Copyright © 2024 by the author(s). Published here under license by the Resilience Alliance. Open Access. CC-BY 4.0
Sakurai, R., T. Uehara, H. Tsunoda, H. Enari, R. C. Stedman, and A. Onuma. 2024. Developing a system model for articulating thesocial-ecological impacts of species reintroduction. Ecology and Society 29(2):9. https://doi.org/10.5751/ES-14952-290209

Synthesis

Developing a system model for articulating the social-ecological impacts ofspecies reintroduction
Ryo Sakurai 1, Takuro Uehara 1, Hiroshi Tsunoda 2, Hiroto Enari 3, Richard C. Stedman 4

and Ayumi Onuma 5

ABSTRACT. Reintroducing locally extinct/extirpated species has been considered as an approach for restoring ecosystems. Althoughsuch projects share the same goals of rebuilding previously affected ecosystems, the overall impacts that such reintroductions generate
on both ecosystems and human society, i.e., on the social-ecological system, are difficult to measure. We propose a system dynamics
approach, a platform on which both natural and social scientists could collaborate to identify the social-ecological impacts of species
reintroduction as well as factors that affect such decision making. We use cases in Japan to demonstrate the potential applicability ofsystem dynamics in terms of (1) understanding the impacts of a previously reintroduced species, the Oriental Stork (Ciconia boyciana),and (2) predicting the impacts of reintroduction of wolves (Canis lupus). We present a causal loop diagram of the social and ecologicaleffects of Oriental Stork reintroduction, and we discuss how the relationships between factors could be articulated based on empirical
data and ongoing projects in Japan. The model demonstrates how local residents began to appreciate the rich biodiversity, including
the Oriental Stork, following its reintroduction, and how public support toward such reintroduction enhanced further projects toreintroduce these species in different parts of Japan. A similar diagram, created to illustrate the social and ecological effects of thepotential reintroduction of wolves to Japan, demonstrates how social factors such as environmental education and public attitudes
could affect decision making as well as ecological factors such as predator-prey dynamics and overall biodiversity. Further, humanwolf conflicts could negatively affect the overall loop. Creating causal loop diagrams can help managers and stakeholders understandthat species reintroduction projects need to be considered via an interdisciplinary approach. The models illustrate that these problems
are dynamic and that the factors affecting or affected by such projects change over time, implying the importance of both the spatial
and temporal scales in managing reintroduction projects.
Key Words: causal loop diagram; environmental education; Oriental Stork; social-ecological system; wolf
INTRODUCTION
Reintroducing locally extinct/extirpated species is considered as
one approach for restoring ecosystems, and various projects havebeen carried out globally, e.g., wolf (Canis lupus) reintroductionin the United States (Smith et al. 2003) and beaver (Castor fiber)
reintroduction in northern England (Campbell-Palmer et al.
2016). Although diverse goals and approaches for ecosystem
restoration have been considered, from minor modification tofacilitating faster changes via taxon substitution and theacceptance of novel ecosystems (Corlett 2016), there seems to beagreement among researchers and practitioners that reintroducing
once-extirpated species can be a valuable way to conserve andrestore biodiversity (Jachowski et al. 2016). Although these
reintroduction projects share the same goals of rebuilding
previously affected ecosystems, the overall impacts that such
reintroductions have on both the ecosystem and the humansociety, i.e., the social-ecological system, are difficult to measure.
The social-ecological system framework incorporates subsystems
such as resource and governance systems that go beyond
disciplinary boundaries such as ecological science and social
science (Ostrom 2009). The increasing knowledge regarding
reintroduction science indicates that a similar multi-disciplinary
approach is required. Ecological research has been conducted tounderstand how reintroduced species affect the behavior andpopulation of other species, as well as the overall ecosystem. In
Yellowstone National Park (hereafter, YNP), after the
reintroduction of wolves from Canada in 1995, the behavior of
the prey species (elk, Cervus canadensis) changed, which in turn1

caused the underbrush to revive and restored populations of otherspecies such as beaver (Beschta and Ripple 2016). Furthermore,social science research has been conducted, for example, tounderstand local residents’ and visitors’ willingness to support
the wolf reintroduction in and around YNP (Bath 1989).
However, such reintroduction projects impact both ecological andsocial systems simultaneously, particularly if the reintroduced
species are keystone species that have considerable impacts on theenvironment. To understand the overall effects of reintroductions,researchers should explore the social-ecological systemframework, a model that highlights how people and nature are
dynamically interconnected and co-evolving (Hanspach et al.
2014). Species reintroductions demonstrate clearly how ecologicaland social factors are connected in the real world. For example,
social factors can have ecological effects, such as when public
opinion (e.g., support for wolf reintroduction—a social factor)
affects the decision making of the agency regarding species
reintroduction, thus affecting the whole ecosystem (e.g., via the
dynamic relationship between the reintroduced predator and its
prey species—an ecological factor). Alternatively, ecologicalfactors can have social effects, such as when a rapid increase in a
species (such as deer) that causes conflicts with people (anecological factor) reduces local residents’ willingness (a social
factor) to accept that species. Therefore, in order to reveal andpredict the overall impacts of reintroduction projects, future
research needs to identify and investigate the key factors involved,
and their relationship with the social-ecological system. Althoughknowledge from different disciplines is indispensable to identify

College of Policy Science, Ritsumeikan University, 2Center for Environmental Science in Saitama, 3Yamagata University, 4Department of Natural
Resources and the Environment, Cornell University, 5Keio University

