological and behavioural mechanisms to increase their chances of
survival. Adaptations related to morphology and physiology are
particularly energetically expensive, and thus prey may display
different responses to a threat depending on how severe they perceive it
to be, known as the threat-sensitive predator avoidance hypothesis
(Chivers \emph{et al.} 2001; Moncl\selectlanguage{ngerman}ús \emph{et al.} 2009).

Some adaptations reduce the chance of initial detection by a predator
including: camouflage (Cott 1940; Ruxton \emph{et al.} 2004; Stevens \&
Merilaita 2008) which includes crypsis where an individual resembles its
surroundings and masquerading, when an individual resembles a single
inanimate object (Ruxton \emph{et al.} 2004; Quicke 2017); and Batesian
mimicry in non-toxic species that have evolved to imitate the appearance
of toxic species (Lindström \emph{et al.} 1997).

Others do not decrease initial detection, and in some cases may increase
the chance of detection, but instead reduce the chance of the predator
successfully carrying out the job and serve as a means of protection
from a predator after detection including: defensive structures such as
spines, shells or scales (Ruxton \emph{et al.} 2004; Yadun \& Halpern
2008); autotomy (loss of a body part) and regeneration (Maginnis 2006);
warning colouration and associated toxins (Fisher 1930; Harvey \emph{et
al.} 1982), both of which are also associated with mimetic species
(Huheey 1961; Brower \& Brower 1963; Brower \emph{et al.} 1963;
Pough\emph{et al.} 1973); venoms (Holding \emph{et al.} 2016);
construction of appropriate refugia e.g. bolt holes (Baugh \& Deacon
1988); and plastic growth and development (Edmunds 1974; Lima \& Dill
1990; Tollrian \& Harvell 1999; Sokolowska \emph{et al.} 2000). For
example, the wide variation in shell structure in marine invertebrates
is believed to be a response against shell-destroying predators (Vermeij
1997). Similarly, the freshwater three-spine stickleback
(\emph{Gasterosteus aculeatus} ) exhibits a complex of distinct
morphological forms, varying in the degree of spinal and lateral plates
as a consequence of predation risk (Colosimo \emph{et al.} 2005).

Group aggregations are another example of an adaption in response to
predator threat, leading to benefits including increased vigilance
(Treves \emph{et al.} 2001), selfish-herd effects (King \emph{et
al.}2012), and active defence e.g. musk ox \emph{Ovibos moschatus}
(Dixon 1998). Furthermore, such group behaviours have given rise to the
development of group warning/alarm calls in some species (Smith 1986),
and sometimes leads to sentinel behaviour in which one individual scans
for predators whilst the other group members forage, observed in vervet
monkeys (\emph{Chlorocebus pygerythrus} ) (Horrocks \& Hunte 1986). In
Gunnison's prairie dogs (\emph{Cynomys gunnisoni)} this has led to
species-specific warning calls, allowing the group to react in
accordance to the threat (Slobodchikoff \emph{et al.} 1991; Kiriazis \&
Slobodchikoff 2006; Slobodchikoff \emph{et al.} 2009).

In addition to the mechanisms outlined above, prey animals use basic
sensory modalities (visual, aural, olfactory) to detect the presence of
predators in the same way that predators use both direct and indirect
cues to identify and locate the position of the prey (Dwernychuk \& Boag
1972; Sugden \& Beyersbergen 1986; Nams 1997; Santisteban \emph{et
al.}2002; Carthey \emph{et al.} 2011). In these contexts, visual, aural
and olfactory cues each offer advantages and disadvantages to predator
and prey in terms of being detected and/or avoiding detection. Vision is
a directional cue and relies upon direct ``line of sight'' between two
individuals, so can be used by prey to indicate the exact position and
identity of the predator, and vice versa (Hemmi 2005). Predators may
reduce the risk of detection while residing amongst vegetation, for
instance. Often, prey may only observe their immediate environment at
the expense of feeding, resulting in a trade-off between foraging and
vigilance. Though social species may deploy sentinels to detect
approaching predators, such individuals in turn experience a foraging
cost by foregoing feeding to protect other group members (Doolan \&
Macdonald 1996; Whittingham \emph{et al.} 2004). The slender-tailed
meerkat (\emph{Suricata suricatta} ), vervet monkey (\emph{Chlorocebus
pygerythrus} ), and black-tailed prairie dog (\emph{Cynomys
ludovicianus} ) exhibit such complexes within their respective social
groups (Magle \emph{et al.} 2005). Furthermore, visual cues are of
limited use for nocturnal species and those living in dense vegetation.

Auditory cues are also directional, although generally to a lesser
extent than visual cues due to sound resonating in a non-linear fashion
from a point source. Predators, therefore, may inadvertently reveal
their proximity when stalking prey, but such signals may not disclose
exact locations and are less valuable to prey than direct sight.
Furthermore, auditory signals are attenuated and reflected by vegetation
and other surfaces, reducing the distance they travel and obscuring
their origin (Goerlitz \emph{et al.} 2008). Similar effects may arise in
poor weather conditions, where sounds are refracted in strong winds
(Clinchy \emph{et al.} 2013). Consequently, predators that stalk prey
must approach conspicuously and, preferably, upwind. Sound, however, is
not likely a significant concern for ambush (sit-and-wait) predators
(Boonstra \emph{et al.} 2013).

