Understanding managers’ and scientists’ perspectives on opportunities to achieve more evolutionarily enlightened management in conservation

Abstract Despite wide acceptance that conservation could benefit from greater attention to principles and processes from evolutionary biology, little attention has been given to quantifying the degree to which relevant evolutionary concepts are being integrated into management practices. There has also been increasing discussion of the potential reasons for a lack of evolutionarily enlightened management, but no attempts to understand the challenges from the perspective of those making management decisions. In this study, we asked conservation managers and scientists for their views on the importance of a range of key evolutionary concepts, the degree to which these concepts are being integrated into management, and what would need to change to support better integration into management practices. We found that while managers recognize the importance of a wide range of evolutionary concepts for conservation outcomes, they acknowledge these concepts are rarely incorporated into management. Managers and scientists were in strong agreement about the range of barriers that need to be overcome, with a lack of knowledge reported as the most important barrier to better integration of evolutionary biology into conservation decision‐making. Although managers tended to be more focused on the need for more training in evolutionary biology, scientists reported greater engagement between managers and evolutionary biologists as most important to achieve the necessary change. Nevertheless, the challenges appear to be multifaceted, and several are outside the control of managers, suggesting solutions will need to be multidimensional.

Recommendations for improved practice often centre on the active management of genetic diversity to maintain resilient populations (Table 1; Sgrò et al., 2011;Weeks et al., 2011). While the goal of conserving genetic diversity is widely recognized within international conservation policies, such as the Convention on Biological Diversity, little, if any, specific direction is provided about how management practices need to change (Cook & Sgrò, 2017). Conservation policies within individual countries appear to do little more than mention genetic diversity, while more specific evolutionary concepts (evolutionary process and principles), such as inbreeding and outbreeding depression or gene flow, are almost entirely absent (Cook & Sgrò, 2017).
The evidence of poor integration of evolutionary concepts into conservation policy documents (e.g., Cook & Sgrò, 2017;Lankau, Jorgensen, Harris, & Sih, 2011;Pierson et al., 2015) supports the widespread criticism from evolutionary biologists that conservation managers (policymakers and on-ground managers) are not changing their management practices in response to the available science. A key concern raised by evolutionary biologists is the general lack of consideration given by managers to how their management practices can act as a selection pressure on wild populations (Hendry et al., 2011;Smith et al., 2014). This is particularly relevant for invasive species management, where the evidence from agricultural science can inform risk assessments for management practice that might promote the evolution of resistance to the toxins used to control pest species (Neve, Busi, Renton, & Vila-Aiub, 2014;Tabashnik, Brevault, & Carriere, 2013).
Despite continued commentary about the long-term implications of management actions that do not consider evolution Smith et al., 2014), the reasons for this gap remain largely speculative. Some of the many reasons proposed by evolutionary biologists to explain the widespread failure to integrate evolution into management practices include poor training of managers in evolutionary biology (e.g., Frankham, 2010) and a lack of support for conservation managers to enable them to change their management practices (e.g., Hoban et al., 2013;Smith et al., 2014).
Anecdotal reports describing misconceptions among managers about evolutionary theory, which might act as impediments to evolutionarily enlightened management, have been blamed on a lack of specific training for conservation managers. For example, some authors have reported that managers often view species as fixed entities that do not change (Ashley et al., 2003), or believe that evolution happens too slowly to be relevant to management practices (Kinnison, Hendry, & Stockwell, 2007;Smith et al., 2014). It has also been suggested that managers have risk adverse attitudes to changing their management practices, viewing the manipulation of evolutionary forces as potentially disrupting the integrity of natural processes (Hendry et al., 2011;Smith et al., 2014). Concerns have also been raised that the actions of many managers remain guided by largely outdated or misinterpreted ideas, for example, anxiety about the risk of outbreeding depression when mixing populations, despite growing evidence that the risks have been overstated (Frankham, 2015;Frankham et al., 2011). Similarly, the focus on the "local is best" paradigm, which places precedence on collecting seed from populations considered to be adapted to local conditions, may limit the genetic diversity of revegetated populations, compromising their long-term viability and success (Broadhurst et al., 2008;Byrne, Stone, & Millar, 2011).
However, despite concerns that conservation managers misunderstand evolutionary concepts, the evidence for this belief remains anecdotal; managers' knowledge of evolutionary biology has not been quantified.
Some authors have identified a lack of appropriate decisionsupport tools as a barrier to better integration of evolutionary biology into conservation management (Frankham, 2010;Hoban et al., 2013). Specifically, changes to management practice require the development of tools that enable managers to identify the practical changes needed for evolutionarily enlightened management.
