The separation between wolves and humans in modified landscapes
There are State Game Lands around where I lived in Pennsylvania in the early 1980s. I knew that there were hunters among my work colleagues who had licences to take deer that they would portion and freeze. It was disturbing to see hunting rifles so openly racked up in the cabs of their pickup trucks during the hunting season, but I didn’t see the crossbows that some of them used (1). I never really thought about what deer it was that they had been hunting, until I came across an article recently on reinstatement of elk (Cervus canadensis) to Kentucky. Elk once had the largest range of any deer species in North America, but overhunting, grazing competition from domestic livestock, habitat destruction from unrestrained timber harvesting, urbanization, and expansion from the east throughout the nineteenth century caused a westward contraction in its range, and reduced its population to less than 100,000 by the early 1900s (2,3). Of the four subspecies of American elk in North America today, the Rocky Mountain elk is the most abundant (800,000 to 900,000) occurring primarily in the mountain ranges east of the Cascade Mountains, and it is this elk that is used in reinstatement translocations. Elk are the big deer I’ve seen in the National Parks like Yellowstone and Rocky Mountain in America, and Jasper and Banff in Canada, whereas Yosemite only has the smaller mule deer (Odocoileus hemionus) but which I have also seen in those Parks that have elk. I find now that it was the even smaller white-tailed deer (Odocoileus virginianus) that were hunted in Pennsylvania, a deer I have since seen in eastern National Parks like Shenandoah, Rock Creek and Great Smoky Mountains. The most recent State-wide harvest of white-tailed deer in Pennsylvania during 2018-2019 was 375,000, the total for the hunting area where I used to live being about 24,000 ((4) and see WMU5B in (5)). Hard to say what the total population of white-tailed deer is in Pennsylvania, as it seems to be the habit of state game authorities not to bother worrying about total population for a species unless there is a marked fall in harvest numbers that may indicate a drop in that total population and, in the case of ubiquitous white-tailed deer in Pennsylvania, that the hunting harvest is ensuring that state forest growth is not being impacted by the deer.
While there used to be elk across Pennsylvania, with concentrations in the north-central mountains, as well as Pocono Mountains in the east (where I would ski during winter) the species had been extirpated by 1867, as it also became extinct throughout its range in New York state and the north east (6). However, a total of 177 elk were released by the Pennsylvania Game Commission between 1913 and 1926, but it seems they started back hunting too soon - 98 were taken between 1923 to 1931, and an unknown number were also killed illegally for crop damage, this combination of killing driving numbers down so that hunting had to be stopped in 1932. By 1936, there were only 14 elk remaining in the state. A further reintroduction saw numbers rebound to 65 by 1971, and then a three-year trap-and-transfer program launched by the Game Commission in 1998 expanded the elk's range from 350 to 800 square miles, leading to a population 100 years after restoration efforts began of about 950 animals, a small number of which are allowed to be taken by hunters. Elk also roamed the southern Appalachians in the early 1800’s, but were lost to hunting until the Great Smoky Mountains National Park that straddles North Carolina and Tennessee began a five-year experimental release program in 2001 (the year I visited the Park) to determine if elk could be successfully reintroduced, and which has led to a small herd in the Cataloochee Valley area in the south-eastern section of the Park (7).
Filling in a gap in trophic structure
What I find interesting about reinstatement of elk to the eastern state of Kentucky, the state above Tennessee, is that it was not solely predicated on reinstating a game species to be hunted, as it most likely was in Pennsylvania, but that it was seen as filling in a gap in trophic structure caused by its extirpation, as was also the most likely reason for reinstatement in the Great Smoky Mountains. David Maehr, an assistant professor of conservation biology in the Department of Forestry at the University of Kentucky, studied elk restoration in Kentucky. He wrote about it in Wild Earth in 2001, and about his involvement in a partnership project between the Rocky Mountain Elk Foundation, the Kentucky Department of Fish and Wildlife Resources, the University of Kentucky, and private landowners (8). At the time, it was nearly halfway to the goal of importing roughly 2,000 elk from wild western populations. He observed that elk restoration could be justified because it added community complexity through returning an interspecific tension (a tension between two different species) by the physical dominance of elk over white-tailed deer, and which led to ecological separation through a behavioural modification that promised to alter the regional distribution of plants and animals, even if only subtly. It was also a different grazing and browsing influence from the primarily forest-dwelling, browsing white-tail deer, whereas elk graze and browse forest edges and are able to consume plants and plant parts that were uneaten or otherwise out of the reach of deer.