\section*{\texorpdfstring{\emph{Use of chemicals in
mammals}}{Use of chemicals in mammals}}

{\label{use-of-chemicals-in-mammals}}

For mammalian predators, chemical cues are particularly problematic in
terms of avoiding detection (Stoddart 1980). Olfactory cues are
especially important to mammalian predators as they are frequently used
to communicate information about sexual condition, group membership,
social status and territory ownership (Ralls 1971; Bel\emph{et al.}
1999; King \& Gurnell 2007; Gelperin 2008; Sankar \& Archunan 2008).
Scent cues are even utilised by marine mammals despite them often being
categorised as anosmic or microsmatic (no/reduced olfactory processing
centres) (Lowell \& Flanigan 1980; Tyack \& Sayigh 1997), such as female
Australian sea lions using scent cues in offspring recognition
(\emph{Neophoca cinerea} ) (Pitcher \emph{et al} . 2010).

{\label{for-mammalian-predators-chemical-cues-are-particularly-problematic-in-terms-of-avoiding-detection-stoddart-1980.-olfactory-cues-are-especially-important-to-mammalian-predators-as-they-are-frequently-used-to-communicate-information-about-sexual-condition-group-membership-social-status-and-territory-ownership-ralls-1971-bel-et-al.1999-king-gurnell-2007-gelperin-2008-sankar-archunan-2008.-scent-cues-are-even-utilised-by-marine-mammals-despite-them-often-being-categorised-as-anosmic-or-microsmatic-noreduced-olfactory-processing-centres-lowell-flanigan-1980-tyack-sayigh-1997-such-as-female-australian-sea-lions-using-scent-cues-in-offspring-recognition-neophoca-cinerea-pitcher-et-al.-2010.}}

Chemoreception also plays a role in optimising the use of food caches,
the defence of shelter and resources, and for active defence (Henry
1977; Medill \emph{et al.} 2011; Piñeiro \& Barja 2015, Appleton \&
Palmer 1988). In these contexts, scent-marks are used actively. In the
case of territory ownership, scent marks confer one significant
advantage; they enable territorial boundaries to be delineated without
the need for direct contact between conspecifics, therefore reducing the
risk of injury. Scent cues are, however, also left inadvertently by
individuals in the form of dander (usually comprised of skin cells or
fur) which is dispersed while traveling (Ferrero \emph{et al.} 2011).
These ``accidental'' scents, as well as those associated with urine,
faeces and the excretions from scent glands, contain a complex mixture
of compounds that vary in their molecular weight, such that their
composition varies over time as different compounds aerosolise and
disappear or reduce in concentration (Seamans \emph{et al.} 2002; Burger
2005). Consequently, conspecifics use the composition of these chemical
cues to indicate the length of time since their deposition and hence the
likeliness of the presence or absence of an individual (Wyatt 2003).

While a conspecific may incorporate chemical indications into its
assessment of social status and mating opportunity, prey species may use
chemical signals to assess the presence of a predator and likelihood of
predation. In predator-prey relationships, chemosensory cues are often
invaluable and the process of communication via chemicals has been
widely studied (Amo \emph{et al.} 2007). Olfactory cues can provide
information on both direct and prospective predatory threats and this
can be particularly important during times of limited visibility (Kats
\& Dill 1998). However, since aerosolised molecules are distributed by
prevailing wind currents, which themselves are susceptible to variation
in landscape topography and weather conditions, olfactory cues are also
less directional than visual cues (Burghardt 1966; Halpern \& Frumin
1979; Redmond \emph{et al.} 1982; Roth \& Hobson 2000; Conover 2007;
Fogarty \emph{et al.} 2018). Olfactory cues pose additional problems for
predators (especially mammalian) where intra-specific chemical
communication is frequent, and scent is inherently difficult to mask
from prey (Venuleo \emph{et al.} 2017).

The detection of kairomones, specific to mammalian carnivores, often
initiates defensive behaviours, reduced foraging, and increased refugia
in mammalian prey (Mathis 2003; Foam \emph{et al.} 2005;
Preisser\emph{et al} . 2005; \selectlanguage{polish}Š\selectlanguage{english}mejkal \emph{et al.} 2018). Similarly,
rodents have been observed to increase evasive behaviours in response to
the detection of 2-phenylethylamine which, though found in all mammalian
urine, exists at significantly higher concentrations in carnivore
excretions (Ferrero \emph{et al.} 2011).