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the relationships between ecological and social factors, there has
been limited effort to tackle this issue via collaboration between
natural and social scientists in the field of reintroduction science(George and Sandhaus 2016).
System dynamics, which aims to identify the dynamic behavior
of complex systems, provides a potential platform in which
natural and social scientists (and potentially researchers fromother academic fields) could collaborate to identify the links
between ecological and social factors. The first step in the system
dynamics approach is to visualize a system’s structure by
constructing causal loop diagrams (CLD) showing the feedback
effects in a system (Brennan et al. 2019). Reintroduction of species
generates complex and diverse effects in the social-ecologicalsystem, and requires system-level thinking, which encourages
consideration of the interrelationships and dynamics of the wholesystem. For this reason, we believe that applying the system
dynamics approach is well-suited to our case studies. Although
the system dynamics and social-ecological system approaches are
similar in considering all of the factors of one system, the former
has advantages in terms of being able to qualitatively andquantitatively identify relationships between the factors. On theother hand, in a system dynamics model, the researcher mustchoose which factors to include or exclude.
Here, we present cases from Japan to demonstrate the potential
applicability of system dynamics. First, we examine the impacts
of a previously reintroduced species, the Oriental Stork (Ciconia
boyciana). Second, we predict the impacts of a potential
reintroduction of wolves in Japan. We developed causal loop
diagrams based on previous studies as well as discussion amongsix authors of this paper on factors to include in models anddirections of influences of each factor. In the field of ecosystem
restoration, various terms, including “restoration,” “conservationtranslocation,” and “rewilding” have been used (Corlett 2016). In
this paper, we use “reintroduction” to mean the release of a species
back into its natural range from which it had previously been
extirpated by humans (Jachowski et al. 2016). In this study,“reintroduction” has similar meaning to “trophic rewilding,” asthe species we discuss here (the Oriental Stork and wolf) are the
top predators in each ecosystem.
LOOP DIAGRAM BASED ON A PRIORREINTRODUCTION
The Oriental Stork is one of the species that went extinct in Japan
during the rapid economic growth period (1960s—1980s). For
multiple reasons such as hunting, habitat loss (including
deforestation), and the decline of its prey in paddy fields (becauseof the introduction of modern rice-farming methods such as theuse of pesticides), the stork population had gradually declined
since the 1900s, and the last stork was captured in Toyooka Cityin Hyogo Prefecture (West Japan) in 1971 (Naito et al. 2011,
Yamada et al. 2019). Meanwhile, efforts to increase the numberof Oriental Storks in captivity continued, and six storks weretranslocated from the former Soviet Union for captive breeding
in 1990. In 2000, the number of Oriental Storks in captivity had
increased to more than 100. Consequently, five storks werereintroduced into the wild area of Toyooka City in 2005; with thesubsequent release of additional storks, there are currently morethan 300 wild storks in and around Toyooka, as well as in otherregions of the country (Hyogo Park of the Oriental White Stork

2024). Oriental Storks can cause conflicts with people as thisspecies frequently walks in paddy fields to feed on aquaticorganisms, and their stamping on the rice angers the farmers
because the rice is then unsaleable (Kikuchi 2006). Oriental Stork
also sometimes nest on utility poles on private property, causing
conflict with local residents (Kobe Shimbun 2021). However,
although human-stork conflicts occur, they seem to be tolerated
by most people, as local residents generally support stork
reintroduction (Honda 2008, Takahashi and Honda 2016).
Here, we present a loop diagram for the Oriental Stork
reintroduction in Toyooka (Fig. 1). A study conducted before thereintroduction revealed that local residents were already positive
about the project, and supported it because they acknowledgedthat the presence of storks is indicative of a rich habitat andenvironment (Honda 2008). Another study reported that attitudes
toward stork reintroduction differed with the respondent’s age,with the younger generation being less supportive (Honda 2008).
However, the support of the younger generation toward stork
reintroduction increased as a result of learning about the projectthrough the school curriculum and media (Honda 2019, Sakurai
et al. 2022). The success of the Oriental Stork reintroduction in
Toyooka, and the fact that local residents supported the project,encouraged the local government in other regions to implement
similar reintroduction projects, and they were reintroduced in2015 into Chiba prefecture (eastern Japan) and Fukui prefecture(central Japan; Ezaki and Ohsako 2019). Figure 1 illustrates how
environmental education (EE) enhanced the positive attitude of
the public toward Oriental Stork reintroduction (R1: Socialecological loop with a specific focus on EE and stork
reintroduction).
Oriental Storks are carnivorous birds; based on their gape size,
they can feed on many creatures from small fish to snakes androdents, and are considered one of the top predators in paddy
fields and the surrounding environment. Following theirreintroduction, local residents began to appreciate the richbiodiversity (Kikuchi 2017; supporting R2 in Fig. 1: EE, cognitiveelements and biodiversity loop). Not only did they recognize thesignificance of living in a landscape with diverse wildlife, but they
took pride in having the Oriental Storks in the region and seeing
Oriental Storks in flight and walking in the paddy fields(Evaluation Committee of Oriental White Stork Reintroduction
Project 2014, Kikuchi 2017).
In Toyooka, after the reintroduction of Oriental Stork, a citywide school curriculum, “Furusato Kyoiku” (meaning “education
about local areas” in Japanese), was developed in 2017; in thiscurriculum, all elementary and junior high school students in thecity study Oriental Storks (Honda 2019). This program enrichedstudents’ understanding about biodiversity and the significanceof wildlife reintroduction, and potentially enhanced their
willingness to conserve biodiversity and Oriental Storks (Honda
2019). This supports the EE, cognitive elements and biodiversity
loop (R2) of Figure 1. The number of Oriental Storks has been
increasing (because of both reproduction of storks in the wild
and additional reintroduction) since their first reintroduction in2005 (Ezaki and Ohsako 2019), and Oriental Storks have showna preference for paddy fields where organic farming has been
implemented, and thus for greater biodiversity (Nishimura andEzaki 2019). This has encouraged farmers who want to conserve

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Fig. 1. Causal loop diagram showing relationships between socialand ecological factors in terms of environmental education (EE)and Oriental Stork (Ciconia boyciana) reintroduction in Toyooka
City (R1). This relationship reflects a reinforcing loop involving
multiple positive relationships. In a reinforcing loop, which lacks
negative relationships (or has an even number of them), changes
in one direction are intensified by further changes in that
direction. In contrast, the relationship between conflicts andpolicy reflects a balancing loop; in such loops, which have an odd
number of negative relationships, a change in one direction iscounterbalanced by one in the opposite direction. This is a
simplified model; in reality, the relationships between factors are
not necessarily linear and are affected by other factors. Human–stork conflicts can include agricultural damage (e.g., stamping in
paddy fields) and property damage (e.g., nesting on poles). The
stakeholders can include local residents, farmers, animal-rights
associations, agencies, and researchers.

