Effects of Human Disturbance on Terrestrial Apex Predators

The effects of human disturbance spread over virtually all ecosystems and ecological communities on Earth. In this review, we focus on the effects of human disturbance on terrestrial apex predators. We summarize their ecological role in nature and how they respond to different sources of human disturbance. Apex predators control their prey and smaller predators numerically and via behavioral changes to avoid predation risk, which in turn can affect lower trophic levels. Crucially, reducing population numbers and triggering behavioral responses are also the effects that human disturbance causes to apex predators, which may in turn influence their ecological role. Some populations continue to be at the brink of extinction, but others are partially recovering former ranges, via natural recolonization and through reintroductions. Carnivore recovery is both good news for conservation and a challenge for management, particularly when recovery occurs in human-dominated landscapes. Therefore, we conclude by discussing several management considerations that, adapted to local contexts, may favor the recovery of apex predator populations and their ecological functions in nature.

[1]  M. Drouilly,et al.  Interactions between livestock guarding dogs and wolves in the southern French Alps , 2020, Journal of Vertebrate Biology.

[2]  H. Sand,et al.  Interspecific Interactions between Brown Bears, Ungulates, and Other Large Carnivores , 2020 .

[3]  J. Andrew Royle,et al.  Estimating and forecasting spatial population dynamics of apex predators using transnational genetic monitoring , 2020, Proceedings of the National Academy of Sciences.

[4]  Sean M. Johnson-Bice,et al.  Outsized effect of predation: Wolves alter wetland creation and recolonization by killing ecosystem engineers , 2020, Science Advances.

[5]  I. Kojola,et al.  Wolf predation risk and moose movement in eastern Finland , 2020, Wildlife Biology.

[6]  U. Schraml,et al.  Human-Large Carnivores Co-existence in Europe – A Comparative Stakeholder Network Analysis , 2020, Frontiers in Ecology and Evolution.

[7]  T. Williams,et al.  Hunters versus hunted: New perspectives on the energetic costs of survival at the top of the food chain , 2020, Functional Ecology.

[8]  N. Kingston,et al.  Area-based conservation in the twenty-first century , 2020, Nature.

[9]  M. Waltert,et al.  Variation and conservation implications of the effectiveness of anti-bear interventions , 2020, Scientific Reports.

[10]  H. Sand,et al.  Individual Variation in Predatory Behavior, Scavenging and Seasonal Prey Availability as Potential Drivers of Coexistence between Wolves and Bears , 2020 .

[11]  C. Krebs,et al.  Climate change increases predation risk for a keystone species of the boreal forest , 2020, Nature Climate Change.

[12]  H. Sand,et al.  Promoting grazing or rewilding initiatives against rural exodus? The return of the wolf and other large carnivores must be considered , 2020, Environmental Conservation.

[13]  M. Huijbregts,et al.  Mechanistic insights into the role of large carnivores for ecosystem structure and functioning , 2020, Ecography.

[14]  Bradley P. Smith,et al.  Co-existing with dingoes: Challenges and solutions to implementing non-lethal management , 2020 .

[15]  Sheng Li,et al.  Retreat of large carnivores across the giant panda distribution range , 2020, Nature Ecology & Evolution.

[16]  Beer Martin,et al.  African swine fever - A review of current knowledge. , 2020, Virus research.

[17]  M. Heurich,et al.  Landscape predictors of human–leopard conflicts within multi-use areas of the Himalayan region , 2020, Scientific Reports.

[18]  A. Ford,et al.  The ecology of human–carnivore coexistence , 2020, Proceedings of the National Academy of Sciences.

[19]  M. Apollonio,et al.  Relative impact of human harvest and wolf predation on two ungulate species in Central Italy , 2020 .

[20]  H. Sand,et al.  Wolf habitat selection when sympatric or allopatric with brown bears in Scandinavia , 2020, Scientific Reports.

[21]  J. Estes,et al.  Costs and benefits of living with predators , 2020, Science.

[22]  J. Fisher,et al.  Move to nocturnality not a universal trend in carnivore species on disturbed landscapes , 2020, Oikos.

[23]  Timothy P. Duane,et al.  Gray wolf habitat use in response to visitor activity along roadways in Yellowstone National Park , 2020, Ecosphere.

[24]  V. Penteriani,et al.  Large carnivores living alongside humans: Brown bears in human-modified landscapes , 2020, Global Ecology and Conservation.