Maehr speculated about the other missing members of a large mammal fauna in Kentucky that had recently included wolf (Canis lupus) black bear (Ursus americanus) and cougar (Puma concolor) and pondered, given the ability of elk populations to grow quickly, whether all of the ecological components were in place to facilitate a naturally regulated herd of elk? Could this work of restoration be considered complete without the additional complexities and regulatory potential imparted by large carnivores that regularly kill and consume an animal as large as an elk, and that might limit the ecological changes that could be caused by unchecked and widespread herbivory? Maehr discounted the black bear as a candidate, as it did not exert a selective force on large, sympatric ungulates such as white-tail deer and elk because it was an omnivore, often only opportunistically scavenging meat, and hibernating for up to half of the year. He noted that although the wolf was making a dramatic return to some parts of the western America, he wondered whether it would be the appropriate choice as the primary top-down regulator for restoring evolutionary relations and landscapes, given that they had such an undeservedly bad reputation because of erroneous legend, and because of the length of their absence from the East. He ruled the wolf out as he considered that successful widespread reintroduction was unlikely, at least in the short term, especially since pack-living habits and diurnal tendencies made them an easy target for intolerant humans who would frustrate their successful reinstatement.
Maehr considered that the cougar (otherwise known as mountain lion, puma, catamount, and panther) was fundamentally different from the wolf in terms of its behaviour and its place in folklore, and was held in higher esteem than its carnivorous canid (wolf) and ursid (bear) relatives, but it likely was also because of the more secretive nature of the cougar – “With these characteristics of North American predators in mind, and in view of the ancient cultural animosity directed toward wolves by Europeans and their descendants, the cougar becomes the most logical flagship for rewilding eastern North America. The return of elk to the East is an important but insufficient step toward recreating the community dynamics under which many of our remaining plants and animals evolved. Herbivory without predation will demand increasing attention from managers as forests suffer the consequences of a missing large carnivore”
A potential change in perception of the wolf in America
Other than game species like the overhunted elk, it’s the strongly interactive species like the large carnivores, the cougar and wolf, which have been extirpated from landscapes in America for being an inconvenience to human land use, as is typically seen in many other continents (9). Cougar and wolf were once distributed across America, but suffered persecution and range contraction such that the main distribution of cougar is in western states, while it is considered functionally extinct in eastern states except for the Florida panther (10-12). The wolf was driven out of America by systematic hunting so that it was essentially extinct by 1960, other than a few hundred roaming the deep woods of upper Michigan and Minnesota (13-15). A slow, natural recolonisation started on two fronts over the 1970s and 1980s, wolves expanding their range southward within those two Great Lake states, and by an incursion across the Canadian border into Glacier National Park in Montana (15). The 1990s saw a second incursion over the border, this time into the Cascade Mountains of Washington State, as well as planned reintroductions into Idaho and Wyoming. Range expansion has continued, predominantly in the redoubts of Washington, Oregon, Montana, Idaho and Wyoming in the west, and Minnesota, Wisconsin and Michigan in the Great Lakes Area (14,15). However, this is still only 10 percent of their historic range in America, and which represents only 30 percent of currently suitable habitat arrived at through modelling (16,17).
It is a pity that David Maehr didn’t live long enough to see recent evidence that points to a potential change in perception of the wolf in America, as well as encouraging news on the spread of Florida panther - he died aged 52 in a light airplane crash in 2008 (18). Maehr was long associated with studying the Florida panther (19) albeit his assertions about its habitat selection were contested (20). However, I am sure he would have been delighted with camera trap evidence in 2016 confirming that a female panther had crossed the Caloosahatchee River for the first time in 40 years, a significant migration barrier that had confined the panther population to southern Florida (21). This female panther must have joined up with one of the males that were known to routinely cross the river, because further camera trap evidence shows her with two kittens, giving hope that panther will naturally expand further north in Florida (22,23).
