A wolf-shaped hole in Britain
Reinstatement of wolves to Britain is an itch I have to scratch. It has to be a strategic but, more importantly, an ecologically rational and valid approach. Even when the discourse is favourable to reinstatement, it rarely gets above the superficial or simplistic because it is never taken seriously enough
I don’t often get to see the outcome when interviewed by students for their dissertations. I think I’m chosen by them (or their advisors) as the outlier, the immoderate viewpoint that makes the others interviewed seem like the voice of reason. It was a surprise then, when looking back through journal articles recently, that the answers I’d given in an interview for a Master’s thesis four years before had popped up instead in a paper from 2018 (1). My memory of the interview is very sketchy, but I am pleased to see that my considerations on the ecological illiteracy of herbivore-driven landscapes were reported, highlighting the fallacy of creating a trophic imbalance in the absence of predators.
Using bison as a tool for species gardening
A more recent interview in July was with a Master’s student in International Journalism at City, University of London (2). His final project was to be a video on the topic of rewilding in the UK. He said he was interested in whether apex predators should be brought back to the UK, and what their impact could be. However, his first question was about whether bison should be reintroduced. I understood why he asked that, as there had been a big fanfare in a newspaper article a few weeks before about the release of bison into a fenced enclosure in Blean Woods in Kent where they would “transform a dense commercial pine forest into a vibrant natural woodland” (3). Smashing up woodland is how I categorised this project when it first announced a couple of years ago that it had received lottery funding (4). I wrote that it was no different from conservation grazing, as it would be using these bison as a tool for species gardening. It would be just like the use of cattle in mainstream nature conservation in Britain, but with the difference that the perpetrators of this project had been seduced by seeing the greater destructive ability of bison compared to cattle in fenced-off enclosures in the Netherlands like the Kraansvlak. Worse though, was the impression given by the project that these bison had an eventual place freely living in British landscapes when there is no ecological justification for their being here.
I explained this to the student, adding that the population density of the bison, even in the largest of the fenced off area in the Blean, would be six times greater than is found in free living populations across Europe. This over population characterises enclosed bison projects in Europe (5). It speaks to the ecological illiteracy of ignoring the life history of free-living bison and their need for large, unconfined spaces, as it does to any number of projects predicated on enfenced herbivores that ignore the life history strategies of wild species, such as deer, or make no attempt at extrapolation for the impact of domesticated species like horses and cattle. It is a recipe for ecological meltdown (6,7).
At the end of the interview, I implored this student to ensure accuracy in his future career in journalism, having in mind that the newspaper article about this bison project had described them as “wild” when they had come from incarceration in wildlife parks in Scotland and Ireland, and that they were “to become the first wild bison to roam in Britain for thousands of years” when their roaming would be curtailed by fencing (3). The latter claim is just such hyperbolic nonsense anyway, as there were roughly 250 bison already present in Britain before this project, mostly farmed for meat, some in wildlife parks (see, for instance, the Melton Borough Council area that has 105, Wiltshire Council area has 60, and Durham County Council area has 40 in (8)). These are all behind fences, as bison come under the Dangerous Wild Animals Act 1976, so that anyone wishing to keep them has to obtain a licence from their local authority and adhere to the requirement that they be “held in accommodation which secures that the animal will not escape” (9)
How wolves use the land
His question then on wolf reintroduction allowed me to rehearse my conviction that the westward voluntary dispersion of wolves in continental Europe was one of the greatest ecological experiments in trophic reinstatement (10); that there was a wolf-shaped whole in Britain (11, 12); that it was the English Channel that prevented voluntary dispersion here, making reinstatement a contestable proposition (13,14); but that reinstating the wolf would drive a process of determining the ecological flow that would exist for the wolf – how it would use the land (15) This would be a hugely revealing process, as well as a great opportunity to engage people in ecological understanding. It would break through the power and ownership structures, its impact forcing land users to face up to human-wildlife conflicts in constructive ways, as the strict protection of the wolf under the Bern Convention would apply here. It would be a catalyst for reframing our outlook and policies on wild nature, and thus had the potential to change the paradigm from nature conservation to nature protection – protecting nature from us.