\emph{The `Landscape of Fear' concept}

Mammalian predators often do not to use their home range or territory
uniformly, and thus areas of high activity (core areas) would be
expected to be associated with a high density of kairomones and other
predatory chemical cues (Atkins \emph{et al.} 2019). Prey individuals,
therefore, should avoid such areas and may be able to assess relative
predation risks in different areas (Atkins \emph{et al.} 2019).
Conversely, the removal or reduction of predation risk may enable the
dispersal of prey populations; declines in Persian leopards
(\emph{Panthera pardus} ) and African hunting dogs (\emph{Lycaon pictus}
) allowed the recovery and distribution of Cape bushbuck
(\emph{Tragelaphus sylvaticus} ) in Mozambique (Atkins \emph{et
al.}2019). Relationships such as these, however, have formed across
multiple generations of predator and prey species and, as such,
alterations may not result in observable ecological changes immediately;
a 50 to 130 year absence of predators may result in the inability of
prey to identify recolonizing predators as a threat (Berger \emph{et
al.} 2001). Such phenomena have given rise to the concept of a
``landscape of fear'' (LOF), where prey may reduce their temporal and
spatial exposure to core predator areas in relation to perceived
predation risk, although a variety of other biological and evolutionary
factors besides fear also influence habitat use (Laundr\selectlanguage{ngerman}é \emph{et al.}
2001; Laundré \emph{et al.}2010; Bleicher 2017).

As a prerequisite for prey to actively avoid predation threats, it must
be possible to identify and associate predatory cues with predation risk
(Griffith 1920; Schaller 1974; Mech 1970: Table 1). In some cases, the
ability to identify scents specific to a predator species may be
advantageous; foraging beavers (\emph{Castor fiber} ) show significantly
fewer defensive behaviours in response to the odours of dogs
(\emph{Canis familiaris} ) compared to those of wolves (\emph{Canis
lupus} ) (Rosell \& Czech 2000); grey kangaroos (\emph{Marcopus
fuliginosis} ) can discriminate the urine of coyotes (\emph{Canis
latrans} ) from dingoes (\emph{Canis lupus dingo} ) (Parsons \emph{et
al.}2007); mule deer (\emph{Odocoileus hemionus} ) respond to the faecal
remnants of coyotes and mountain lion (\emph{Felis concolor} ), but not
those of African lions (\emph{Panthera leo} ), leopards, or tigers
(\emph{Panthera tigris} ) (Mulller-Schwarze 1972). The type of kairomone
left by a predator is often species-specific; weasel (\emph{Mustela
nivalis} ) tracks are rich in sulphur compounds, while those of beech
martens (\emph{Martes foina} ) contain no sulphur-rich components
(Schildknecht \& Birkner 1983; Apfelbach \emph{et al.} 2015). The
ability to detect kairomones also differs between prey species; the
abundance of trace amine-associated receptors (TAARs), a group of
olfactory receptors used to detect kairomone presence, differs between
prey species (Borowsky \emph{et al.} 2001; Ferrero \emph{et al.} 2011).
Rodents may possess up to seventeen TAARs, while primates only six
(Borowsky \emph{et al.} 2001; Ferrero \emph{et al.} 2011). The ability
to differentiate between predatory threats and benign, albeit from
closely related predators, chemical cues may reduce the energetic
expenditure spent on vigilance and evasion.