As to the wolf, evidence has arisen from video camera footage of surprising behaviour in the Greater Voyageurs Ecosystem that encompasses the Voyageurs National Park in northern Minnesota (24). The Voyageurs Wolf Project, a collaboration between the University of Minnesota and Voyageurs National Park, was started to address one of the biggest knowledge gaps in wolf ecology—what do wolves do during the summer?(25). It was known that wolves hunt as packs throughout the winter, killing large prey such as moose and deer. However, once wolf pups are born in the spring they became more solitary predators, switching to smaller mammals such as beaver and deer fawns, and returning to and from the den or rendezvous site between hunting bouts. It was that summer predation behaviour that was poorly understood, and which was difficult to observe due to seasonal growth of dense vegetation. Moreover, kill sites were less obvious as wolves can almost wholly consume beavers and fawns in 20-40 min leaving little evidence of the kill except for hair, a few bone fragments, and maybe stomach contents. This is in contrast to the much greater evidence of hiatus and remains that are left after wolves bring down a moose or deer during winter. Thus the Project had to develop more rigorous search methods for kill sites that entailed fitting wolves with GPS-collars and intensively searching clusters of GPS-locations where a collared wolf remained stationary for more than 20 minutes. One of these cluster locations turned out to be where a couple of wolves from the Bowman Bay Pack spent time next to Irwin Creek over April and May in 2017 (26). When a researcher went to check it out, he observed over 30 minutes one of the wolves hunting and consuming fish. In May the following year, a remote camera trap caught the first-ever recorded video footage of a wolf hunting and killing freshwater fish (27).
If that was not surprising enough, another cluster location identified for a collared wolf in the Moose River Pack in 2017 threw up even more startling behaviour (28). As a researcher approached the area in a drained beaver meadow approximately 1km south of Voyageurs National Park, he expected it to be a kill site, but instead observed five pups gathering around an adult wolf. This was a rendezvous site, a place where adult wolves bring food to their pups when they are no longer dependent on using a den. The pups licked up at the adult wolf’s mouth for approximately 30 seconds, after which the adult wolf began to regurgitate food that some of the pups ate directly from the adult wolf’s mouth as it slowly walked around, while other pups followed behind the adult wolf consuming the regurgitated food that fell to the ground. Investigation of the area after the wolves had left showed several small piles of chewed and whole wild blueberries mixed with a foamy liquid, presumably stomach fluid, on top of the matted grass - the nearest area of berry forage was approximately 360m away. There were no prey remains (animal hair or bone fragments) present in any of the regurgitated material. It took two years before the Project finally got footage in 2019 of wolves eating blueberries (29). It begs the question of how often wolves feed pups with blueberries, and does this improve the survival of the pups? It is known that by mid-to-late summer deer fawns are quick enough to evade wolves, and this period of reduced prey vulnerability coincides with a period of abundant wild berries (28).
Coloradans get to vote on wolf reinstatement
Surely these behaviours should prompt a reassessment of the villainous reputation ascribed to wolves? Fortunately, it did not need this evidence to persuade people in Colorado that now is the time to seriously consider reinstatement of wolves. A petition process exists within the state system to initiate an addition to the state statutes that, if it receives sufficient signatories, is put to a ballot of the people of Colorado (30). Those pursuing wolf restoration to Colorado saw this as a way to incorporate a clause in the state’s revised statutes that would require the wolf to be reinstated. The auguries were good. Just over a year ago, 900 registered voters across the state who were likely to vote in the 2020 election, and vote on ballot proposals, were interviewed on the presumption that a vote would take place for restoring wolves to western Colorado (31). Two-thirds (67%) favoured wolf restoration, with 39% strongly in favour. Just 15% opposed the idea (only 8% strongly) and 18% said they had no opinion. Support was broadly based and widespread, encompassing both urban and rural (both Western Slope and Eastern Plain) and included ranchers, farmers and hunters. Going further, the interviewees were presented with specific proposals that could appear on the ballot proposal, such as the requirement to hold state-wide hearings; develop a science-based plan for restoration; and reintroduce the grey wolf to the west of the continental divide in Colorado by December 31, 2023. In response, 78% said they would vote yes, and in a another version where compensation for injury to livestock caused by wolf restoration was added, 75% would vote yes, rising to 84% in Western Slope region where the reinstatement would take place. Interestingly, Colorado voters were nearly four times more likely to say they would vote for their State Senator or Representative if he or she backed wolf restoration.