I explained that a first step would be an ecological feasibility study to identify the likely locations where wolves could disperse to and secure their living. This is the approach that Denmark and the Netherlands had employed in advance of wolves voluntarily dispersing into their countries (16) although these studies did not address possible routes for wolf dispersal as they recolonised those countries, only predicting where they may end up staying (10). The student, however, brought up a proposal to confine wolves behind fencing in a reserve as a means to enlist their predatory behaviour in reducing the number of Red deer. While I didn’t ask, I thought I knew the paper where this proposal may have come from (17) having written about the many failings it contained, not least in distracting from the real task in Scotland, its proposed location, as it may do elsewhere in Britain, of debating the return of the wolf to free-living rather than captive enslavement (19). It was a non-starter anyway, which you think the authors should have been aware of, in that these fenced reserves would be regarded as open air zoos and thus would be subject to the Standards of Modern Zoo Practice that do not allow predator and prey to be in the same enclosure.
I gave the student the chilling vision that could follow this proposal, of the reaction people would have when they saw a wolf pack chase and manoeuvre a Red deer into the fence so that its further escape is blocked, it becomes trapped and its fate inescapable. Moreover, I explained that I don’t tend to justify reinstatements on the basis of ecosystem service or function (use value) because I take the view that it’s not about gain but absence when considering the intrinsic value of lost wild nature ((20) and see (21,22)) that absence or incompleteness having consequences for the trophic ecology of our landscapes. I don’t want to see wolves just as a tool to manage our native wild deer. In concluding, the student gave his own thoughts; that it was unlikely that wolf reinstatement would take place given what he thought were the potential conflicts with farming.
Reinstatement of wolves to Britain is an itch I have to scratch. It has to be a strategic but, more importantly, an ecologically rational and valid approach, unlike the egocentrism of the personal agendas that are driving the bison project (see above). I can’t stand the thought of another abortive attempt like lynx reinstatement where the process was wrested away from the ecologically rational and valid through a vainglorious agenda that so carelessly and violently sloughed off the remarkably invested and self-motivated team working with the project (23). As it is, there are elitist, self-regarding forces circling in Britain at the moment to claim and degrade wolf reinstatement. Thus, in typical grandstanding form, animal trader Derek Gow has proclaimed that he is thinking of organising a wolf conference next year while openly soliciting funds that Ben Goldsmith, his perennial bankroller, seems inclined to supply, as does one of the egocentrics behind the bison project (24). You have to look at animal trader Gow’s track record to understand his motive here, whether its water voles, beaver, wildcat, tree frogs etc. he is always seeking to make money out of wild nature (25-28). Hence also his safari park farm with its farming subsidy and agri-environment payments (29). In similar view to not wanting Gow anywhere near taking responsibility for a national strategy for reinstatement of wildcat (25) I don’t want him anywhere near the reinstatement of wolves. Contracting nature recovery out to the private sector is not our only or even remotely best option.
A predominant hypocrisy
I believe that predation above all else – and at every level of trophic occupancy up to the highest – is the most important component of our wild nature, even though our wild nature is rarely seen in relation to the natural process of predation in trophic ecology (30). Yet predators, whether fox and badger today - or wolf in the past – are spitefully persecuted in an essentially lawless countryside. There is a predominant hypocrisy in all this, but it will take some explaining.
Man’s best friend is a common phrase used to describe the dogs we keep as companions. It’s a sentiment first attributed to Frederick the Second, King of Prussia, when the Italian greyhound that accompanied him on one of his military campaigns during the 1740s did not bark and give him away when he had to hide under a bridge to avoid the advancing enemy (31). Those best friends are also utilised as working dogs in the countryside, such as sheep dogs and gun dogs, by the very people whose forebear’s intolerance of the wolf led to its extirpation.
James Harting’s book from 1880 on extinct British animals meticulously collects together records and sources of information on the presence and eventual elimination of the wolf from Britain, noting that osteological remains (bones) indicated “there was perhaps scarcely a county in England or Wales in which, at one time or another, Wolves did not abound, while in Scotland and Ireland they must have been even more numerous” (see from pg. 115 (32)). Harting’s history of oppression of wolves provides a more comprehensive account of the repeated attempts over centuries by the power and ownership structures to enforce elimination than I found when writing about this previously (13,15). Paradoxically, there are many accounts in Harting’s book of the hunting of wolves with dogs, often for pleasure by the ennobled, or for the determined and systematic destruction of wolves. Wolf hounds used for hunting were considered very valuable and were “highly thought of by those who received them as presents”. Today, large dogs like the patou (Pyrenean Mountain Dog) and other mountain dogs are used to guard sheep against wolf attacks (14,33).