Furthermore, the extent of predation risk likely varies, and prey
species should possess the ability to distinguish between the least
threatening and more ominous indicators. These abilities are often
demonstrated in the occurrence of ``giving up distributions'' (GUDs),
with the emphasis on whether prey animals sacrifice foraging behaviour
in relation to the perceived risk of predation (Brown 1988;
Brown\emph{et al.} 1994; Laundré \emph{et al.} 2001). For instance,
moose (\emph{Alces alces} ) quit more than half of experimental areas
following repeated exposure to urine from wolf and grizzly bears
(\emph{Ursus arctos} ) (Pyare \& Berger 2003). More specifically, a
forager should terminate feeding in a food patch when the value of its
harvest rate (H) no longer exceeds the sum of the energetic cost of
foraging (C), predation risk (P) and missed opportunity cost (MOC):
H=C+P+MOC (Brown 1988), and has been shown to apply in a range of taxa
under varying ecological conditions (Stokes \emph{et al.} 2004; Merwe \&
Brown 2008; Shrader \emph{et al.} 2008; Valeix \emph{et al.} 2009;
Iribarren \& Kotler 2012; Clinchy \emph{et al.} 2013; Gallagher \emph{et
al.} 2017).\selectlanguage{english}
\begin{longtable}[]{@{}llllll@{}}
\toprule
\begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
\textbf{Predator species}\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
\textbf{Prey species}\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
\textbf{Sensory} \textbf{modality}\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
\textbf{Behaviour response}\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
\textbf{Result}\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
\textbf{Reference(s)}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Black bear (\emph{Ursus americanus})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Elk (\emph{Cervus canadensis})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Visual\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Habitat use\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
black bear predation as well as mountain lion sites (0.636, 95\%
CI=0.531--0.741) had higher canopy over the capture sites (0.186, 95\%
CI=0.152-- 0.220).\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Quintana 2016\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Dogs (\emph{Canis lupus familiaris})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Raccoon (\emph{Procyon lotor})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Sound\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Foraging on beaches; vigilance behaviour\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Raccoons reduced activity on beaches by 50\%; increased vigilance by
66\%\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Brown (2018)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Wolf (\emph{Canis lupus})\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Elk and Bison (\emph{Bison bison)}\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Visual\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Foraging on habitat; vigilance behaviour\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Male elk \& bison showed no response, low levels of vigilance (7\%) but
female elk \& bison showed significantly higher vigilance (47.5\%)\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Laundr\selectlanguage{ngerman}é \emph{et al.} (2001)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Golden eagle (\emph{Aquila chrysaetos})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Greater sage grouse (\emph{Centrocercus urophasianus})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Visual\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Lekking behaviour\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Across all 26 lek years, vigilance behaviour in males and females
decreased as male attendance increased\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Boyko \emph{et al.} (2004)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Saharan horned viper (\emph{Cerastes cerastes}) and Sidewinder
rattlesnake (\emph{Crotalus cerastes})\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Allenby's gerbil (\emph{Gerbillus andersoni})\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Moonlight\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Foraging on resource patches\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
In response to both snakes, giving-up densities of the gerbils were
higher in the bush than open microhabitat. In response to moonlight,
GUDs were higher on full than on the new moon\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Bleicher \emph{et al.} (2016)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Coyote (\emph{Canis latrans})\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Black-tailed jackrabbit (\emph{Lepus californicus}) and Desert
cottontail rabbit (\emph{Sylvilagus audubonii})\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Visual\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Foraging in habitat; vigilance behaviour\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Jackrabbit and cottontails not only just balance food resources but
reciprocally alternate levels of predation risk and escaping success in
decision making\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Razo \emph{et al.} (2012)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Leopard (\emph{Panthera pardus}), wolf, striped hyena (\emph{Hyaena
hyaena}) and humans\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Nubian ibex (\emph{Capra nubiana})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Visual\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Foraging in habitat; vigilance behaviour\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
GUD was lowest at 40\% vegetation cover, indicating that thick
vegetation might obstruct vigilance or escape opportunities.\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Iribarren \& Kotler (2012)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Large raptors, black-backed jackal (\emph{Canis mesomelas}), caracal
(\emph{Caracal caracal}) and Leopard\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Cape ground squirrel (\emph{Xerus inauris})\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Visual\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Foraging in habitat; vigilance behaviour\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Among the colonies, only 3- 22\% colonies of the landscape resulted in
low foraging costs (\textless{}2,500 J/min)\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Merwe \& Brown (2008)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Humans\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Samango monkey (\emph{Cercopithecus albogularis})\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Visual\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Foraging in habitat; vigilance behaviour\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
The GUD were greatest at ground level (0.1 m) relative to the three tree
canopy levels (2.5,5 and 7 m)\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Nowak \emph{et al.} (2014)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Puma (\emph{Puma concolor})\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Vicu\selectlanguage{ngerman}ña (\emph{Vicugna vicugna})\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Movement\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Foraging on habitat; vigilance behaviour\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Puma moved less during the day (176.2 \selectlanguage{ngerman}± 3.8 SE m/h) than at dusk/dawn
(290.5 ± 5.9)\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Smith \emph{et al.} (2019)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Barn owl (\emph{Tyto alba}), Red fox (\emph{Vulpes vulpes}), Sidewinder
rattlesnake (\emph{Crotalus cerastes})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Negev desert gerbils (\emph{Gerbillus pyramidum} and \emph{G. andersoni
allenbyi})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Direct sight of predator\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Vigilance behaviour\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Higher GUD when sharing enclosure with caged predator compared to an
enclosure without a predator\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Kotler \emph{et al.} 2016\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Lion (\emph{Panthera leo}), Hyaena (\emph{Crocuta crocuta})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Juvenile cheetah (\emph{Acinonyx jubatus})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Vocalization recordings\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Habitat use\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Reduced prey occurrence in predator-dense areas\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Durant 2000\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Coyote\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
White-tailed deer (\emph{Odocoileus virginianus})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Real predator\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Daily activity levels\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Bucks and does shared a crepuscular lifestyle with coyotes, but adults
with offspring adapted activity levels and were most active at
midday.\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Higdon \emph{et al.} 2019\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Mountain lion (\emph{Puma concolor})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Mule deer (\emph{Odocoileus hemionus})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Predator faeces (chemical)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Vigilance behaviour (giving up food)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Altered habitat usage; higher vigilance when on outskirts of
forest\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Altendorf \emph{et al.} 2001\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Raccoon, Corvids (Family Corvidae), Owl (Family Strigidae), Hawk (Family
Accipitridae)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Song sparrow (\emph{Melospiza melodia})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Vocalization recordings\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Reproduction\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
40\% reduction in offspring and increased offspring death\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Zanette \emph{et al.} 2011\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Red fox\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Wood mouse (\emph{Apodemus sylvaticus})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Predator faeces (Chemical) \& Moonlight\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Use of live traps as refuge\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
To analyse the predation risk 67.5\% of the bottles (N=51) showed bite
marks, treated with fox faeces lower than in the absence of predator
cues (50.0\%, N = 27). Interaction of the food access and moonlight
showed a less frequent result for the open bottles during new moon
nights (27.8\%, N = 20) than during waxing/waning crescent moon nights
(58.8\%, N = 10).\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Hernandez \emph{et al.} 2019\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Red fox\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Wood mouse, field vole (\emph{Microtus agrestis}), bank vole
(\emph{Clethrionomys glareolus}), common shrew (\emph{Sorex
araneus})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Predator faeces and urine (chemical)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Use of traps treated with rodent odour as refuge\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Strong avoidance of areas treated with predator scents, especially in
wood mice and bank voles and least in field voles.\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Dickman \& Doncaster 1984\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Red fox\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Northern pocket gopher (\emph{Thomomys talpoides})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Predator faeces\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Preference of treated or control enclosures\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Significant; \textasciitilde{}66\% of gophers opted for the control
enclosures.\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Sullivan \emph{et al.} 1988a\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Dog\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Crested porcupine (\emph{Hystrix cristata})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Real predator\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Feeding behaviour\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
In areas where hunting dogs were used, porcupines altered home range and
opted for more accessible food which required less digging and distance
from refuge.\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Mori 2017\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Harris hawk (\emph{Parabuteo unicinctus})\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Egyptian geese (\emph{Alopochen aegyptiaca})\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Real predator\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Goose vigilance behaviour and abundance\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Goose vigilance increased by 76\%. Decreases in abundance (73\%) were
considerably larger than the number killed, indicating declines due to
mere presence of predator\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Atkins \emph{et al.} 2017\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Multiple raptor species\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
House mice (\emph{Mus domesticus})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Real predator\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Feeding in open habitat\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
High predator counts coincided with increased mouse feeding in
vegetation cover. This cover offered little nutrition and indicates a
trade-off of food quality versus predation risk.\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Y\selectlanguage{ngerman}ïonen \emph{et al.} 2002\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Coyote\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
White-tailed deer, squirrel (\emph{Sciurus spp.}), Eastern cottontail
rabbit (\emph{Sylvilagus floridanus})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Real predator\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Habitat preference\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Presence of coyotes caused deer to seek dense forest for refuge, while
rabbits and squirrels used urban areas as refuge.\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Jones \emph{et al.} 2016\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Wolf\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Elk\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Real predator\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Habitat preference\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Presence of wolves caused elk to shelter in wooded areas.\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Creel \emph{et al.} (2005).\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Wolf\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Elk\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Real predator\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Habitat preference\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Presence of wolves correlated with elk changing their habitat use from
open meadow to forest edge despite this habitat providing food of lower
quality. This habitat use was not observed in wolf-free areas.\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Hern\selectlanguage{ngerman}ández \& Laundré (2005)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Lion\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Plains zebra (\emph{Equus burchelli})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Real predator\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Vigilance, habitat preference\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Negative correlation between lion and zebra presence on a particular
patch on the same day. Zebra were observed using woodland at night as
lions increased the use of woodland during the day. Zebra exhibited
faster and sharper movements at night when lion activity is
elevated.\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Fischhoff \emph{et al}. (2007)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Leopard\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Vervet monkey (\emph{Chlorocebus aethiops})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Auditory (alarm call recordings)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Habitat use\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
LOF from leopards was the strongest driver of their use of the habitat
vervet monkey. Produced specific alarm calls to the threat from
leopards.\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Coleman \& Hill (2014)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Wild dog (\emph{Lycaon pictus})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Greater kudu (\emph{Tragelaphus strepsiceros}), sable antelope
(\emph{Hippotragus niger}), warthogs (\emph{Phacochoerus
africanus})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Real predator\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Giving-up densities (amount of food left in the patch once foraging has
ended), vigilance\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
In experimental areas Kudu GUDs and vigilance increased significantly,
whereas sable antelope and warthogs stopped feeding completely\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Makin \emph{et al.} (2017)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Brown bear\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Reindeer (\emph{Rangifer tarandus})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Real predator\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Access to growing vegetation (following the green-up).\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
When bear density was higher, reindeer deviated more from the path
towards lower quality food. In some cases this meant they missed the
green-wave peak. Increased vigilance (faster movements) also exhibited
when in closer proximity to high bear densities.\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Rivrud \emph{et al.} (2018)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Cougar (\emph{Puma concolor})\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Mule deer\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Real predator\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Access to growing vegetation (following the green-up)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Access to green-up vegetation by mule deer reduced by presence of
cougars, particularly in Spring\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Lowrey \emph{et al}. (2019)\strut
\end{minipage}\tabularnewline
\bottomrule
\end{longtable}