A final wording of a proposed statute was accepted for circulation last June, after which signatories were sought to get it above the threshold of 124,632 valid signatures needed for it to succeed onto the ballot (32). It received 215,370 signatories by last December within the six-month time limit, of which 139,333 were deemed to be valid, and so it was confirmed in early January that it would be put to a vote of the people of Colorado this November when congressional elections take place. If passed, it will add a follow-on section specifically about wolves to the Colorado Revised Statute that deals with reintroduction of species (33). It will require the Colorado Parks and Wildlife Commission to create and carry out a plan to reintroduce and manage gray wolves on Colorado lands west of the continental divide (the Western Slope) by the end of 2023, and would direct the state legislature to make appropriations to fund the reintroduction program (34). The Commission would also be required to assist owners of livestock in preventing and resolving conflicts with gray wolves and livestock, and pay compensation to owners of livestock for any losses that could be directly attributed to wolves. Overall, the wording of the proposed statute addition is exemplary in that it recognises restoration of wolves will help restore a critical balance in nature. It also urges using the best scientific data available in formulating the restoration plan; tasks the Colorado Parks and Wildlife Commission to hold state-wide hearings to acquire information to be considered in developing such a plan, including scientific, economic, and social considerations; and to periodically obtain public input to update the plan. As to specifics, it notes that the Commission’s Plan must comply with the requirements of the existing statue on reintroduction of species (33) and must include selection of donor populations of gray wolves; the timing and locations west of the continental divide in Colorado for reintroductions; and the actions necessary for establishing and maintaining a self-sustaining population (34).
Last August, while signatures were being sought for the petition, another state-wide poll took place, conducted by Colorado State University and survey company Qualtrics, the results being released a couple of weeks after it was confirmed in January that the initiative had been placed on the ballot (35). It was found that 84% of Coloradans would vote in favour of gray wolf reintroduction with 16% voting against. Voting intentions were similar across the different regions of Colorado: 84.9% of sampled respondents in the Front Range, 79.8% on the Western Slope, and 79.3% on the Eastern Plains would vote yes. The proportion that would vote yes was also similar among residents in cities, towns, or rural areas. There was 58% support for compensation of landowners for loss of livestock caused by a wolf using state hunting and fishing license income, but this dropped to 42% if the compensation came from state tax revenue. It’s worth reading the open-ended responses received when asked whether wolves would have a positive impact on their life. These were grouped under a range of themes, from a balancing the ecosystem; the opportunity to observe wolves; having an emotional connection; that there was a moral argument for reinstatement; that they would control other animals that come into conflict with humans; that they had an existence value; that they would increase wildlife diversity; that wolf reintroduction would increase environmental learning and care; and that wolf reintroduction would enhance Colorado State Pride – “It would increase the wildness of Colorado”. You can read the responses to negative reactions for yourself, hackneyed as they are.