Where do these hunting dogs, livestock guarding dogs, working dogs and man’s best friend come from? From domestication of the wolf, and there is the hypocrisy. The domestication of the wolf began even before the domestication of food or livestock species, such as sheep and goats (34). It started sometime between 40,000 and 15,000 years ago and possibly independently in more than one region of Eurasia (35). One recent paper favours Siberia as being the origin ~23,000 years ago, noting that domestic dogs accompanied the dispersal of early human groups (36). The wolf was the earliest animal domesticated, but the specific wolf population from which dogs were derived appears to be extinct. Two theories have been proposed for how that domestication occurred: commensalism where wolves essentially domesticated themselves by invading early human settlements, scavenging animal remains and other edible waste discarded by hunter-gatherers, so that in time a tolerance was developed by humans and wolves that turned into a relationship; or cross-species adoption where wolf cubs were taken by early humans and socialised through nurture. The scavenging hypothesis was considered unlikely in a recent review, based on safety concerns for wolf predatory behaviour when in close proximity, and what is known about recent and ancient hunter-gatherer societies in avoiding waste and disposing of unused remains (34). Instead, it was considered that wolf domestication was probably predicated on the establishment of cooperative social relations between humans and wolves based on the early socialization of wolf cubs.
What do you see when you look at a dog?
Is it a wolf? Could it be a wolf? A recent study compared the perceptions regarding the reintroduction of river otters and oriental storks in Japan (37). There was a more positive attitude to reintroduction of storks because this was the species that was more familiar to respondents. The authors noted that this was consistent with the theory of mere-exposure effect - a psychological phenomenon by which people tend to develop a preference for things merely because they have repeated exposure to them (38). If you were to see a dog as a wolf, albeit with pretty much of its wild instincts removed, then sentiment towards reinstatement of its wild forbear should benefit from the mere-exposure effect. That it doesn’t comes down to control, the clearly subordinate character of dogs compared to the free-living uncontrollable wildness of the wolf.
I think this lack of control is shown by a study that explored peoples’ preferences for the conservation firstly of large carnivores as a whole, and then distinguishing preference between wild canids and wild felids (39). Each participant was asked to record how familiar they were with the given species (familiarity) and how much they liked the species (affinity). No canids appeared in the top ten for preference among large carnivores, the top two places occupied by the White Tiger and Giant panda, animals adjudged elsewhere to be amongst the twenty most charismatic species (40). When just the preference for felids and canid was tested, only the Artic fox appeared in the top ten – at tenth place (39). The authors noted that familiarity and preference were not always linked. Thus preference scores increased with affinity for species which were liked, but declined with familiarity for species which were not liked. The authors concluded that this decline reflected negative experiences or cultural histories – it was a familiarity with consistently negative depictions. As the authors say, to know them is not always to love them, but I don’t understand why felids appear to people to be less dangerous than canids.
Changing perceptions of the wolf in a vacuum of understanding of its ecological flow in Britain – as was the case when only the social feasibility of reinstating lynx to Scotland was recently assessed (41) – is a futile pursuit. I have written before of the spatial and temporal avoidance strategies that wolves employ in human dominated landscapes to reduce potential conflict with humans (10). It seemed logical to me that we should determine that ecological flow - how wolves would use the landscapes of Britain, where they would choose to reside, and what routes they would take to disperse – so that we could work out what measures must be taken to ensure the separation that undoubtedly does take place between wolves and humans in modified landscapes. It would go towards lessening the negative experiences, both physically and psychologically. We have the Danish and Dutch mapping studies to learn from (see above) and there is a more recent study that maps the potential areas for expanding the distribution of wolves in Sweden based on prey availability, and the impact it would have on prey species (42). This is against a backdrop of acute cultural intolerance that saw Sweden divided into three predator management areas in 2009 along with the resumption of licensed wolf hunting (43). The hunting was subsequently deemed a form of population control that is an infringement of the strict protection of wolves under the EU Habitats Directive. The planned cull to halve the Swedish wolf population announced by its Parliament in May provoked the ire of scientists (44). Nevertheless, the authors of the mapping study hope that their data on forecasting the potential impact of a dispersing large carnivore can contribute to more constructive discussions rather than heated debates about conservation conflicts (42).