[25]  J. Belant,et al.  Mortality of a large wide-ranging mammal largely caused by anthropogenic activities , 2020, Scientific Reports.

[26]  T. Ruf,et al.  How climate change and wildlife management affect population structure in wild boars , 2020, Scientific Reports.

[27]  Michel T. Kohl,et al.  How climate impacts the composition of wolf‐killed elk in northern Yellowstone National Park , 2020, The Journal of animal ecology.

[28]  L. Boitani,et al.  Anthropogenic food subsidies hinder the ecological role of wolves: Insights for conservation of apex predators in human-modified landscapes , 2020, Global Ecology and Conservation.

[29]  Kaitlyn M. Gaynor,et al.  Ecological impacts of human-induced animal behaviour change. , 2020, Ecology letters.

[30]  A. Flykt,et al.  Believed effect - A prerequisite but not a guarantee for acceptance of carnivore management interventions , 2020, Biological Conservation.

[31]  Erlend B. Nilsen,et al.  Harvest models of small populations of a large carnivore using Bayesian forecasting , 2019, Ecological applications : a publication of the Ecological Society of America.

[32]  S. Creel,et al.  Carnivores, competition and genetic connectivity in the Anthropocene , 2019, Scientific Reports.

[33]  T. Masaki,et al.  Downhill seed dispersal by temperate mammals: a potential threat to plant escape from global warming , 2019, Scientific Reports.

[34]  J. Swenson,et al.  Endozoochory by brown bears stimulates germination in bilberry , 2019, Wildlife Biology.

[35]  R. Kansky,et al.  Human-carnivore relations: A systematic review , 2019, Biological Conservation.

[36]  L. Boitani,et al.  High levels of recent wolf × dog introgressive hybridization in agricultural landscapes of central Italy , 2019, European Journal of Wildlife Research.

[37]  P. Beja,et al.  Designing the landscape of coexistence: Integrating risk avoidance, habitat selection and functional connectivity to inform large carnivore conservation , 2019, Biological Conservation.

[38]  L. Boitani,et al.  Unravelling the Scientific Debate on How to Address Wolf-Dog Hybridization in Europe , 2019, Front. Ecol. Evol..

[39]  J. M. Rey Benayas,et al.  Rewilding complex ecosystems , 2019, Science.

[40]  J. Swenson,et al.  Habituation, sensitization, or consistent behavioral responses? Brown bear responses after repeated approaches by humans on foot , 2019, Biological Conservation.

[41]  G. Mowat,et al.  Mechanisms influencing the winter distribution of wolverine Gulo gulo luscus in the southern Columbia Mountains, Canada , 2019, Wildlife Biology.

[42]  J. Goheen,et al.  Antipredator response diminishes during periods of resource deficit for a large herbivore. , 2019, Ecology.

[43]  Joseph D. Holbrook,et al.  Wolverines in winter: indirect habitat loss and functional responses to backcountry recreation , 2019, Ecosphere.

[44]  E. Revilla,et al.  Patterns of brown bear damages on apiaries and management recommendations in the Cantabrian Mountains, Spain , 2018, PloS one.

[45]  J. Månsson,et al.  Habitat segregation between brown bears and gray wolves in a human‐dominated landscape , 2018, Ecology and evolution.

[46]  T. Ramakrishna Rao.  Trophic Cascades , 2018, Resonance.

[47]  J. Grinath.  Short‐term, low‐level nitrogen deposition dampens a trophic cascade between bears and plants , 2018, Ecology and evolution.

[48]  G. Stenhouse,et al.  Grizzly bear response to spatio‐temporal variability in human recreational activity , 2018, Journal of Applied Ecology.

[49]  M. Boyce.  Wolves for Yellowstone: dynamics in time and space , 2018, Journal of Mammalogy.

[50]  B. Jędrzejewska,et al.  A meta-analysis of ungulate predation and prey selection by the brown bear Ursus arctos in Eurasia , 2018, Mammal Research.

[51]  Lily M. van Eeden,et al.  Carnivore conservation needs evidence-based livestock protection , 2018, PLoS biology.

[52]  M. Heurich,et al.  Illegal hunting as a major driver of the source-sink dynamics of a reintroduced lynx population in Central Europe , 2018, Biological Conservation.