However, criticisms have been made that evolutionary theory continues to be ignored within existing decision tools, such as population viability analysis (Ashley et al., 2003;Frankham, 2010) and species distribution models (Urban et al., 2016). In other cases, there is a failure to translate theory into practical management recommendations that managers could implement (Cook & Sgrò, 2017). For example, recommendations that managers should monitor genetic diversity are increasingly the norm (Cook & Sgrò, 2017), yet practical guidance about how this should be done and how to distinguish between neutral and adaptive diversity is largely lacking (Hoffmann et al., 2015).
It has also been suggested that the slow integration of evolutionary biology into conservation practice may stem from a general failure by evolutionary biologists to engage with conservation managers (Hendry et al., 2010;Hoban et al., 2013;Mace & Purvis, 2008).
The increasing attention in the literature given to the need for evolutionarily enlightened management, and the potential reasons for this slow uptake have helped raise awareness of this problem in conservation biology (Hendry, Gotanda, & Svensson, 2017).
However, moving beyond the current speculation requires a clear understanding of the scale of the problem, quantifying the degree to which different evolutionary concepts are being integrated into management practice. Likewise, these data need to be paired with evidence for the causes of poor uptake in order to identify the most efficient ways to achieve change. This evidence needs to come from the conservation management community, to gain a true appreciation of the challenges they face in making changes to management practices. Because this evidence is currently lacking, our objective was to fill this important gap by surveying conservation managers, along with conservation scientists involved in conducting management-relevant science, about their views on (i) the importance of a range of key evolutionary concepts to conservation management, (ii) the current level of integration of these concepts into management decisions and (iii) the barriers and opportunities they see to achieving the necessary change.
TA B L E 1 Areas of environmental management in which evolutionary principles and processes could be used to inform management actions  (Ellstrand & Elam, 1993). Genetic diversity must be actively managed within threatened populations, potentially by facilitating gene flow via maintaining connectivity between populations or translocating individuals (Weeks et al., 2011).
Selecting individuals for captive breeding should focus on maximizing genetic diversity To ensure the long-term viability of restoration efforts, seed should be sourced from multiple populations (Broadhurst et al., 2008), preferably spanning an environmental gradient, to maximize genetic diversity and adaptive potential in the face of environmental change  When managing invasive species, the goal is to minimize genetic diversity. Biosecurity measures can be used to avoid multiple introductions that increase the genetic diversity of invasive populations (Simberloff, 2009   Management activities function as a selection pressure on invasive populations, which could lead to adaptations that reduce management effectiveness (Neve et al., 2014). This is particularly a problem for the use of toxins (e.g., herbicides, pesticides). Rotating or stacking toxins can help to avoid strong directional selection that leads to rapid adaptation (Beckie & Reboud, 2009 Gene flow can also be negative if it works to disrupt local adaptation or facilitates adaptation of pest species The small, often fragmented nature of threatened populations means that gene flow is important to maintain or restore genetic diversity (Weeks et al., 2011). Where connectivity has been disrupted, managed gene flow through translations can act to restore genetic diversity and reverse the impacts of inbreeding depression (e.g., genetic rescue; Weeks et al., 2017) Sourcing seed from populations that occur along an environmental gradient can maximize the adaptive capacity and resilience of restoration effects (Prober et al., 2016). Maximizing connectivity along an environmental gradient can enable the movement of adaptive alleles (Jordan, Hoffmann, Dillon, & Prober, 2017) Restrict gene flow between populations to limit genetic diversity and adaptive potential (Hendry et al., 2011).
It is important to assess the risk that genetically modified organisms transfer herbicide resistance to invasive species (Andow & Zwahlen, 2006).
Use gene flow to disrupt the evolution of resistance by increasing gene flow from susceptible populations (Tabashnik et al., 2013) Ensure protected areas are well connected to facilitate gene flow among populations (Ridley & Alexander, 2016).
Situating protected areas to allow gene flow across environmental gradients can help facilitate adaptation to changing environments (Lankau et al., 2011) Gene flow can serve to increase the adaptive capacity of pathogens, but can also be used to disrupt the evolution of resistance by increasing gene flow from susceptible populations (Alphey, Coleman, Donnelly, & Alphey, 2007) Inbreeding depression Inbreeding depression, the decreased fitness of offspring due to increased levels of homozygosity leading to the expression of deleterious recessive alleles, leads to loss of genetic diversity and demographic suppression Small, isolated populations are prone to inbreeding depression, which can increase extinction risk (Hedrick & Kalinowski, 2000). Mating systems influence genetic diversity (e.g., lowest in asexual reproducing species) and therefore the adaptive capacity of invasive species (Frankham, 2010) Not currently being discussed in the literature Not currently being discussed in the literature (Continues) TA B L E 1 (Continued)