No indication about potential wolf movement
There is guidance in the literature on the different ways to explore species reintroduction through spatial modelling (i.e (45,46)) that will prove useful in that it may need a number of iterations using different approaches and varying attributes (vegetation cover, prey availability, connectivity, level of human disturbance) before we can be satisfied. As it is, a rule-based habitat suitability model for wolves on the Scottish mainland came out in March, a pre-print of which is available but which has yet to undergo peer review (47). The study took land cover, prey density, low road and human density as the most important factors in determining wolf habitat. Use of land cover was problematic because of the difficulty of assigning suitability values to some of the vegetation cover types, and the dataset for prey density was incomplete (including missing out various species in Scotland) so a series of models were used where the values of these were varied. The Highlands and Grampian mountains emerged as a concentration of most suitable areas and, depending on which model, a contiguous area of between 10,139km2 and 18,857km2 was shown to have between 80 to 100% suitability. I estimate that top end area to be just under a third of mainland Scotland (see Fig 4 in (47)). The authors claimed this could be sufficient to support between 50 and 94 packs of four wolves, if the average pack range size was taken to be 200km2.
The study attracted some press coverage in Scotland: one newspaper was neutral (48) another commented that it “may send a chill down the spine of many who live and work rurally”; that rural sceptics were unlikely to be persuaded by another set of figures and graphs; and that NatureScot, the statutory nature agency, had said that there were no plans to consider the wolf’s return (49). The latter newspaper also ran a poll on whether wolves should be reinstated in Scotland: out of 611 votes, only 30% said Yes, 64% said No, and 7% they should only be reinstated behind fences (50). I don’t take these kinds of polls seriously, as respondents are self-selecting. I’m more concerned with what appears to be the grossness of the outcome in the study in terms of the mapped area of suitability.
It struck me as counter intuitive that there were only two large blocks with little deviation within them except for inland water bodies ((see Fig 4 in (47)) and that there was a considerable difference between the upper and lower estimate in terms of overall area of suitability (see above). Forest was assigned the highest value for land cover suitability score in the study (see Table 2 in (47)). I would thus have expected to see something turn up in other areas of the mapping away from those two large blocks, as could be predicted from the distribution of woodland in Scotland. Have a look at the map of woodland by size class used in a study for the potential of reintroduction of Eurasian lynx to Britain, and which shows three large areas of woodland cover to the south of Grampian and Highlands: around Oban and the peninsula below; in the area of Dumfries and Galloway and up into Strathclyde; and eastern Dumfries and Galloway across into the Borders (see Fig 2 in (51)).
The authors claim there was a high level of covariance between three of the variables, prey density, road density and human density, in that they predicted suitable area in the same regions. However, if you compare the overall map that combines all four variables, it almost perfectly mimics the map of suitability in relation to low road density, even to the point of replicating a few small spots and other areas away to the south of the large blocks (compare Fig 3f with Fig 4d in (47)). This would suggest that the threshold of low road density for suitability has the effect of outweighing any influence from the other three variables. I have my colleague Steve’s voice in my ear telling me about the difficulties from uncertainty in the validity of chosen weighting schemes in multi-criteria evaluation (52). It would suggest that this variable of road density needs revisiting. The authors rely on data from N. America and Eastern Europe (47). However, data from countries in Western Europe, including newly colonised countries such as Denmark, Belgium, and the Netherlands, would probably provide more real world appraisal of the influence of road density in places like Scotland and the rest of Britain. No distinction was made between major roads and smaller ones, which the Danish study (see above) saw as a factor, indicating that smaller roads will likely have lower rates of traffic, especially at night, and will thus pose a lower threat to the dispersal of wolves (see temporal avoidance strategies above).