nature (including storks) to engage in organic farming, thereby
enriching the biodiversity of the agricultural landscape in the region(Kikuchi 2017). These results support the biodiversity/stork loop
reflected in R3 in Figure 1.
Further, the recognition by government ministries (such as the
Ministry of the Environment, Ministry of Land, Infrastructure,
Transport and Tourism, and Toyooka City) of public support for
Oriental Stork reintroduction has enhanced the movement toreintroduce storks in different parts of the country (Evaluation
Committee of Oriental White Stork Reintroduction Project 2014).
Restoration of stork populations is currently taught in junior highand high school science courses across the country, and the successof reintroduction is repeatedly reported in the media (Sakurai et al.
2022). This increase in environmental education, as well as media
reports on stork reintroduction, have increased awareness andpotentially support for reintroduction in different regions (Kikuchi
2017, Sakurai et al. 2022). This relationship among social-ecologicalfactors (R1, the social-ecological loop, with a specific focus on EE
and stork reintroduction) is supported by the existing data andliterature. Interviews with government officials from Toyooka
City and from the Ministry of the Environment provide evidence
that public support enhances decision making regarding Oriental
Stork reintroduction.
LOOP DIAGRAM BASED ON THE POTENTIAL
REINTRODUCTION OF WOLVESThe Oriental Stork reintroduction loop model, which reveals therelationships among ecological and social factors, may be usefulin predicting the impacts of a potential species reintroduction,that of wolves into Japan. The role of wolves as a top predator
first requires clarification. Not only are predator populations
associated with prey populations, they also affect the entire
ecosystem (Ripple et al. 2014), including ecosystem nutrient
cycling (Schmitz et al. 2010) and herbivore biomass (Letnic and
Ripple 2017), via direct and indirect effects. Loss of top predatorscauses the prey population to increase, which potentially changes
the entire ecosystem (Schmitz et al. 2010, Estes et al. 2011). In
YNP, since the extirpation of wolves in the 1920s, woody species
have declined, primarily owing to heavy browsing pressure by theelk; this in turn destroyed riparian forest, causing the populationof beavers (Castor canadensis) to decline (Ripple and Beschta
2003). In contrast, following the reintroduction of wolves in thepark in 1995, riparian vegetation has been restored, as the elk
population has declined and their behavior has changed; thisdemonstrates the potential effects of a top carnivore in a terrestrial
food chain (Beschta and Ripple 2016). Wolf reintroduction has
also affected bison (Bison bison), coyote (Canis latrans), beavers,
and songbird assemblages, emphasizing the significance of
reintroducing top predators in terms of restoring and increasing
biodiversity (Ripple et al. 2014, Beschta and Ripple 2016).
The Japanese wolf (C. l. hodophilax), the endemic wolf subspecieson the Honshu, Shikoku, and Kyushu islands, disappeared by
1905. It is now classified as an “extinct species” in Japan. The
populations of species hunted by wolves, such as sika deer (Cervus
nippon) and boar (Sus scrofa), have been rapidly increasing in the
country, partially because the ecosystem lacks predators that
could suppress their populations. Other reasons for the increasein ungulate populations include changes in land-use; for instance,
the aging of local residents and depopulation of rural
mountainous regions in Japan have increased the number ofabandoned fields that become overgrown with brush, ultimately
increasing the habitat available for those species (Sakurai 2019).
The population density of deer has increased to more than 20
deer/km² in some regions (e.g., Kishimoto et al. 2010). The risein the wildlife population has also increased human-wildlife
conflict, and the current annual cost of agricultural damage
caused by terrestrial mammals is estimated to be approximately
13 billion yen (about US$130 million; Ministry of Agriculture,
Forestry, and Fisheries 2021).
Although an intermediate density of deer contributes toenhancing local species richness (Suzuki et al. 2013), their
overpopulation can alter mountain forest vegetation throughovergrazing, resulting in fewer evergreen trees and less diversityof the understory (Takatsuki 2009, Ohashi et al. 2014).
Overgrazing owing to deer overpopulation has affected otherfauna in the forest ecosystem, e.g., the faunal composition offorest-dwelling birds and small mammals such as Apodemus mice

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has declined because of a reduction in food resources (Seki et al.
2014). The populations of endemic serow (Capricornis crispus)
have declined, potentially owing to resource competition with sika
deer (Koganezawa 1999, Seki and Hayama 2021). Although large
herbivores can negatively affect insects, a meta-analysis on theindirect effects of herbivore density on phytophagous insects
revealed no clear trends (Takagi and Miyashita 2014).
Based on the situation in Japan and case studies worldwide, wecreated a simple ecological system loop diagram (Fig. 2). Wolves
affect the deer population via predation, while, as its primary prey,
the number of deer affects the wolf population (illustrated byrelationship B3, the predator/prey loop, Fig. 2). Although wolves
also prey on other species such as boar and hares (Lepus
brachyurus), in this study we focus on the relationship between
wolves and deer, because sika deer was historically their staple
prey (Matsubayashi et al. 2017). Wolves can reduce the prevalenceof infections such as tuberculosis in prey populations (e.g., boars)
via predation, thus contributing to maintaining healthy prey
populations without reducing their density (Tanner et al. 2019),
although the relationships between predators and their prey are
more complex than this.
Fig. 2. Causal loop diagram of the relationships between
biodiversity level, deer and wolf populations, and hunters. The
relationships between biodiversity and deer population, thewolf and deer populations, and hunters and deer populations
show balancing loops (B2: Biodiversity/deer loop; B3: Predator/
prey loop; B4: Hunter/deer loop). The relationships among
biodiversity, deer and wolf populations, and biodiversity andthe wolf population show reinforcing loops (R4: Biodiversity/
predator/prey loop; R5: Biodiversity/wolf loop). Other factors
also affect the dynamics of each factor.