Barriers to voluntary wolf migration into Colorado
I keep an interest in wolves and their prospects in America since I first saw them in Yellowstone National Park in 2008 (36). I first heard about the process in Colorado through a newsletter I had signed up to from the Rocky Mountain Wolf Action Fund that led the support campaign for the initiative, eventually delivering enough signatories to the petition to get it on the ballot (37). It’s a sister project to the Rocky Mountain Wolf Project (38). The recognition of the importance of wolves in the trophic ecology of Colorado, as expressed by the Rocky Mountain Wolf Project (39-42) had certainly moved on from when I wrote a few years ago about the deliberations on how to reduce the impact of elk on aspen in the Rocky Mountain National Park, the Park authorities ruling in favour of culling rather than reintroduction of wolves (43). It was my understanding then that the outlook for voluntary reinstatement in to Colorado was uncertain, and it still seems to be the case because there are two significant barriers that largely prevent wolf range expansion from Wyoming in to Colorado. John Murtaugh of Defenders of Wildlife, which assisted in the delivery of signatures, recently explained that a “Predator Zone” exists throughout 85% of Wyoming (see the map in (44)) an area designated by state legislature in which there are no regulations on the hunting of wolves so that they can be killed in any number, at any time, and by any means (45,46). This arose after the U.S. Court of Appeals reversed a U.S. District Court decision and issued a final mandate in 2017 that took away their protection by delisting wolves as an Endangered Species in Wyoming, and which gave back management authority to the State (47). Thus according to the 2018 Wyoming Wolf Report – “Wolves could also be taken in any legal manner in Wyoming where they are designated as predatory animals. Fortytwo wolves were taken by the public under predatory animal status in 2018”(46). Furthermore, Murtaugh noted that the southern portion of Wyoming is dominated by the Red Desert, over 9,000 square miles of arid desert that sits on the Wyoming/Colorado border, and which is an inhospitable habitat for the wolves to cross (44). I can testify to that landscape, having driven from Grand Teton National Park in Wyoming down to Fort Collins in Colorado. It thus makes sense to bridge that gap by jumping over it through reintroducing the wolves into Colorado.
I would also like to point out that a ready source of wolves for translocation could have been an alternative to the mindlessly stupid mass cull of over 800 wolves in west-central Alberta, Canada, just to see if that would cause a rebound in falling numbers of woodland caribou – it didn’t (48,49). That was back in 2005 to 2011, but the stupidity persists with a plan this year by the provincial government to cull 90 wolves in Chilcotin Country in British Columbia, Canada, to supposedly prevent the number of caribou from dropping further (50). Perhaps the decline in number since 2003 might have been less if the licensed hunt for caribou had been stopped earlier than last year.
The Colorado Parks and Wildlife Commission does have a successful track record in reinstating a native species, the lynx (Lynx canadensis) to the southern portion of their former range, having been lost to Colorado by 1974. Starting in 1999, 41 lynx were brought from Canada and Alaska and released into the San Juan Mountains in southwest Colorado (51,52). A total of 218 lynx were brought to the area over the years to 2006, each of which were fitted with radio and satellite collars, allowing researchers to monitor movement patterns, survival, male and female proximity during breeding season, and female denning and births. In 2008, I was staying in Dillon in Summit County, as a base to walk Colorado wilderness, and I remember reading a local newspaper account about concerns at a shortage of snow-shoe hares (Lepus americanus) that had led to a failure of lynx to give birth to any kittens in both 2007 and 2008 (53). I was relieved to later see from an annual report that reproduction had resumed in 2009 and 2010 (54). By the summer of 2010, all benchmarks established prior to the start of the project for successful lynx reintroduction had been met, with lynx established as a self-sustaining population (51,52).
The location data collected from the collared lynx by the Colorado Division of Wildlife from 1999 to 2010 enabled an analysis of habitat use by the lynx (55). The analysis provided a range of landscape variables that showed a broad preference for high elevation wet spruce/fir and mixed spruce/fir forests, but avoidance of lower elevation, drier montane forests and montane shrublands, but they may use the latter infrequently as corridors for migration. Thus it’s not surprising that the majority of lynx use areas were located on National Forest lands, with the two largest contiguous use areas being in the San Juan Mountain range, and the Collegiate Peaks ranging from Monarch Pass up to Vail Pass. I walked two wilderness in the Collegiate Peaks in 2008 - Hunter-Fryingpan Wilderness and Collegiate Peaks Wilderness – but I don’t think I was at a high enough elevation, even if there was a chance of seeing lynx. What this analysis has made possible is the production of a map of predicted lynx use on a statewide scale, and which identified areas within Colorado that should contain high quality lynx habitat (56). If suitable corridors for migration exist, then there is potential for substantial range expansion within Colorado.