The mapping of that study gave me no indication about potential wolf movement. It’s not just a disappointment at how little complexity it reveals, but a doubt that this study has got past the stage of a too simplistic exploration. Its failure to provide much insight - other than a quandary over suitability of some landscape covers; a known deficiency in data for prey; and road density appearing to be the overriding factor irrespective of the other variables - has little value in our understanding of ecological flow for wolves, or in overcoming negative assumptions. We need better than this. I have seen an unpublished landscape connectivity analysis of Britain that focuses on identifying areas likely to facilitate ecological flow—particularly movement, dispersal, gene flow, and distributional range shifts for terrestrial plants and animals. The mapping was a resistance-surface-based connectivity approach (15) that combined a spatial layer of land cover classified in terms of naturalness on a relative scale (eg. woodland versus urban) with a layer on human influence based on distance to roads and built density. It was a species agnostic modelling of landscape connectivity, because the weightings given to the various land cover types were not dependent on the behavioural responses of a specific species to landscape structure. I ground truthed this connectivity analysis against an area I know well, using street and aerial maps, and found remarkable congruence (53). With the addition of prey availability, and perhaps some adjustment of land cover weightings, this resistance surface approach could be used for identifying sufficiently large areas for colonisation by wolves as well as their ecological flow.
A process of public engagement
I’ve been watching progress since the success of the ballot in Colorado on requiring the state wildlife agency, Colorado Parks and Wildlife, to reinstate wolves on designated Federal land in the west of the state (10, 54). A process of public engagement is being carried out by Colorado Parks and Wildlife and Keystone Policy Center, an independent facilitator, on behalf of the Colorado Parks and Wildlife Commission (55). The outcome feeds into the development of a Colorado Wolf Restoration and Management Plan that will see the restoration of wolves begin by the end of 2023. As well as a summer of public engagement last year that reached more than 3,400 participants through 47 meetings, along with an online comment form (56,57) a Technical Working Group and a Stakeholder Advisory Group were convened as advisory bodies to the state agency (58). These two bodies started meeting regularly from June last year, and continued up until last month when they submitted final reports on a range of issues that need to be part of the Restoration and Management Plan, such as outreach and education, conflict minimisation, restoration logistics (59) impact assessment, and livestock compensation. Colorado Parks and Wildlife staff will now develop a draft plan by November, which will be presented to the Commission and released to the broader public during a series of meetings in January and February next year (60).
It’s been an education to follow and learn from this, not least in accepting that consensus was not always possible in the advisory bodies; and in the dynamic of perceptions, the public engagement report noting that holders of diametrically opposed viewpoints frequently felt that “the other side was perpetuating a myth or misperception of the likely ecological impacts of wolves in Colorado” (61) – hence the sensible approach of setting up the two advisory bodies that could resolve these issues. It’s been known for some time that western Colorado was potentially suitable habitat for recolonising with wolves ((see Fig 2 in (62)). The ballot in Colorado provided a remarkable opportunity to map ecological and socially suitable habitat for reinstatement of the wolf. A proxy measure of tolerance for wolves was based on voting patterns across the state (votes at precinct level) and then combined with livestock distribution and land ownership to map socially acceptable space (63). This was then set against ecological suitability based on prey availability, wolf use space (road and housing density) and slope combined with snow cover as a proxy for prey vulnerability/hunting success. The overall mapping identifies a spine (~32.5m acres) running up through western Colorado that is both ecologically and socially acceptable habitat (see Fig 3, 4A in (63)). As importantly, the mapping identified areas (8.3m acres) to the west of that spine that were likely conflict hotspots (see Fig 4B in (63)). The authors recommend that these potential conflict hotspots would benefit from proactive management and outreach efforts to reduce conflict in advance and during the wolf restoration process.
There are about 20 wolves at non-zoo locations in Britain (8) all of them living out their lives behind fences while being gawped at, such as during a safari drive-through at Longleat (64) on a tour at Wolf Watch (65) or feeding the wolves at the Paradise Wildlife Park (66). Such an indignity when they should be living free in Britain. There is much to pull together if we are to chart a robust course for wolf reinstatement.