Hunting by humans plays a significant role in controlling deer
populations (e.g., Kaji et al. 2010). The growth of deer
populations could increase hunters’ motivation to hunt them,promoting the recruitment of new hunters (relationship B4, Fig.
2: the hunter/deer loop). In Japan, the overpopulation of deer
incentivized government ministries to increase the budget for
deer-population management and for providing hunting subsidies
to hunters; this in turn increased hunters’ motivation and provided
incentives for people to obtain hunting licenses (Igota and Suzuki2008). However, it is important to remember that increase in gamespecies populations does not necessarily lead to an increase inhunters’ motivations, as is illustrated by a case in Spain where

hunters perceived the need to reduce overpopulation by European
rabbits (Oryctolagus cuniculus) as an obligation rather than apleasant activity (Delibes-Mateos et al. 2020). Therefore, we
believe that a combination of hunters and wolves will decrease
deer populations.
The deer population affects biodiversity both quantitatively andqualitatively, as illustrated by case studies worldwide, with a rapid
increase in deer numbers reducing biodiversity. Biodiversity also
affects the deer population, because deer thrive in diverse
landscapes including forests and grasslands, but not inmonoculture plantation forests (Martin et al. 2020). Althoughbiodiversity could affect the wolf population, such a relationshipmight not be linear. Theoretical and empirical research has shownthat greater diversity contributes to greater resilience in theecosystem community (Tilman 1996, Krebs 2001). In a lessbiodiverse environment, wolves might depend solely on deer for
their prey, hence their population could fluctuate rapidly with thechange in the deer population. In many parts of the world, in theabsence of natural prey, wolves have started to rely on livestockand other anthropogenic inputs such as garbage (Newsome et al.
2016). In contrast, a highly biodiverse ecosystem could provide
wolves other prey besides deer (e.g., boars and hares), making it
more likely that the wolf population would stabilize. In that sense,
the R5 and B2 loops in Figure 2 should be interpreted as follows:
the presence of sufficient biodiversity would stabilize (but notnecessarily increase) both the wolf and deer population. The
relationships of biodiversity to deer, on the one hand, and towolves, on the other, differ in that although an increase in the wolf
population could increase biodiversity, deer overpopulation could
reduce biodiversity. In contrast, deer overpopulation would
increase the wolf population by providing additional prey. We
discuss this further in the following section. Japan currently lacks
a top predator (in this case, wolves); therefore, in the CLD
illustrated in Figure 2, only the biodiversity/deer (B2) and hunter/
deer (B4) loops currently occur in Japan, illustrating that deer
continue to negatively affect biodiversity.
In various parts of the world, ungulate population sizes havedeclined or been controlled by epizootic diseases; for instance,mange has impacted the Alpine chamois (Rupicapra rupicapra
rupicapra; Rossi et al. 1995) and tuberculosis has impacted wildboar in Spain (Tanner et al. 2019). Therefore, we have added“Disease” as a potential factor affecting the deer population (Fig.
2). However, in Japan, hunters are considered to be the main factor
controlling the deer population, as indicated by historical deer
population dynamics: in the 1900s, deer were consideredthreatened throughout Japan, because of overhunting; as huntingpressure decreased in the 1990s, deer became overpopulated; sincethe 2000s, the population has been controlled by intense hunting
activity (Kaji and Iijima 2017).
From a social-science perspective, we first present a simple CLD
illustrating the relationship between environmental education andpublic intention to conserve biodiversity (Fig. 3) as a benchmark
for a combined social-ecological model. Assuming that thedecisions of agencies about species reintroduction are affected bypublic opinion and support for reintroduction (e.g., voting infavor of reintroduction, which can be regarded as proenvironmental behavior), it is necessary to understand how such
public opinion and pro-environmental behavior is formed.

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People’s opinions and perceptions regarding a certain topic areshaped and affected by various factors, including education(Ajzen and Fishbein 1980), the media (Takeshita 2008), pastexperiences (Ormrod 2008), and information from peers (Decker
et al. 2012). Environmental education can increase knowledge
about, and foster awareness of, environmental issues, and possibly
encourage pro-environmental behavior (Liddicoat and Krasny
2012, Zint 2012). Although knowledge is thought to have limited
effects in terms of changing behavior (Ajzen 2001, Kollmuss andAgyeman 2002), the combination of three types of knowledge—
systems, action-related, and effectiveness-based—could lead topro-environmental behavior (Krasny 2020). In this case, people
could gain system knowledge by learning about, for example, how
the ecosystem functions, in terms of species interactions andpredator/prey relationships. People can gain action-related
knowledge by discovering, for example, that they can participate
in nature observation and restoration activities in theirneighborhoods. Finally, effectiveness-based knowledge can begained by understanding that, for example, voting for ecologicallyliterate candidates and communicating with elected officials (orin this specific case, supporting organizations that promote
species reintroduction) are effective ways to promote biodiversity.
Fig. 3. Causal loop diagram illustrating the relationships
between environmental education (EE), knowledge (systems,
action-related, and effectiveness-based) about biodiversity andconservation, intention to conserve biodiversity, and demandfor environmental education. The relationships among thesefactors are in a reinforcing loop (R6: Reinforcing intention
through EE loop). Other factors also affect the dynamics ofeach factor.

Based on previous studies, a simple CLD regarding the socialscience process can be created (Fig. 3). Environmental education
can increase public understanding about biodiversity andconservation (Ernst et al. 2009). With the increase in public
understanding of biodiversity (in terms of all three types ofknowledge about biodiversity conservation), their intention toconserve biodiversity as well as actual behavior could be