A spoiler to the ballot initiative
It would seem perverse to mention the opposition to reintroduction of the wolf when it has been couched in such hyperbolic terms by the Colorado Stop the Wolf Coalition – “Extreme activist groups want to force introduction of destructive non-native wolves onto public lands in Colorado” (57,58). The Coalition can, of course, make its own case to the Coloradan electorate. However, within a couple of weeks of the initiative making the ballot, a Colorado state Senator introduced a Senate Bill that seeks to wrest the decision process on reintroduction away from the people, the Senator claiming that the state legislature would give a forum for a discussion of a complex issue for the Western Slope – “I don’t think that happens with two opposing campaigns, which does not always provide the right forum for the discussions and compromises that these kinds of complicated issues require” (59). It’s a spoiler, a side-step around the ballot initiative, as well as being patronising to the voters. The proposed bill authorizes the management and, if necessary (see later for why that qualifier is there) the reintroduction of the gray wolf into suitable habitat in Colorado so that it can develop a self-sustaining population in accordance with a science-based plan adopted by the Colorado Parks and Wildlife Commission, the latter financing programs to educate livestock owners on avoidance and mitigation of damages potentially caused by gray wolves. (60). However, the reintroduction is to begin by 31 December 2025, a start date that is much later than the completion date of the ballot initiative (see above) and which can be further postponed until a new source of revenue is identified to pay for damages caused by gray wolves. On that, a study group would be convened to consider how to verify and estimate damages caused by gray wolves, as well as identifying one or more of those new sources of revenue to pay the damages.
The reintroduction can also be cancelled if the gray wolf achieves in the meantime a self-sustaining population in Colorado (60). The Senator would have been aware that there had been sightings in early January of a pack of six wolves around an elk kill in Moffat County in the far north-western corner of Colorado (61-63) later DNA testing of scat samples taken near an elk carcass confirming they came from wolves (64). These wolves must have sneaked down past the outside western edge of both the Predator Zone and the Red Desert in Wyoming (see above and Fig. 1 in (46)). This was foreseen as a potential migration route by Defenders of Wildlife in its report Places for Wolves (65). Based on the seven places in America that had wolves, the report showed maps for each that speculated on their movement from the occupied areas via potential dispersal corridors into other nearby ecological regions that had the potential to support thriving wolf populations. The map for the Northern Rockies did show a direct route southward through Wyoming to the Medicine Bow-Routt National Forests on the border with Colorado, but this report was compiled in 2006 and thus before the Predator Zone came into force. The map also showed a dispersal corridor moving southward on the western edge of Wyoming, but into an ecological region in NE Utah first before another corridor linked eastward into Colorado (see map on pg. 17 in (65)). That this migration route was not fanciful was proven when a satellite-collared, year old female wolf of the Mill Creek Pack from an area just north of Yellowstone National Park was tracked moving southward through Wyoming before heading west for a short journey into Idaho, and then on southward into Utah, finally hooking around eastward through Ashley National Forest and moving into Colorado (see the graphic in (66)). The wolf deserved better after her epic journey, being found illegally poisoned in April 2009 with Compound 1080, a substance banned in Colorado. It would, however, be wishful thinking yet to presume that this presaged a migration corridor that would stay open, and which coupled with breeding in Colorado would establish a self-sustaining population. It is interesting to note that Colorado Governor Jared Polis welcomed these wolves to the state – “This is a historic sighting. While lone wolves have visited our state periodically including last fall, this is very likely the first pack to call our state home since the 1930s. I am honored to welcome our canine friends back to Colorado after their long absence” (667). He also warned against their persecution, urging people to give them room – “It’s important that Coloradans understand that the gray wolf is under the protection of the Endangered Species Act. While the animals have naturally migrated to our state and their presence draws public interest, it’s important that people give them space. Due to their Protected status, there are severe federal penalties for anyone that intentionally harms or kills wolves in our state” (and see this protection confirmed (68)).
There has been no progress yet of the Senator’s bill through the Colorado State legislature (69) but I should point out that state legislation does not automatically override voter-approved initiatives, and thus if passed it would rely on the ballot initiative being withdrawn before the vote (59). The Rocky Mountain Wolf Action Fund said it was happy to discuss compensation plans and proactive measures ranchers can take to prevent predation of livestock, but it wouldn’t support the bill, particularly the delays and ambiguity inherent in it about what constituted a sustainable population, so that withdrawal of the ballot initiative is unlikely to happen without major changes in the bill.