Mark Fisher 6 September 2022
(1) Root-Bernstein, M., Gooden, J., & Boyes, A. (2018). Rewilding in practice: projects and policy. Geoforum, 97, 292-304
(2) International Journalism MA, City, University of London
(3) The rangers expect the bison to breed, with females producing one calf a year, and the Wilder Blean site is licensed for up to 10 animals, Damian Carrington and Nicola Davis, Guardian 18 July 2022
(4) Where have all the woodland flowers gone? Self-willed land August 2020
(5) Movement ecology and rewilding, Self-willed land September 2019
(6) Patterns and disconnections in nature, Self-willed land August 2016
(7) Moving past process to outcome – the manifestation of wild land, Self-willed land September 2017
(8) Dangerous wild animal map, Born Free Foundation
(9) Dangerous Wild Animals Act 1976
(10) The separation between wolves and humans in modified landscapes, Self-willed land March 2020
(11) Fisher, M. (2009) Ecological incompleteness and our missing top predators: learning the lessons from abroad Wilder Horizons 1(1): 14-16
(12) Requiem for rewilding, Self-willed land July 2021
(13) Large carnivores as the focal species for reinstatement of natural processes in Britain, Self-willed land November 2014
(14) Cry wolf - the return of Britain's top predator, Self-willed land February 2015
(15) Ecological flow, nature protection, and the wolf, Self-willed land July 2020
(16) Ecological consequence of predator removal, Self-willed land July 2014
(17) Bull, J. W., Ejrnæs, R., Macdonald, D. W., Svenning, J. C., & Sandom, C. J. (2018). Fences can support restoration of human-dominated ecosystems when rewilding with large predators. Restoration Ecology 27(1) 198-209
(19) Using functional traits - walking rewilding and wolves straight into the criticism of Goldilocks Standards, Self-willed land June 2018
(20) The greatest challenge for living with wolves rests within the human mind, Self-willed land November 2017
(21) Vucetich, J. A., Bruskotter, J. T., & Nelson, M. P. (2015). Evaluating whether nature's intrinsic value is an axiom of or anathema to conservation. Conservation Biology, 29(2), 321-332.
(22) Batavia, C., & Nelson, M. P. (2017). For goodness sake! What is intrinsic value and why should we care? Biological Conservation, 209: 366-376
(23) A false start for lynx reinstatement, Self-willed land December 2018
(24) Derek Gow (@gow_derek) Twitter 14 August 2022
(25) What future Wildcat in Britain? Self-willed land August 2019
(26) Captive wildcats sent to 'training camp' to equip them for life after release, Helena Horton, Daily Telegraph 13 March 2021
(27) Dawn chorus of the tree frog may return to the UK under major rewilding plan, Helena Horton, Daily Telegraph 1 February 2021
(28) Farmers in England to be allowed to use ‘lethal force’ on beavers, Helena Horton and Gemma McSherry, Guardian 3 September 2022
(29) Faking the wild – safari park rewilding, Self-willed land May 2020
(30) The most important component to preserve is predation itself, Self-willed land July 2019
(31) Jean-Charles Laveaux (1789) The Life of Frederick the Second, King of Prussia: To which are Added Observations, Authentic Documents, and a Variety of Anecdotes, Volume 2. J. Derbett – Prussia. Pg. 521
(32) Harting, J. E. (1880) British animals extinct within historic times: with some account of British wild white cattle; with illustrations by J. Wolf, C. Whymper, R.W. Sherwin, and others. London: Trübner and Company, Ludgate Hil.
(33) Carnivore Damage Prevention News, Issue 15 SUMMER 2017
(34) Serpell, J. A. (2021). Commensalism or cross-species adoption? A critical review of theories of wolf domestication. Frontiers in Veterinary Science, 8: 662370.