encouraged (Jacobson 2009, Sakurai et al. 2019). Meanwhile, anincrease in public intention to conserve biodiversity could increasedemand for environmental education programs, potentially
increasing the number of environmental education programs
conducted nationwide. As more people engage in conservationrelated behavior, this could alter social norms, such as byincreasing the social pressure from family, neighbors, and others
to engage in pro-environmental behaviors (Hayabuchi 2008). This
could in turn cause additional environmental education programs
to be implemented to meet this need (Sasaki 2016). Althoughconservation initiatives and projects are sometimes implemented
without specific environmental education programs, we believethat including environmental education programs could makesuch initiatives more effective and sustainable, as the projects may
then have long-term public support (George and Sandhaus 2016).
The Oriental Stork reintroduction case demonstrates thepotential connections between the frequency of environmental
education, public knowledge about biodiversity, public intention
to conserve, and the demand for environmental education (Fig.
1).
In the real world, ecological and social factors are connected. For
example, increased public participation in conservation(potentially via an increase in the number of environmental
education programs) could preserve or increase the biodiversityof the surrounding environment. Citizen science projects
worldwide have demonstrated how local environment andbiodiversity can be conserved through active involvement by
citizens (Bonney et al. 2014, Kobori et al. 2016).
Although the reintroduction of once-extirpated top predators canrestore self-sustaining ecosystems and enrich their biodiversity, itremains a policy decision whether or not to reintroduce such a
species. Such policy decisions are affected by public opinion(especially in democratic countries such as Japan). The model that
integrates ecological and social causal loops (Figs. 2 and 3)
illustrates how social factors affect ecological factors and viceversa (Fig. 4).
People who understand the ecological roles of wolves are more
likely to support wolf reintroduction (Gosling et al. 2019, Sakurai
et al. 2020, Sakurai et al. 2023); in many cases, such knowledge
is acquired through education (Jacobson 1999, Musiani andPaquet 2004). Knowledge of wolves’ roles in the ecosystem(systems knowledge), of the types of activity required to support
NGOs that promote wolf reintroduction (activity-basedknowledge), and of how supporting such NGOs can influence
policy (effectiveness-based knowledge) is likely to foster public
support for wolf reintroduction. People with positive attitudes
toward reintroduction are more likely to act in support of suchpolicies (e.g., voting for reintroduction; Sakurai et al. 2023).
Although we started our discussion here focusing on a simple
model considering the general public, stakeholders that arepotentially affected by wolf reintroduction, including hunters andranchers, would have different opinions toward wolves that are
not necessarily affected by environmental education. We discuss
this in detail in the following section.
In Japan, wolf reintroduction will depend on the amount of public
support for the government’s policy (Yamanaka and Kaji 2006,
Kaji 2017). In the U.S., which has a similar democratic system,
Colorado has decided to reintroduce wolves by 2024 (and initialEcology and Society 29(2): 9
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Fig. 4. Causal loop diagram illustrating the relationships between social and ecological factors in terms ofenvironmental education (EE) and wolf reintroduction (R7). Other factors also affect the dynamics of each factor.
Human–wolf conflict can include the loss of pets and livestock, polarization over support for wolf conservation, andurban/rural differences. The stakeholders include local residents, farmers, hunters, animal-rights associations, agencies,and researchers.

reintroductions were completed in December 2023), as a 2020
ballot indicated increased public support for it (NPR News 2020).
In Japan, it is not uncommon for the public to influence policyby showing their preferences in a ballot or signature campaign(Murayama 2009). The Japanese public increasingly expects toparticipate in decision making, and to have their values and beliefs
reflected in decisions; acceptance of the ballot procedure as animportant process in structuring and implementing policies is
growing (Murayama 2009). It is therefore likely that the public
will decide on wolf reintroduction in Japan by ballot, as inColorado.

decisions. (In the different CLDs presented, the same loop
numbers are used when the contents of the loop are same, suchas for relationship B2 [the “Biodiversity/deer loop”] in Figs. 2 and
4). If the government decides to reintroduce wolves as a national
policy, a new wolf population will be released and established in
the country. The presence of wolves would directly and indirectly
affect deer behavior and the deer population; once the deer
population has relatively stabilized given wolf presence, otherspecies populations are expected to increase, and the overallbiodiversity level could increase (as indicated in R4: Biodiversity/
predator/prey loop).

Lobbying can also play an important role in environmental
decision making. In Spain, a formal request by a conservationrelated NGO to the government succeeded in having wolves listed
in the Spanish Red list of Threatened Species, thus banning
recreational hunting as a wolf management option (Ordiz et al.
2022). Environmental issues are among the top lobbying agendas
in the U.S. (Baumgartner et al. 2009). For Japan, the potential
effects of lobbying on decision making need to be considered.

An increase in biodiversity could increase opportunities for the
public to learn about and experience biodiversity through, forinstance, eco-tours or citizen science activities. This, in turn, would
foster public awareness of the benefits of a biodiverse
environment (as illustrated in R2: EE, cognitive elements andbiodiversity loop in Figs. 1 and 4). This is because people begin
to understand ecological processes and how biodiversity
functions by learning about changes in the country’s biodiversity
via the media or by visiting such areas. In YNP, the wolf
reintroduction program provided visitors with opportunities tolearn about the ecological roles of wolves and about wholeecosystem functioning, irrespective of whether they actually saw
wild wolves in the park (Montag et al. 2005). Public attitudes

With this in mind, relationship R7 (the social-ecological loop witha specific focus on EE and wolf reintroduction) in Figure 4
illustrates how environmental education could increase publicknowledge of the role of wolves in ecosystems, and how public
support for wolf reintroduction policies could affect government