Under this LOF model, in addition to the pressure exerted by predators
in terms of mortality rate, predators will also influence the
distribution of prey species within the landscape. Consequently,
fear-mediated responses to predation risk can be mapped both as a
``landscape of fear'', which emphasizes areas prey should avoid, to
``landscapes of opportunity'' (Willems \& Hill 2009), which emphasizes
areas of low predation risk and increased foraging opportunities (Figure
1); lagomorphs, including rabbits (\emph{Oryctolagus cuniculus} ) and
jackrabbits (\emph{Lepus californicus} ), adopt a crepuscular lifestyle
to avoid the peak diurnal activity levels of coyotes (\emph{Canis
latrans} ) (Razo \emph{et al.} 2009). Such patterns will vary spatially
and temporally in relation to factors such as the diversity of the
predator community, conspecific density, predator activity and natural
environmental fluctuations (Bleicher \emph{et al.} 2017). In addition,
they will be susceptible to the influence of human activities such as
habitat destruction and alteration; the lethal control of predators; the
introduction of non-native species, including competitors and predators;
conservation activities; and climate change (Brown \emph{et al.} 1988;
Brown \emph{et al.} 2004; Bleicher \emph{et al.} 2016; Razo \emph{et
al.} 2012, Laundr\selectlanguage{ngerman}é \emph{et al.} 2001; Merwe \& Brown 2008).~\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image1/image1}