The greatest natural experiment in reinstatement of trophic ecology
I’ve gone to great lengths in documenting this process of wolf reintroduction in Colorado, along with the preceding information on species restoration and trophic ecology, as it is just something completely missing in discussion and aspiration here in Britain, let alone the opportunity for its citizens to determine for itself that wolf reinstatement should happen. We don’t even want to learn the lessons from continental Europe where the greatest natural experiment in reinstatement of trophic ecology has taken place by the voluntary westward expansion of wolves so that there is now a presence in every country. It’s a big research question in Europe, but a brief exploration of the literature reveals that no one has yet studied this. Thus a great chance is being missed for a better understanding of how wolves have distributed, how they use landscapes in Europe, and what it says about transcontinental wildlife movement. Knowledge of their land use would get us away from the dead hand of the almost triumphalist but specious and simplistic assertion that the recovery of wolves in Europe’s modern human-dominated landscapes proves that they coexist with people rather than depend on wilderness, an inference that is impossible to sustain from the gross mapping of the study (70). There is some data that can begin to fill this void in knowledge, but which I believe would also give insights into how you could have predicted how wolves moved westward in Europe, and which could then have been tested and verified by using camera traps, DNA sampling in faeces etc. in advance of the leading edge in locations where you might have expected migration to occur.
I’ve written before of the report from 2013 that mapped the potential of wolf recolonisation in Denmark in anticipation of its voluntary return across the border from Germany (71). The evaluation was based on the distribution of sufficiently large enough patches of inter-connected forest and heath, and with those having good populations of Red and roe deer, from which were identified a total of 10 suitable breeding areas in Denmark for packs consisting of about eight wolves, including pups (see Fig 7 in (72)). Interestingly, the official webpage on wolves in Denmark now carries information from Germany about the diet of wolves, showing that livestock make up only 0.6% of the wolf's food choices – “There are very few known examples of wolves attacking larger livestock such as horses, cows and donkeys” (73). I’ve remarked on this and other dietary studies that consistently show a high reliance on wild species, and which suggests to me that wolves certainly seek out wild food in preference to domestic livestock, irrespective of whether the landscape is wilderness or not (74,75).
So what has happened since that evaluation – have the in-migrating wolves followed the predicted distribution? The number of wolves coming over from Germany into Denmark - and not getting killed – are still low (less than 20) although two wolves have had litters of at least eight pups that will disperse from inside Denmark, all these wolves individually confirmed by observation and DNA analyses from wolf faeces (76). There is a status report on wolves from last September based on the national surveillance of wolves in Denmark under the Danish Environmental Protection Agency during the period 1 April -30 June 2019, and which maps the geographical distribution of secure and confirmed, probable, and possible findings of wolf (see Fig 1 in (77)). In addition, there is a rolling dynamic map of wolf observations from 2012 until now, maintained by the Natural History Museum in Aarhus (78). It may be a bit early to make such a comparison because of the low numbers, but the similarities between the predicted location of wolves (see Fig 7 in (72)) and their actual location is quite striking. Two spatial studies were also carried out in the Netherlands in 2012 based on prey availability, habitat suitability, and areas with low human disturbance, again in anticipation of the arrival of wolves (71) but there just aren’t enough wolves there yet to make a comparison.
Spatial and temporal avoidance strategies
Neither the Denmark nor Netherlands study addressed possible routes for wolf dispersal as they recolonised those countries, only predicting where they may end up staying. Two recent studies of Iberian wolves (Spain and Portugal) both predicated on their existence in human dominated landscapes, offer real answers on the choices made by wolves for where they live and the characteristics of the routes through which they travel and disperse. Unfortunately, neither study is freely available, but you can get a good impression of their findings from the abstracts. One study compiled presence/absence data at different spatial scales that characterised wolf presence, habitat selection, and breeding sites, from such as the Portuguese wolf census and the atlas of Spanish mammals, and used that to identify associated environmental variables, such as topography (altitude, slope) water and prey availability, openness of vegetation, and extent of human disturbance (79). Their observation was that refuge availability, as defined by topography, seemed to be the key factor determining wolf presence at the multiple scales analysed. As a result, wolf populations may coexist with humans in modified landscapes when the topography is complex. Based on those environmental variables, they also used modelling to predict across Iberia where range expansion could occur. You can at least see the graphic of the outcome of that modelling, and which shows that a significant amount of favourable habitat is currently unoccupied (80) the authors averring that it was most likely due to direct persecution and other sources of anthropogenic mortality that are hampering its expansion. They suggest that priorities for conservation and coexistence should be protecting good quality habitat within the current range, the refuges away from human disturbance; and by allowing dispersal to unoccupied areas of good quality habitat by reducing human-induced mortality rates.