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toward wolves are expected to become more positive once theylearn about the ecosystem services that wolves provide. Seeing theincrease in biodiversity and the value of reintroduced species canshape people’s vision of the desired state of biodiversity. For
instance, once species have been reintroduced or their populations
restored, local residents are more likely to accept those species
and recognize their importance in the ecosystem, such as for the
Oriental Stork reintroduction in Japan (Honda 2008, Kikuchi
2017, Sakurai et al. 2022) and human-wolf coexistence in Spain(Pettersson et al. 2021).
Wolves can contribute to disease control in their prey populations,in turn reducing transmission from wildlife reservoirs to otheranimals such as cattle; wolves could thus potentially be recognized
as allies, even by those who are expected to oppose wolf
reintroduction, such as farmers (Tanner et al. 2019). Species
reintroduction could increase the number of environmental
education programs implemented, as reflected in the various wolfwatching tours and educational programs offered in YPN related
to the wolf reintroduction (Miller 2017).
LIMITATIONS IN CREATING A REALISTIC CLD
Creating diagram-based models helps stakeholders understandthe implications of management decisions (van den Belt 2004).
Such diagrams enable policy makers to see the “big picture,”
overcoming the typical tendency toward short-term linear
thinking in decision making (van den Belt 2004). However,diagrams are incomplete and cannot fully explain the real world(Bueno 2014). Here, we selected the factors to construct the CLDs,
but there are other factors that also affect ecological and social
processes. For example, deer populations are affected not only bynatural predators, human hunters, and disease, but also by otherfactors such as the weather. For instance, increasing snow depth
interacted with wolf predation to slow the population growth of
elk in Alberta, Canada (Hebblewhite et al. 2002). Further, while
human-wolf conflicts are included and explained in Figure 4, thecomplexity and depth of the conflict cannot be fully captured andexplained in this model. Adding additional factors to the diagram
would provide a better representation of reality, but would
increase the model’s complexity. However, to be able tounderstand the dynamics of the system and predict futurescenarios, it is more important to identify the relationships among
the variables (thus explaining the system’s dynamic complexity),rather than accurately verifying the linear cause-and-effectrelationships between the factors (van den Belt 2004, Hovmand2014).
The validity of some of the relationships illustrated in Figure 4
is debatable. For instance, considering relationship R5 (Fig. 4),
while wolves could affect the deer population, and deer
overpopulation could reduce biodiversity, more research isnecessary to clarify whether and how biodiversity levels affectwolf population size. When wild ungulate densities are low, or
human-subsidized food sources (such as refuse) are easily
accessible, wolves can live near human settlements by feeding on
discarded household waste, or by preying on livestock (Newsome
et al. 2016). However, this potentially provokes conflicts withhumans (Kuijper et al. 2019). It also increases the risk of pathogen
infection (Plaza and Lambertucci 2017), potentially increasing
the risk of casualties from attacks by rabid wolves (Linnell et al.

2021). Historically, such increases in human-wolf conflict haveoften led the authorities to eradicate wolves that live near humansettlements (Nie 2003). Nonetheless, in many parts of the world,
wolf populations have not only been maintained but haveincreased in spite of human-wolf conflicts such as livestock
predation (Newsome et al. 2016). In contrast, high biodiversity
encourages the long-term and sustainable survival of a stable wolf
population (as simplified by “+ (positive)” in Figs. 2 and 4).
Although some studies have revealed how wolves affect the levelof biodiversity via cascade effects (Smith and Bangs 2009, Ripple
et al. 2014), fewer have examined how the level of biodiversity
affects wolf populations. Further studies are necessary toarticulate this relationship in greater detail. The factors
potentially affecting public attitudes toward wolf reintroduction,
as described in the literature, are summarized in Table 1.
The CLD in Figure 4 assumes that wolf reintroduction would
have positive outcomes, such as increasing biodiversity andgenerating public support for even more wolf conservation andreintroduction. However, it could also negatively affect humans
if, for example, human-wolf conflicts increased after the project.
In many parts of the world, ranchers, livestock farmers, andhunters typically have negative attitudes toward wolf
reintroduction or conservation (Bath 1989, Williams et al. 2002).
An increase in the wolf population can increase human-wolf
conflict and consequently reduce public support for wolf
conservation (Eriksson and Herberlein 2003, Pettersson et al.
2021). If this occurs in Japan, conflict such as wolf attacks onpets and livestock could reduce public support for wolf
reintroduction and conservation (Fig. 4). Support for the
reintroduction of large carnivores depends both on humantolerance toward them and conflict reduction (Forbes et al. 2020).
Unlike in the U.S., where wolves have been reintroduced, there isrelatively little livestock production in Japan (the U.S. produces
about 36 times more beef than Japan annually, while it has 2.6
times more residents; U.S. Meat Export Federation 2021).
Therefore, the reintroduction of wolves in Japan could lead toless livestock predation (both in frequency and quantity) than incountries such as the U.S. and European countries.
In Japan, people’s perceptions toward wolf reintroduction areaffected by factors such as fear of wolves, trust in researchers, andwhether or not they believe that reintroduced wolves will becomeinvasive (Sakurai et al. 2020, 2023). In order for a model (such as
the CLD in Fig. 4) to be useful and meaningful for stakeholders,it should reflect context-specific information. For example,
conflicts related to wildlife reintroduction could be classified as
disputes (e.g., disagreements), underlying conflicts (e.g., distrust),
and deep-rooted conflicts (e.g., a lack of recognition of culturalvalues in decision making; Glikman et al. 2022). Therefore,
increasing public support for wolf reintroduction requires morethan simply environmental education; it also requires the creationof diverse opportunities to enhance governance, including
outreach and civic meetings; increasing the public trust in agencies
and researchers; promoting the dissemination of fact-based
information about wolf-attack frequency; and for decision makers
to learn about the local culture and context in the areas where
wolf reintroductions are planned (Massarella et al. 2021, Glikman
et al. 2022).

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Table 1. Factors potentially affecting public attitudes toward wolf reintroduction (WR) and conservation.
Items

Relationships identified by previous studies

Knowledge about wolves

Effects on public
attitudes toward WR

More knowledge about wolves leads to more support for WR and wolf conservation (Angeli et al. 1998, Enckand Brown 2002, Gosling et al. 2019, Sakurai et al. 2020). In contrast, hunters with the most accurate
knowledge about wolves have the most negative attitudes (Eriksson and Heberlein 2003).
Attitudes toward wolves
More positive attitudes toward wolves lead to more support for WR and wolf conservation (Bath 1989, Enckand Brown 2002, Sakurai et al. 2020, 2023).
Knowledge about wildlife issues Increasing public realization that deer populations have grown and cause damage might increase support for
WR (Angeli et al. 1998).
Beliefs related to the role wolves The more critically people think about the ecological role of wolves, the more they support WR (Enck andplay in ecosystem
Brown 2002, Sakurai et al. 2020).
Wildlife-related values andGreater adoption of mutualism as a value leads to more support for WR and wolf conservation (Hermann etbeliefs
al. 2013, Gosling et al. 2019)
Trust in government and other Less trust in politicians leads to less support for WR as a policy (Eriksson 2017) whereas more trust inentities
researchers at universities leads to more support to WR (Sakurai et al. 2020)
Risk perception
The perceived risk of wolf attacks reduces support for WR (Enck and Brown 2002, Sakurai et al. 2020).
Experiences with wolves
More direct experience with wolves reduces support for WR or wolf conservation (Eriksson and Herberlein2013). On the other hand, more experience of wolves could increase support for wolf conservation (Petterson
et al. 2022).
Socio-demographic factors
Livestock farmers oppose WR (Bath 1989) wherease members of environmental groups are more likely tosupport it (Williams et al. 2002).
Local residents living adjacent to wolf habitats or WR sites are less likely to support it, whereas city residents
are more likely to support it (Bath 1989, Bath and Buchanan 1989, Williams et al. 2002, Eriksson andHeberlein 2003, Gosling et al. 2019).
Elder respondents are less likely to support WR (Gosling et al. 2019).
Women are more likely to support WR (Williams et al. 2002)
People with higher incomes are more likely to support WR (Williams et al. 2002)
People with more education are more likely support WR and wolf conservation (Eriksson and Herberlein2003).