\end{center}
\end{figure}

\emph{Energetics related with LOF model}

Recent publications proposed the incorporation of energetics to
understand the movement in space with the conventional model of LOF
(Bleicher \emph{et al.} 2017). Previous studies quantified the energetic
expenditure and foraging trade-offs in relation to a LOF; research on
cape ground squirrels (\emph{Xerus inauris} ) proposed the existence of
a correlation between the distance from shelter and the relative risk of
predation within a habitat (Van Der Merwe \& Brown, 2008). Multiple
variables may alter the extent of predation risk and avoidance
behaviours within a LOF; individuals will tolerate higher risk upon
exposure to stress imposed by drought, blight, infectious disease, and
parasites (Real \& Caraco 1986; Raveh \emph{et al.} 2011;
Bleicher\emph{et al.} 2017). Prey may reduce predator avoidance if the
predation is less threatening than death by another means; African
ungulates will visit predator-infested watering holes to avoid
dehydration (Lima \& Dill 1990; Haythornthwaite \& Dickman 2006; Tadesse
\& Kotler 2011; Iribarren \& Kotler 2012; Bleicher \& Dickman, 2016).

\textbf{Methodology}

For the preparation of manuscripts and research, we conducted an
extensive literature search for manuscripts that covered the terms
`landscape of fear', `animal communication', `giving up density',
`predation risk' and `ecology of fear', and related terms. Searches were
conducted via Google Scholar, Web of Science, JSTOR, PeerJ, and Wiley
Interscience Online Library. Literature deemed relevant in the aspect of
calculation of landscape of fear, interpretation of the landscape of
fear, the spatial distribution of predation risk, future recommendation
of the concept and the so-called limitation of the concept were included
in the database and proceeding reference ist. Every article was studied
based on the (1) type of the article (factual, review, opinion), (2)
types and numbers of predator/prey studied, (3) sensory modality used in
the study and (4) results/interpretations that bring larger
understanding about the concept. Much of this publication was inspired
by three manuscripts not attached in the reference list (since they are
unpublished and not peer-reviewed); Kathan Bandyopadhyay's MSc thesis
(Bandyopadhyay 2019), Sam Ashby's BSc thesis (Ashby 2020) and Warren's
BSc thesis (Warren 2020).

\textbf{Discussion}

The ecology of fear drives demographic and community level changes in
wildlife systems (Zanette \emph{et al.} 2011; LaManna \& Martin 2016;
Suraci \emph{et al.} 2016). The impacts of the landscape of fear may be
measured, in their simplest form, by alterations in prey behaviour as a
result of the perceived predation risk across a given landscape (Schmitz
et al. 1997; Brown \& Kotler 2004; Gaynor \emph{et al} . 2019). While
research has covered the more advanced aspects of the LOF concept and
its implications, such as energy expenditure and trade-offs, the
following aspects remain somewhat overlooked:

\emph{Anthropogenic disturbance}

Darimont \emph{et al.} (2009) describe humans as a `super-predator',
having removed single or multiple species from a multitude of
landscapes. The elimination of such species, particularly mammalian
megafauna, may be a significant driver of change in a predator-prey
relationship and prey behaviour within a LOF (Oriol-Cotterill \emph{et
al.} 2015; Suraci \emph{et al.} 2019). Recently, research has indicated
the significance of a fear response to human activity
(Fern\selectlanguage{ngerman}ández-Juricic\emph{et al.} 2005; Stankowich \& Blumstein 2005;
Bateman \& Fleming 2017). It is yet to be ascertained, however, whether
the presence of humans as a `super-predator'(Kotler 1984) , the noise
associated with human activities, or a combination of both results in
behavioural alterations (Frid \& Drill 2002; Stankowich 2008).