The other study focussed on wolves in north-western Portugal, using movement data from 15 individuals to model how human-related risks constrain the spatial distribution of habitats, their temporal use, and the functional connectivity in modified landscapes (81). The results showed that wolves avoided potential sources of human disturbance, particularly settlements, roads, trails and windfarms, avoidance being much stronger for resident wolves than dispersers, and avoidance was greater in daytime than in twilight and night. There was strong segregation based on altitude, particularly at night, with resident wolves favouring higher elevations, whereas dispersing wolves would move through lower elevations. Resident wolves were largely restricted to fragmented mountainous areas less used by people, while they faced strong resistance to movement in more densely populated lowlands. There were well-defined dispersal corridors potentially connecting most wolf packs, a greater tolerance to humans facilitating the movement of dispersers through the landscape, though they had the additional constraint of avoiding resident wolves. The authors say that their results reinforce the need to prevent new sources of human disturbance, such roads and wind farms, in the remnant areas used by resident wolves, and to preserve the continuous dispersal corridors through areas heavily used by humans. Their formula for designing landscapes of coexistence between wolves and humans would be ensuring the spatial and temporal segregation of human structures and activities from large carnivore breeding and dispersal habitats.
These fascinating observations are summarised in a visually compelling infographic provided as an addition to the published article (82). Shockingly, this paper hasn’t received much attention since it was published! I find it immensely powerful and urge you to study the infographic for the insights it has to the movement ecology of wolves, and the implications it poses for any area facing up to - but why not say welcoming the return of wolves. It gives the lie to the prejudiced fantasists that want you to believe that wolves will dominate our landscapes – the wolves want to avoid us as much as we may wish to avoid them. Dietary studies also give the lie to assertions that wolves are mindless predators of livestock. If that were not enough, then a recent study from Germany, a country that has seen wolves return and establish over the last decade or so, strongly shows that the majority of people reported positive experiences when encountering wolves, regardless of whether wolves were encountered in the wild within Germany (57%) in the wild abroad (67%) or in captivity (64%)(83). Contrary to their expectations, a higher frequency of encounters with wolves was not associated with more negative feelings, which anyway only ranged between 5-14%, but it was associated with greater tolerance of living in proximity to wolves. The authors concluded that direct personal experiences play an important role in shaping attitudes toward large carnivores, noting that their results highlight the existence and potential importance of positive experiences. They also considered the positive or negative nature of indirect experiences, such as the various sources of information like the print and online media, and how they shaped people's attitudes toward large carnivores, and what that means – especially if there is a disconnect between direct and indirect experiences - in paving the way toward long-term coexistence with large carnivore species.
Given that few people in Britain will have had an encounter with wolves – but maybe they should – then the prevailing attitude that characterises the indirect experiences is monopolised by those that just scream against wolves. Other voices must come forward and be heard. A mature society should be using every piece of scientific evidence to predict how wolves would use the landscapes of Britain, where they would choose to reside, and what routes they would take to disperse. We should then work out what measures must be taken to ensure that the separation that undoubtedly does take place between wolves and humans in modified landscapes is unhindered and works to the benefit of both wolves and humans. Would you not feel that the reinstatement and the presence of wolves here in Britain would more likely be a positive experience if we put some effort into understanding it, if we took that more mature approach? How much support does there have to be to kickstart this, because I think the survey early in January that showed that 36% of people questioned wanted to reintroduce wolves to Britain is enough to build on (84).
Mark Fisher 19, 31 March 2020
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