| MATERIAL S AND ME THODS
To explore the degree to which evolutionary biology has been integrated into conservation management, we developed an anonymous, online questionnaire (Appendix 1) focused on a set of key evolutionary concepts (Table 2). These concepts were selected to represent evolutionary processes and principles from evolutionary theory and population genetics that are considered to be highly relevant to conservation management decisions. The selection of the concepts was informed by the frequency of their mention in the literature and in consultation with experts in applied evolutionary biology (withheld for review).
We targeted two groups of respondents: 1. conservation managers in policy or on-ground management roles; and 2. applied ecologists whose research programmes focus on management-relevant science.
The conservation managers were protected area or natural resource managers drawn from across all jurisdictions in Australia (five states, two territories and the federal government). Senior managers within protected area management agencies (n = 8) and national resource management organizations (n = 56) were contacted and asked to distribute the link to an online questionnaire to all relevant staff members, including those in policy and on-ground management roles.
We also contacted scientists whose research addresses conservation management problems. These individuals were identified by searching staff profiles from universities and government research institutes across Australia (n = 23). Relevant individuals were contacted via email (n = 78) and invited to participate in the study. They were also asked to distribute the survey to relevant members of their research groups, following a snowball sampling approach (Patton, 2002).

| Questionnaire development
We asked respondents to determine how important each of the key concepts in Table 2 was for conservation management, and to what degree they believed each concept was being integrated into current management practices. Respondents were asked to score their responses according to a 5-point Likert scale (Table 3), with an option to indicate if they were unsure about any of the concepts.
Respondents were not provided with definitions of the concepts, in order to avoid any temptation to speculate about the importance or integration of concepts they were not familiar with. However, this would not preclude respondents from looking up the meaning of concepts they were unsure about. It may also mean that respondents had varying interpretation of the concepts, which may have influenced their responses.
We also asked respondents to describe their perception of the most important barriers that inhibit integration of evolutionary the- Life history strategy also impacts the relationship between census and effective population size (e.g., when individuals reach sexual maturity) Life history traits influence the ability of species to adapt to changing environmental conditions (Hendry et al., 2011), influencing their vulnerability to extinction (Gallagher, Hammerschlag, Cooke, Costa, & Irschick, 2015).
Life history information can inform estimates of viable population size (Lankau et al., 2011) (Simberloff, 2009) Protected areas can be used as refuges for exploited species to reduce the impacts of life history evolution driven by selective harvesting (Baskett, Levin, Gaines, & Dushoff, 2005) Understanding life history strategy can inform the timing of treatments because those that act after first reproduction will slow the evolution of resistance in pathogens (Hendry et al., 2011) TA B L E 1 (Continued) need to change within their organizations for evolutionary theory to be adopted more broadly. Respondents were asked to provide a range of demographic information, including their role, age, years of experience, gender, level of education and exposure to evolutionary theory during their formal education (Appendix 1). Scientists were also asked whether they spend time engaging with managers about effective management practices.
Before distributing the questionnaire, the tool was piloted with seven individuals who confirmed the face validity (i.e., whether the meaning of the questions is clear to respondents and results of the survey will provide a meaningful outcome; Wainer & Braun, 1988) of the survey tool.