Accuracy and correctness are not necessarily the most important
objectives when creating a system dynamics model, although suchmodels must be applicable (Bueno 2014). In that sense,
stakeholders could include factors and relationships in suchmodels to reflect the types of ecological and social processes thatthey want to understand and predict. Therefore, models are
created based on the outcomes of subjective judgment (Stedman
2016); thus, the reliability and validity of the CLDs should beanalyzed and discussed when identifying potential strategies for
improving such models (Cornwell 2004, Hovmand 2014).
CHALLENGES AND POSSIBILITIES IN QUANTIFYINGTHESE RELATIONSHIPSThe system dynamics approach qualitatively and quantitatively
predicts the relationships among factors, providing insights into
the magnitude of change that would occur in the system (van den
Belt 2004). The explanatory power of the model can be
determined based on the adequacy of the data used in the analysis.
For example, previous studies have quantitatively determined how
much knowledge about wolf reintroduction is necessary to change
public attitudes toward wolf reintroduction (Enck and Brown
2002). A study in Japan, based on a survey of university students,
found that increasing knowledge about wolves (e.g., their roles inthe ecosystem and the fact that the reintroduced wolves would be
the same species as continental wolves) would increase support
for reintroduction by on average 0.6 points on a five-point scale(Sakurai et al. 2020).
A recent survey (Sakurai et al. 2023) revealed diverse public
attitudes toward wolf reintroduction: about 40% disagreed withreintroduction while another 40% were unsure about it, and therest (approximately 20%) agreed with it. In the same study,
knowledge that Japanese wolves would be the same species as+ and −

+
+
+
+
+
−
+ and −

+ and −
+ and −−+++

continental wolves, that they can control deer populations, andthat they are necessary for a healthy ecosystem, increased public
support for reintroduction by a total of 0.7 points on a five-point
scale. Those who agreed with wolf reintroduction had greater
intention to engage in behaviors to support it, such as signing apetition to support it. Majority rule is the basis for decision
making in many democratic countries, and whether the ratio of
citizens who agree to a certain topic increased to more than 50%
would be assumed as one social tipping point that could change
the policy. Based on previous studies (e.g., Niemiec et al. 2020,
Sakurai et al. 2020, 2023), we hypothesized that once a majorityof the public obtains scientific information about the identity androle of wolves, they would be more likely to support wolf
reintroduction. Further, as the public shifts toward favoring wolf
reintroduction, more people would engage in activities to support
the reintroduction, and this in turn would encourage or force thegovernment to start considering wolf reintroduction as a national
policy.
By including numerical values into the diagram, the proposedmodel (Fig. 4) could become quantitative. For example, based on
current research on public attitudes toward wolf reintroductionin Japan (17% of citizen supported it; Sakurai et al. 2023), we canassume a value of 0.17 for “Support for wolf reintroduction
policy,” and assume that this value would change as theopportunities for public participation in wolf-related
environmental education would increase. Because there are
currently few opportunities where the public can learn about theecological roles of wolves in Japan, the value for “Environmental
education” is assumed to be 0; however, as more information
becomes disseminated via the media and community lectures, thisvalue (the number of environmental education events each year)