(35) Frantz, L.A., Mullin, V.E., Pionnier-Capitan, M., Lebrasseur, O., Ollivier, M., Perri, A., Linderholm, A., Mattiangeli, V., Teasdale, M.D., Dimopoulos, E.A. and Tresset, A., (2016) Genomic and archaeological evidence suggest a dual origin of domestic dogs. Science, 352(6290): 1228-1231
(36) Perri, A. R., Feuerborn, T. R., Frantz, L. A., Larson, G., Malhi, R. S., Meltzer, D. J., & Witt, K. E. (2021). Dog domestication and the dual dispersal of people and dogs into the Americas. Proceedings of the National Academy of Sciences, 118(6): e2010083118
(37) Sakurai, R., Stedman, R.C., Tsunoda, H., Enari, H. & Uehara, T. (2022) Comparison of perceptions regarding the reintroduction of river otters and oriental storks in Japan, Cogent Social Sciences, 8(1): 2115656
(38) Zajonc, R. B. (2001). Mere exposure: A gateway to the subliminal. Current directions in psychological science, 10(6): 224-228
(39) Macdonald, D. W., Johnson, P. J., Burnham, D., Dickman, A., Hinks, A., Sillero-Zubiri, C., & Macdonald, E. A. (2022). Understanding Nuanced Preferences For Carnivore Conservation: To Know Them Is Not Always To Love Them. Global Ecology and Conservation, 37: e02150
(40) Albert C., Luque G.M., Courchamp F. (2018) The twenty most charismatic species. PLoS ONE
(41) Assessing the social feasibility of returning Eurasian lynx to Scotland, Lynx to Scotland partnership
(42) Rodríguez-Recio, M., Wikenros, C., Zimmermann, B., & Sand, H. (2022). Rewilding by Wolf Recolonisation, Consequences for Ungulate Populations and Game Hunting. Biology, 11(2), 317
(43) When nature dies - the impact of the human species, Self-willed land July 2015
(44) Laikre, L. and others (2022) Planned cull endangers Swedish wolf population. Science 377(6602): 162
(45) Hunter-Ayad, J., Ohlemuller, R., Recio, M. R., & Seddon, P. J. (2020). Reintroduction modelling: A guide to choosing and combining models for species reintroductions. Journal of Applied Ecology.57:1233–1243
(46) Zurell, D., König, C., Malchow, A. K., Kapitza, S., Bocedi, G., Travis, J., & Fandos, G. (2022). Spatially explicit models for decision‐making in animal conservation and restoration. Ecography, 2022(4): e05787
(47) Gwynn, V., & Symeonakis, E. (2022). Rule-based habitat suitability modelling for the reintroduction of the grey wolf (Canis lupus) in Scotland. bioRxiv.
(48) Highlands and Grampian mountains are best places to bring back wolves to Scotland, study finds, Ilona Amos, The Scotsman 13th March 2022
(49) Rewilding: Wolves could save Scotland's environment, Vicky Allan, The Herald 15 March 2022
(50) Poll: Should wolves be reintroduced to Scotland? Vicky Allan, The Herald 15th March 2022
(51) Milner, J.M. & Irvine, R.J. (2015) The potential for reintroduction of Eurasian lynx to Great Britain: a summary of the evidence, British Deer Society Commissioned Report
(52) Carver, S.J. (1991) Integrating multi-criteria evaluation with geographical information systems, International Journal of Geographical Information Systems, 5:(3): 321-339
(53) GROUND TRUTHING CONNECTIVITY ANALYSIS – SOLENT COAST MAY 2022, Wildland Research Institute
(54) Commodification of nature, Self-willed land January 2021
(55) Colorado Parks and Wildlife Commission, Colorado's Wolf Restoration and Management Plan Public Engagement Website
(56) Summer 2021 Public Engagement, Colorado's Wolf Restoration and Management Plan Public Engagement Website
(57) Summer 2021 Public Engagement Report, Colorado's Wolf Restoration and Management Plan Public Engagement Website
(58) Advisory Groups, Colorado's Wolf Restoration and Management Plan Public Engagement Website
(59) Colorado Wolf Restoration and Management Plan. Final Report on Wolf Restoration Logistics Recommendations, Technical Working Group (TWG) to Colorado Parks and Wildlife (CPW) November 2021
(60) Colorado Wolf Restoration and Management Plan Stakeholder Advisory Group (SAG) to Colorado Parks and Wildlife (CPW) Meeting Summary, July 27-28, 2022 Fort Lewis College 1000 Rim Dr, Durango, CO 81301
(61) Colorado Wolf Restoration and Management Plan. Summer 2021 Public Engagement Report, Keystone Policy Center November 2021
(62) Carroll, C., Phillips, M. K., Lopez-Gonzalez, C. A., & Schumaker, N. H. (2006). Defining recovery goals and strategies for endangered species: the wolf as a case study. BioScience, 56(1): 25-37.
(63) Ditmer, M. A., Wittemyer, G., Breck, S. W., & Crooks, K. R. (2022). Defining ecological and socially suitable habitat for the reintroduction of an apex predator. Global Ecology and Conservation, 38, e02192
(64) WOLF WOOD, Longleat
(65) Visit Us, the Wolf Watch Centre
(66) Feed a Wolf experience, Paradise Wildlife Park