| Data analyses
We coded responses (Table 3)

| RE SULTS
We received 150 responses to the questionnaire (107 managers; 43 scientists) drawn from across Australia. Managers were mostly in onground management (n = 58) or policy or strategy roles (n = 39) (10 did not indicate their role). These respondents were mostly males (57%) and ranged in experience from 6 months to 29 years (μ = 8.7). The scientists who responded were in research only (n = 18) or teaching and research roles (n = 21), and just over half of scientists (56%) indicated that they engaged directly with managers. Most scientists were female (56%), and their experience ranged from 1 to 43 years (μ = 9.3).
The level of education of managers varied from high school graduates to postgraduate degrees (n = 6 declined to answer), while scientists had a bachelor's degree or higher (n = 4 declined to answer; Figure S1). Most respondents in each group reported having taken at least some subjects relevant to evolutionary theory, although 34%

General concepts
Genetic diversity Genetic differences between individuals of the same species

Adaptation
The condition where the phenotype of individuals is well suited to the environmental conditions, such that the individuals have higher reproductive fitness.

Evolution
The process by which populations or species change over successive generations.

Specific concepts
Gene flow Movement of alleles between populations through mating between individuals from different populations

Inbreeding depression
Mating between closely related individuals that leads to a loss of genetic diversity and corresponding reduction in reproductive fitness

Outbreeding depression
Mating between genetically distinct individuals that introduces new alleles that disrupt local adaptation and lead to reduced reproductive fitness

Mating system
The way in which a population is structured in relation to sexual behaviour

Life history strategy
The way in which individuals invest in growth, reproduction and survivorship TA B L E 2 Key concepts relevant to integrating evolutionary theory into conservation practice of managers had not ( Figure S1). Ten managers and four scientists declined to answer.

| The importance of evolutionary concepts
There were significant differences in the level of importance assigned by respondents to the different evolutionary concepts (H 7 = 46.70; p < .001). While there was variation in which concepts managers and scientists considered most important to conservation management, this difference was only significant for the importance of life history strategy (U 2 = 588.0; z = −2.02; p = .044; Figure 1; Table S1). Among managers, there was no difference in the level of importance ascribed to the evolutionary concepts by those in strategic roles and those in on-ground roles (U 2 = 24676.00; z = −0.43; p = .668; Table S2).
Respondents with different levels of education had significantly different views on how important the different concepts are to conservation management (H 3 = 9.32; p = .025; Figure 2a), with greater importance ascribed by those with a diploma or bachelor's degree. Prior exposure to evolutionary theory during their education influenced how important respondents considered evolutionary concepts to be (H 3 = 10.71; p = .013; Figure 2b), with those who indicated they had only taken subjects relating to genetics being less likely to consider the concepts important. Further evidence that lower importance scores may relate to the level of understanding of these concepts comes from the correspondence between more respondents failing to score the importance of a concept (i.e., selecting unsure) and a lower importance score ascribed to the concept by other participants (Table S3).

| Integration of evolutionary concepts
There was a significant difference in the degree to which respondents considered the different evolutionary concepts were being integrated in conservation management (H 7 = 77.43; p < .001).
While there was variation in the views of managers and scientists about the degree to which different concepts were being integrated into conservation management, overall these differences were not significant (U 2 = 49409.0; z = −0.38; p = .705; Figure 3a; Table S4). Within managers, those in policy or strategy roles and those in on-ground roles had similar views of how well the evolutionary concepts were implemented within conservation management (U 2 = 23416.0; z = −1.56; p = .118; Table S5), while scientists who indicated they engaged directly with managers tended to show closer alignment to the views of managers ( Figure S2).
The level of education of respondents did not influence the degree to which they believed evolutionary concepts were being integrated (H 3 = 1.70; p = .638). However, those who had not taken subjects specifically related to evolutionary biology during their degrees considered the concepts to be better integrated into conservation management (H 3 = 16.38; p = .001; Figure 3b).

| Barriers to and opportunities for greater integration of evolutionary theory
The most commonly reported barrier to greater integration of evolutionary theory by managers and scientists was a lack of understanding of the concepts (Figure 4a). Respondents from both groups also commonly suggested that conservation management is not being prioritized within management agencies, with an increasing emphasis being given to visitor management (Figure 4a; Table 3).
Managers showed greater concern with a lack of resources to implement the necessary management actions, while scientists ranked a lack of communication between managers and researchers more highly (Figure 4a). Scientists were more concerned that a failure to demonstrate clear benefits of changed management practices was a barrier, while managers were concerned that a focus on short-term outcomes of management prevented greater integration of evolutionary theory (Figure 4; Table 4).