\emph{Multiple landscapes}

It should also be acknowledged that the aim of a landscape scale study
is to investigate responses of a number or group of individuals with the
intent to apply the results to concepts on a large scale. Though there
are many publications describing the empirical findings of a LOF applied
to an entire habitat or community, it becomes difficult to blanket a set
of results to the global population of a given species (Fahrig 2005).
Unless a species is endemic to a small territory with limited
disturbance from foreign communities or human activities, each
population of a globally distributed species is likely to experience
varied geographic and ecological aspects affecting LOF responses.
Therefore, single-landscape LOF studies must be reviewed and applied
with caution, especially when conservation efforts are being considered.
Additionally, maximizing the number of landscapes and landscape types
studied can lessen the logistical effects, such as random sample errors,
occurring in a single landscape investigation.

\emph{Neurobiological adaptations of LOF}

Post-traumatic stress disorder (PTSD) is, arguably, the most salient
example of fear and chronic stress shaping the condition of prey
individuals (Clinchy \emph{et al.} 2013). A suitable animal model should
utilize stressors that emulate behavioural, physiological and
neurological responses elicited by a predator (Rosen and Schulkin, 1998;
Rosenboom \emph{et al.} , 2007; Stam, 2007; Armario\emph{et al.} 2008).
Predator exposure often elicits long-lasting effects, including
anxiety-like behaviours, glucocorticoid alteration, dendritic
morphological change, gene expression and the release of
corticotrophin-releasing hormone (CRH) in the amygdala (Adamec \&
Shallow 1993; Schulkin \emph{et al.} 2005; Rosenboom \emph{et al.} 2007;
Takahashi \emph{et al.} 2005; Staples \emph{et al.} 2009).

{\label{post-traumatic-stress-disorder-ptsd-is-arguably-the-most-salient-example-of-fear-and-chronic-stress-shaping-the-condition-of-prey-individuals-clinchy-et-al.-2013.-a-suitable-animal-model-should-utilize-stressors-that-emulate-behavioural-physiological-and-neurological-responses-elicited-by-a-predator-rosen-and-schulkin-1998-rosenboom-et-al.-2007-stam-2007-armario-et-al.-2008.-predator-exposure-often-elicits-long-lasting-effects-including-anxiety-like-behaviours-glucocorticoid-alteration-dendritic-morphological-change-gene-expression-and-the-release-of-corticotrophin-releasing-hormone-crh-in-the-amygdala-adamec-shallow-1993-schulkin-et-al.-2005-rosenboom-et-al.-2007-takahashi-et-al.-2005-staples-et-al.-2009.}}

Traditionally, the perspectives of a wildlife ecologist and
endocrinologist regarding the psychological effects of predator exposure
differed. Immunohistochemistry techniques were developed to understand
the expression of \emph{c-fos} in response to predator induced stress
and exposure (Dielenberg \emph{et al.} 2001). Staples \emph{et al.}
(2009) and Mackenzie \emph{et al.} (2004) successfully mapped the
expression of fosB and its protein product as an alternative to c-fos.
Moreover, global gene expression in response to predator exposure was
assessed using cDNA microarray (Rosenboom \emph{et al.} 2007). Recently,
Yehuda \& Bierer (2009) stated the epigenetic modification of individual
differences to the susceptibility of PTSD. Behavioural and physiological
stress studies suggested predator exposure can lead to chronic stress
and long-lasting detriments in prey individuals (Boonstra \emph{et al.}
1998). Few studies, however, determine the impacts of predator exposure
on population structure and reproduction. Primarily, time and space
limitations obstruct the quantification of reproductive depression as a
result of exposure-induced stress in experimental studies; while
free-ranging individuals can avoid and, therefore, minimize predator
encounters, those in artificial or manipulated enclosures are less able
to do so (Stankowich \& Blumstein 2005; Clinchy \emph{et al.} 2013).
Artificial LOF manipulations enable the recording of immediate or
lasting neurological impacts of exposure. For instance, magnetic
resonance imaging (MRI) data have evaluated the immediate neurological
effects of predator odour exposure on rodents (Chen \emph{et al.} 2009;
Febo \& Pira 2011). Implications upon a community scale and across
generations, however, may remain unattended as a result of limited time
and materials.