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could increase to, for instance, two per year. Such opportunities
for environmental education about wolves are expected toincrease the “Public knowledge of role of wolves in ecosystem”
parameter (currently at 3.27 on a five-point scale; Sakurai et al.
2023), in turn affecting public support for reintroduction.
Similarly, human-wolf conflicts can be quantified using multiple
values, including the number of casualties and depredation ofpets and livestock. Factors such as the deer population andbiodiversity levels have recently been quantified (e.g., theestimated deer population in Japan in 2021 was 2.46 million;
Ministry of the Environment 2021). However, more research isnecessary to determine the potential sustainable wolf populationof Japan and how it would affect the deer population, consideringfactors such as habitat availability and the rate of fragmentation.
ADDITIONAL FACTORS TO CONSIDER IN
DEVELOPING THE SYSTEM MODEL
To better understand the implications of the proposed model andto account for uncertainty regarding the future scenario, both thespatial and temporal scales should be considered. The spatial scalemust be considered because wolf reintroduction is aimed at
restoring the entire ecosystem; prior reintroduction projects havedemonstrated how reintroduced species (such as Oriental Storks
in Japan and wolves in YNP) have dispersed into other regions.
When considering project feasibility, the Japanese government
therefore needs to understand the attitudes and preferences of
not only the local residents near the potential reintroduction sites,
but also of the broader public. In the case of the stork
reintroduction, while we have focused primarily on Toyooka City
(Fig. 1), the reintroduced storks have dispersed beyond theboundaries of this city, and now occur in various parts of westernJapan. Following stork reintroduction in eastern Japan, storks
now occur there, such as in Chiba prefecture (Takahashi and
Honda 2016). Public perceptions of storks have become morepositive and willingness to accept storks has increased not onlyin Toyooka but also in other parts of Japan (Takahashi and
Honda 2016, Sakurai et al. 2022). Figure 1 thus illustrates not
only what happened in Toyooka, but also what is likely to happen
at a larger spatial scale in future.
The temporal scale should also be considered when developing amodel and predicting relationships between factors. Disseminationof accurate information regarding wolves might occur relatively
quickly because a one-time program can increase participants’knowledge and even change their attitudes (e.g., support for
natural energy increased after one seminar; Takano et al. 2018).
A study in California on the potential reintroduction of grizzly
bears revealed that the public generally lacked knowledge of thespecies and reintroduction, and that wildlife managers thus
needed to communicate the ecological and social benefits ofreintroduction to the public before introducing management
proposals, to prevent the issue from becoming polarizing(Hiroyasu et al. 2019).
At government level, it could take longer to decide to adopt a
policy to reintroduce an extirpated species. In the U.S., although
the importance and possibility of reintroducing wolves in the
YNP had been discussed by researchers and the public since the1960s (Fritts et al. 1997), it took about 30 years for the government(specifically, The U.S. Fish and Wildlife Service) to actually planand implement this project (in 1995). In the UK, although thepotential to reintroduce wolves has been discussed by researchers(e.g., Wilson 2004, Nilsen et al. 2007) and organizations(Rewilding Britain 2021) for more than 30 years, such projects
have not yet become government policy. In Japan, although
researchers and NGOs have been discussing the importance andsignificance of reintroducing wolves for several decades (e.g.,
Knight 2006, Maruyama et al. 2007), this seems to have had
limited influence on the government’s decision making, because
introducing a species that could cause new problems (such as
human-wolf conflicts) is considered a risky project for thegovernment (Kawata 2014).
Once wolves have been reintroduced, the effects on other species
rapidly become visible. In YNP, changes in elk behavior andhabitat selection were reported half a year after the reintroduction(Ripple and Beschta 2003). Reductions in the deer population
were reported a few years after the wolf reintroduction in YNP,
although there were multiple potential reasons for this, including
the weather (Smith and Bangs 2009). The ecosystem of YNP haschanged since the reintroduction, mostly positively, in terms ofgreater ecosystem resilience owing to the increased biodiversity(Smith and Bangs 2009). Such changes in ecosystems andbiodiversity could affect people’s understanding and attitudes
toward reintroduction projects (e.g., Montag et al. 2005). Mostof the millions of visitors to the YNP wish to see wild wolves
during their visit (Miller 2017). Environmental education
programs and outdoor activities have been established to teachvisitors about the importance of wolves in the ecosystem in andaround the YNP (National Park Service 2020), potentiallyinfluencing the participants’ perspectives. In Spain, local
businesses and producers have started using wolf-related
branding, thus successfully attracting tourists (Pettersson et al.
2021).
The number and frequency of human-wolf conflicts can changeover time. Conflicts could increase as the wolf population
increases; once measures to prevent damage (e.g., fencing) or
farmers’ compensation systems have been established, however,such conflicts could be tolerated by those stakeholders (Pettersson
et al. 2022). Once the public start to accept wolves as a species
that belongs in the landscape, their attitudes toward wolves andreintroduction projects can become positive (Gosling et al. 2019,
Pettersson et al. 2022). With ongoing modernization andurbanization, the environmental values of the public havechanged from being domination-based to mutualism-based andfavoring conservation (Manfredo et al. 2021), and this value-shift
is expected to continue. In Japan, people who value mutualism
are more likely to support wolf reintroduction (Sakurai et al.
2023).
It could take more than half a century to observe all the effects
proposed in our models. In terms of the stork reintroduction in
2005, it took less than 20 years for local residents to recognize thebenefits of biodiversity; environmental education activities
increased in the region, encouraging biodiversity conservation
behavior. The reintroduction occurred about 30 years after thelast stork was captured in the region in 1971. The amount of timerequired for a decision on reintroduction therefore varies
depending on the target species, the ecological and social context

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that led to the extirpation, the level of conflict, and public
perceptions toward the species. As has been reported for many
species reintroductions around the world, some species are more
likely to be reintroduced than others (Seddon et al. 2005).
Meanwhile, participatory processes encouraging the involvementof various stakeholders (“Participatory process of management”
in Figs. 1 and 4) can make management decisions more effectiveby ensuring that they have public support. For the stork
reintroduction, collaboration among stakeholders—including
local residents, the city administration, Hyogo Park of the
Oriental White Stork staff and volunteers, local farmers, andnatural and social science researchers—ensured ongoing research
into storks, demonstrated that storks and human can coexist, andled to the success of the reintroduction project (Kikuchi 2006).
In addition, climate change should also be considered. The
climate of Japan might change in the next 50 years.
Economic factors should also be considered in such a model. The
cost of species reintroduction and management is massive. The
reintroduction as well as management of wolves cost more than
US$1 million per state (Hoag et al. 2023). However, the ecosystemservices generated by wolf reintroduction could outweigh thecosts: for instance, the presence of wolves reduced deer-vehicle
collisions by 24%, bringing economic benefits 63 times larger thancosts of wolf depredation on livestock (Raynor et al. 2021). In
Japan, while it cost 5.9 billion yen (about US$59 million) to build
and run the Park of the Oriental Stork, the annual benefits ofthis project have been estimated at about 1 billion yen (about
US$10 million) from tourism and about 8 billion yen (about
US$80 million) from related projects (Onuma and Yamamoto2009). Further environmental economics research is necessary to
clarify the potential economic value of species reintroductions
considering both their resulting ecosystem services and costs.
SUMMARY
In summary, our model predicts that environmental education
programs will enhance people’s attitudes toward wolf
reintroduction; this will influence government’s decision making
regarding wolf reintroduction; the deer population will becontrolled, and total biodiversity will thus increase; this will inturn alter people’s attitudes. Causal link diagrams are valuable
because they can help managers and stakeholders to realize thatthe problems are integrated within the system, highlighting theimportance of an interdisciplinary approach to understanding
the “bigger picture.” Further, CLDs help stakeholders to see
problems as dynamic, providing insights into how the
relationships between factors (and the factors themselves) changeover time. For example, it is easy to imagine that wolf and deer
populations change over time; however, social factors, such as
public perceptions and support for reintroduction, also changeover time, and this can be more difficult for stakeholders to grasp.
A recent survey in Japan revealed that the younger generation
holds more positive attitudes toward wolf reintroduction than theolder generation and that public attitudes could shift (Sakurai etal. 2020). The CLD could be further improved by quantitatively
verifying the relationships represented in each loop; it could thenbe used to describe a set of potential future scenarios.

Data Availability:
Data sharing not applicable to this article as no datasets weregenerated or analyzed during the current study.
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