F I G U R E 1
The mean (±SE) importance score for different evolutionary concepts as reported by managers (black circles) and scientists (grey circles). Asterisk indicates significant difference between groups The opportunities identified for greater integration of evolutionary theory into conservation management were directly related to the key barriers, essentially being potential solutions (Figure 4b). Managers and scientists differed somewhat in which solutions they saw as highest priority to improve the uptake of evolutionary theory. Scientists were more focused on greater engagement with managers and the need to demonstrate the benefits of changed management practices in improving conservation outcomes (Figure 4b). Managers tended to be more focused on the need for more training in evolutionary biology, a greater emphasis on conservation management within their agencies and additional funding to facilitate the required changes ( Figure 4b). We reveal that most managers strongly support the view that evolutionary biology is important for conservation management (Figure 1), but demonstrate there is a long way to go to achieve routine integration of these concepts into management practices (Figure 3). Our data also clearly show that managers face a range of practical constraints in trying to achieve the required changes ( Figure 4). By identifying these barriers and opportunities, we provide important insights into how the scientific community can better support managers, but also the changes that must occur within management organizations, and society more generally, to achieve real change.

| Importance of evolutionary concepts and their integration into conservation management
Managers consider evolutionary biology to be highly relevant to their management practices (Figure 1), suggesting that the concerns raised by evolutionary biologists have been heard. In particular, the messages about the importance of general evolutionary concepts, such as genetic diversity (Hoban et al., 2013;Santamaria & Mendez, 2012;Sgrò et al., 2011;Weeks et al., 2011) and adaptation , appear to have reached managers. These general concepts have also received more attention than other concepts within the evolutionary applications literature (Cook & Sgrò, 2017). Interestingly, evolution as a concept was not reported to be as important as adaptation and genetic diversity, potentially supporting concerns that rapid evolution is not being perceived as relevant to management (Ashley et al., 2003;Kinnison et al., 2007;Smith et al., 2014). This pattern also seems to suggest a poor understanding of the links between genetic diversity, adaptive capacity and evolution, which has also been observed in conservation policy (Cook & Sgrò, 2017).
While some evolutionary concepts were considered more important to conservation management than others, we found evidence that these differences may have been driven by poorer understanding of the relevance of some concepts. Managers who had a greater exposure to evolutionary biology in their training gave greater emphasis to the importance of these concepts (Figure 2b), supporting calls for the value of increased training for managers (Frankham, 2010). We also saw qualitative evidence for a lack of understanding of some concepts because managers were more likely to abstain from rating the importance for concepts that were given lower importance across the rest of the sample (Table S3). This may suggest a reluctance by respondents to rate concepts with which they were less familiar. However, without asking respondents to define each of the concepts, it is not possible to truly judge their specific level of understanding, or the consistency of interpretations among respondents. Poor understanding of evolutionary biology has been repeatedly suggested as a barrier to evolutionarily enlightened management (Ashley et al., 2003;Frankham, 2010;Mace & Purvis, 2008). Our results suggest that increased training in evolutionary biology may increase the value managers place on these concepts, and possibly their potential to integrate them into management.  . However, the lower importance ascribed to outbreeding depression and the poorer application of the concept to management do not support this being a major concern for managers ( Figures 1 and 2). The finding that managers consider outbreeding depression to be the least important concept for management decisions may reflect a poor understanding of the concept.
We found that more than half of managers in the sample chose not to provide an importance score for outbreeding depression (Table S3), suggesting most managers were unfamiliar with this concept. The poor integration of outbreeding depression into management decisions suggests managers are also not aware of the risk assessment frameworks available to guide decisions about when to mix populations (Frankham, 2015;Weeks et al., 2011). These decision-support tools also appear to have little penetration into conservation policy (Cook & Sgrò, 2017), making recent work that demonstrates the benefits of gene pool mixing (Frankham, 2016;Weeks et al., 2017) all the more important.
Both managers and scientists consistently considered the integration of evolutionary concepts to be out of step with their importance to management, with concepts rarely being integrated into management decisions (Figure 2). This supports the view that the level of integration of evolutionary concepts is generally quite poor (Hoban et al., 2013;Mace & Purvis, 2008;Smith et al., 2014).
Both managers and scientists were in agreement that the greatest discrepancy between the importance and implementation of concepts occurs for outbreeding depression and evolution (Figures 1   and 2). Management agencies were considered to be doing better at integrating genetic diversity and life history strategies into management decisions (Figure 2), and there is certainly increasing guidance in the literature for how managers can take account of these concepts within revegetation (Breed, Stead, Ottewell, Gardner, & Lowe, 2013;Byrne et al., 2011;Sgrò et al., 2011) and threatened species management Weeks et al., 2011), which may be filtering through to managers. However, further work is needed to understand exactly how managers are applying these concepts to determine whether more effective long-term management outcomes are likely to be achieved.