{\label{traditionally-the-perspectives-of-a-wildlife-ecologist-and-endocrinologist-regarding-the-psychological-effects-of-predator-exposure-differed.-immunohistochemistry-techniques-were-developed-to-understand-the-expression-of-c-fos-in-response-to-predator-induced-stress-and-exposure-dielenberg-et-al.-2001.-staples-et-al.-2009-and-mackenzie-et-al.-2004-successfully-mapped-the-expression-of-fosb-and-its-protein-product-as-an-alternative-to-c-fos.-moreover-global-gene-expression-in-response-to-predator-exposure-was-assessed-using-cdna-microarray-rosenboom-et-al.-2007.-recently-yehuda-bierer-2009-stated-the-epigenetic-modification-of-individual-differences-to-the-susceptibility-of-ptsd.-behavioural-and-physiological-stress-studies-suggested-predator-exposure-can-lead-to-chronic-stress-and-long-lasting-detriments-in-prey-individuals-boonstra-et-al.-1998.-few-studies-however-determine-the-impacts-of-predator-exposure-on-population-structure-and-reproduction.-primarily-time-and-space-limitations-obstruct-the-quantification-of-reproductive-depression-as-a-result-of-exposure-induced-stress-in-experimental-studies-while-free-ranging-individuals-can-avoid-and-therefore-minimize-predator-encounters-those-in-artificial-or-manipulated-enclosures-are-less-able-to-do-so-stankowich-blumstein-2005-clinchy-et-al.-2013.-artificial-lof-manipulations-enable-the-recording-of-immediate-or-lasting-neurological-impacts-of-exposure.-for-instance-magnetic-resonance-imaging-mri-data-have-evaluated-the-immediate-neurological-effects-of-predator-odour-exposure-on-rodents-chen-et-al.-2009-febo-pira-2011.-implications-upon-a-community-scale-and-across-generations-however-may-remain-unattended-as-a-result-of-limited-time-and-materials.}}

\section*{\texorpdfstring{\emph{Use of Automated Behaviour Response
System}}{Use of Automated Behaviour Response System}}

{\label{use-of-automated-behaviour-response-system}}

As documented by Suraci \emph{et al.} (2016), most of the modern camera
traps are capable to record both audio and video at the same time,
though a small subset of the camera traps studies have made use of the
capability to record the behavioural responses of the animals in
real-time (Rowcliffe \emph{et al.} 2012; Weinstein 2015). Justin Suraci
and their research team provided ABR (Automated behavioural response
system) which can be used as an acoustic cue to design the LOF study.
Using the ABR, they successfully tested the responses of species ranging
from elephants (5250 kg) to squirrels (0.2 kg). Since 2016, playbacks
experiments have been used to test the behavioural responses of species
with an impressive acoustic range, including the vocalizations of the
other species (Blumstein 2014; King 2015), and anthropogenic noises
(Francis \& Barber 2013). Through this study we can integrate recent
models of LOF to real-time conservation practices (Berger-Tal \emph{et
al.} 2011)

\textbf{Conclusion}

Fear driven by both real or perceived predation risks influences the use
of habitat across a landscape for a wide array of prey species, which
can have important consequential effects on the quality of habitat
available. The concept of a ``landscape of fear'' provides an insight
into the habitat preferences of an individual across the landscape and
can reveal important aspects of the ecology of a species, with
potentially important consequences for conservation.

The ecology of fear plays a dual role in prey habitat selection. The
presence of a predator directly impacts the movement of a prey
individual as they alter their behaviour to reduce the chance of
predation, However, even in the absence of any predators, a prey
individual/population may begin to avoid an area over time if they have
encountered a predator there previously. This indirect influence on
habitat and movement ecology can be equally as impactful as direct
predator presence.

Historically, studies have used LOF models to understand the concept of
GUDs and quitting harvest rate within a certain landscape. Today, LOF
models are advanced and incorporate the vigilance behaviours that
mammalian and non-mammalian prey species elicit in response to the
perceived presence of predators. However, the use of audible or chemical
cues to determine the change in prey harvest rate and feeding time is
still vastly understudied. Whilst LOF models often understand the
cause-effect scenario in order to identify and minimize the predation
risk, they are so far unable to sense the neurological and/or chemical
trigger of the vigilance behaviour and the source of fear. In this
review we have laid out the conventional use of chemicals through
pheromone and animal communication studies. For instance, many predators
use pheromones and scent marking in territorial behaviours, profiling
his/her age, health status, social status but how this induces a fear
driven response in prey species is not conclusive. Recent studies of LOF
models include three-dimensional (scatterplot) contour maps,
encompassing spatial statistics in current ecology. However, future
studies should test four-dimensional or five-dimensional models to cover
altitude, time, weight, windspeed, waterflow and other ecological
phenomena. Lastly, studies on the ecology of fear are extremely
effective at signifying population trends and, thus, can be implemented
within conservation strategies for species of conservation concern.
Ecological implications of a fear-mediated response not only provide an
insight into interspecific interactions and movement ecology but, also,
aid conservation agencies and citizen scientists in the protection of
indigenous, charismatic, and keystone species.



\textbf{Acknowledgements}

We would like to thank Dr Philip J. Baker, Dr Manuela Gonzalez Suarez,
Dr Chris Foster and Dr Graham Holloway for preservation and assistance
that inspired this research. We would like to extend a special thanks to
our fellow colleagues and peers for insightful discussions about the
content of the paper.

\textbf{Additional Information and Declaration}

\emph{Funding}

The authors received no funding for this work.

\emph{Competing Interests}

The authors declare there are no competing interests.

\emph{Author Contributions}

Kathan Bandyopadhyay, Samuel S. Ashby and Georgia M. Warren formulated
and designed the paper, wrote the paper and prepared figures and/or
tables, reviewed drafts of the paper.

\emph{Data Availability}

This is a review manuscript. The research in this article did not
generate, collect or analyse any raw data or code.

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