| Changes required to facilitate greater integration of evolutionary theory into conservation management
While the message that evolutionary theory can benefit effective management appears to have been received by conservation managers, we found that both managers and scientists perceive a wide range of barriers that will need to be overcome to achieve greater integration ( Figure 4a). Some of these barriers are within the control of scientists and managers to address, such as a lack of understanding of the relevant concepts (Ashley et al., 2003;Frankham, 2010) and poor engagement between scientists and managers (Hoban et al., 2013;Mace & Purvis, 2008). However, some barriers are beyond the control of both groups, such as insufficient resources to make changes to existing management practices (Figure 4a). This is not surprising, given the widely acknowledged shortfall in funding for conservation management (Waldron et al., 2013) is frequently cited as a barrier to implementing more effective management (Addison, Cook, & Bie, 2016;Leverington, Costa, Pavese, Lisle, & Hockings, 2010).
Other barriers to greater integration of evolutionary biology that are beyond the control of managers and scientists include a shift in the emphasis of management roles away from biodiversity towards visitor management (Figure 4). The concern that conservation management is being given lower priority by politicians (Addison, Flander, & Cook, 2017;Addison et al., 2016) and the general community (McCallum & Bury, 2013) seems to be translating to more prominence for tourism and economic objectives for natural areas, particularly within protected areas (Balmford et al., 2009;Eagles, 2002). Managers reported that the ability to achieve best practice in management is being impeded by this lack of support, an idea that scientists appear to corroborate (Figure 4a). This reduced emphasis on conservation management may also be reflected in some managers' concerns that conservation management is overly focused on short-term outcomes (Figure 4a), at the expense of the longer term benefits that can be achieved Smith et al., 2014).

F I G U R E 4 The (a) barriers to and (b) opportunities for greater integration of evolutionary theory into conservation management as reported by scientists (black bars) and managers (grey bars)
While a focus on longer term outcomes is certainly important, this must be balanced with the understanding that evolution can occur at management-relevant timescales (Kinnison et al., 2007), and negative outcomes can occur rapidly (Smith et al., 2014). Greater engagement between evolutionary biologists and conservation managers may need to focus on how to achieve a balance between managing immediate threats and achieving the best long-term outcomes.
While we found broad agreement between managers and scientists about the barriers preventing greater integration of evolutionary theory in conservation, the two groups placed slightly different emphasis on the solutions needed to achieve change ( Figure 4b). Managers focused on the need for more training to better understand evolutionary biology, while scientists emphasized the need to improve the science-policy interface to better support managers to make good decisions (Figure 4b). These two solutions could be seen as two sides of the same coin and have certainly been suggested as a key plank in improving the integration of evolutionary theory (Frankham, 2010;Mace & Purvis, 2008), and scientific evidence more generally (Cook, Mascia, Schwartz, Possingham, & Fuller, 2013), into conservation decisions. Evolutionary biologists have arguably been less proactive than ecologists when it comes to engaging with conservation managers about improving management practices (Hendry et al., 2010;Hoban et al., 2013;Smith & Bernatchez, 2008). Therefore, by playing a greater role in the science-policy interface, evolutionary biologists could help to significantly improve management practices and outcomes.
Both groups also suggested that there needs to be greater emphasis on research that demonstrates how better conservation outcomes can be achieved by changing management practices ( Figure 4b). Many of the scientific advances in evolutionary theory have come through research on model systems, which allow greater traction when testing the impact of evolutionary processes (Frankham, 2015). However, it may be hard for managers to translate this evidence into their management contexts, or even to see the relevance of these studies to management (Cook & Sgrò, 2017;Frankham, 2010;Hoban et al., 2013). The evolutionary biology community appear to be heeding calls to demonstrate how theory can be translated into more effective management practices (Frankham, 2010;Hoban et al., 2013;Lankau et al., 2011), with more experimental studies being undertaken that demonstrate the benefits of evolutionarily enlightened management within conservation contexts (e.g., Frankham, 2015;Hedrick & Fredrickson, 2010

Barriers to better integration
Lack of education A lack of training and basic understanding of the relevant concepts, their importance and/or how they should be applied to management

Conservation not prioritized
A lack of support for conservation from governments and the broader community, and a shift in emphasis to towards visitor management rather than conservation management Lack of resources Declines in the resources for conservation management mean funds are inadequate for on-ground management. There are competing priorities for resources, and funding is uncertain and short-term. Too few managers.
Lack of communication with scientists A lack of engagement by scientists means there is poor translation of primary research into management programmes Short-term not long-term The focus of management is on dealing with immediate problems, not long-term outcomes. There is poor understanding of how concepts can be applied to short management time frames. Evolution and management occur over different time horizons.

Benefit not demonstrated
There is little evidence for the application of relevant concepts to conservation management. There are no case studies that show chances are beneficial.

Research and monitoring
Not enough research or funding for necessary research. Research is generally academic and not focused on conservation management. Unclear what to monitor.

Legislation, policy and guidelines
Legislation is interpreted too narrowly and provides impediments (e.g., managing across borders) to including evolutionary processes. There is no mandate within policy to change management practices. Managers do not know how to integrate these ideas into their practices.
Other Conflict between evolutionary theory and religious beliefs. Need to engage other landholders and managers for integrated management.

Mismatch in spatial scale
Management occurs at small scales (e.g., small, isolated areas), but evolutionary process often need to be managed at a landscape scale.

None
There is currently nothing preventing greater integration.
& Whitlock, 2013), for example. Drawing managers' attention to the findings of these studies could help demonstrate the practical application of what could be perceived to be more abstract theory.

Greater interaction between managers and evolutionary biologists
would support greater knowledge transfer and potentially provide a platform for research demonstrating the outcomes of alternative management practices.

| CON CLUS IONS
Improving the uptake of evolutionary biology within conservation management requires a detailed understanding of the current level of integration into management practice. However, this knowledge must be paired with the views of conservation managers on the value they place on evolutionary theory, their level of understanding of relevant concepts and what needs to change to facilitate greater uptake. We reveal that managers do generally understand the importance of evolutionary theory to conservation management, although managers hinted through their survey responses that there are some concepts (e.g., outbreeding depression) that they do not fully understand. Further investigation of the depth of understanding of core evolutionary concepts would help to better assess the extent to which limited understanding acts as a barrier to evolutionarily enlightened conservation. We also show that managers face many barriers to improving the currently poor level of integration of evolutionary concepts into their management practices. There are practical things that can be done to improve the adoption of evolutionary theory, such as increasing the exposure of managers to evolutionary theory during their training, and greater engagement with managers by evolutionary biologists. However, there are also issues beyond the control of managers, such as the level of public and political support for conservation management and the resources available to improve management practices, which require more systemic changes.
Likewise, when making management decisions, managers must also consider scientific evidence relating to a range of other disciplines (e.g., ecology, hydrology, geology), in addition to influential social, political and economic factors, which may limit their ability to achieve evolutionarily enlightened management. Therefore, it is likely that a multidimensional approach is needed to achieve the necessary change.

ACK N OWLED G EM ENTS
This research was conducted in accordance with the requirements of the Monash University Human Ethics Committee (MUHRECapproved project: CF15/774-2015000348). The authors thank D.
Coates for useful discussion in the planning stage of this work and R.
Valkan for assistance with data collation. The authors would also like to thank all the managers and scientists who generously gave their time to participate in this study. Thanks also go to three anonymous reviewers for comments on this manuscript.

CO N FLI C T O F I NTE R E S T
None Declared.

DATA A R C H I V I N G S TAT E M E N T
We have been granted an exemption from data archiving some or all of the data associated with this paper.