[53]  K. Jerina,et al.  Evolutionary and ecological traps for brown bears Ursus arctos in human‐modified landscapes , 2018 .

[54]  Justin S. Brashares,et al.  The influence of human disturbance on wildlife nocturnality , 2018, Science.

[55]  Taal Levi,et al.  Salmon-supported bears, seed dispersal, and extensive resource subsidies to granivores , 2018, Ecosphere.

[56]  Lily M. van Eeden,et al.  Managing conflict between large carnivores and livestock , 2018, Conservation biology : the journal of the Society for Conservation Biology.

[57]  P. Ciucci,et al.  Ants as food for Apennine brown bears , 2018 .

[58]  Atle Mysterud,et al.  Regulated hunting re-shapes the life history of brown bears , 2017, Nature Ecology & Evolution.

[59]  Richard Bischof,et al.  Caught in the mesh: roads and their network‐scale impediment to animal movement , 2017 .

[60]  Michel T. Kohl,et al.  Diel predator activity drives a dynamic landscape of fear , 2017, bioRxiv.

[61]  D. Landis,et al.  The role of ants in north temperate grasslands: a review , 2017, Oecologia.

[62]  S. Boutin,et al.  Experimental moose reduction lowers wolf density and stops decline of endangered caribou , 2017, PeerJ.

[63]  W. Ripple,et al.  Range contractions of the world's large carnivores , 2017, Royal Society Open Science.

[64]  C. Wilmers,et al.  Fear of the human ‘super predator’ reduces feeding time in large carnivores , 2017, Proceedings of the Royal Society B: Biological Sciences.

[65]  D. Thornhill,et al.  Adaptive use of nonlethal strategies for minimizing wolf–sheep conflict in Idaho , 2017, Journal of Mammalogy.

[66]  J. Swenson,et al.  Seasonality and human disturbance alter brown bear activity patterns: implications for circumpolar carnivore conservation? , 2017 .

[67]  K. Jerina,et al.  Consequences of brown bear viewing tourism : A review , 2017 .

[68]  H. Sand,et al.  Paws without claws? Ecological effects of large carnivores in anthropogenic landscapes , 2016, Proceedings of the Royal Society B: Biological Sciences.

[69]  David W. Macdonald,et al.  Animal behaviour and its role in carnivore conservation: examples of seven deadly threats , 2016, Animal Behaviour.

[70]  H. Andrén,et al.  National Parks in Northern Sweden as Refuges for Illegal Killing of Large Carnivores , 2016 .

[71]  H. Andrén,et al.  Predation or scavenging? Prey body condition influences decision‐making in a facultative predator, the wolverine , 2016 .

[72]  Jameal F. Samhouri,et al.  Ecosystem context and historical contingency in apex predator recoveries , 2016, Science Advances.

[73]  Anne Tolvanen,et al.  Tourism, biodiversity and protected areas--Review from northern Fennoscandia. , 2016, Journal of environmental management.

[74]  E. Revilla,et al.  Conflict Misleads Large Carnivore Management and Conservation: Brown Bears and Wolves in Spain , 2016, PloS one.

[75]  Anthony P. Clevenger,et al.  Wolverine behavior varies spatially with anthropogenic footprint: implications for conservation and inferences about declines , 2016, Ecology and evolution.

[76]  S. Herrero,et al.  Human behaviour can trigger large carnivore attacks in developed countries , 2016, Scientific Reports.

[77]  W. Ripple,et al.  Hypercarnivorous apex predator could provide ecosystem services by dispersing seeds , 2016, Scientific Reports.

[78]  C. DePerno,et al.  Phantoms of the forest: legacy risk effects of a regionally extinct large carnivore , 2016, Ecology and evolution.

[79]  Bruce G. Marcot,et al.  Impacts of Human Recreation on Brown Bears (Ursus arctos): A Review and New Management Tool , 2016, PloS one.

[80]  S. Creel,et al.  Questionable policy for large carnivore hunting , 2015, Science.

[81]  G. Belovsky,et al.  Natural and experimental tests of trophic cascades: gray wolves and white-tailed deer in a Great Lakes forest , 2015, Oecologia.

[82]  Jon M. Arnemo,et al.  Physiological evidence for a human-induced landscape of fear in brown bears (Ursus arctos) , 2015, Physiology & Behavior.

[83]  J. Månsson,et al.  Wolves, people, and brown bears influence the expansion of the recolonizing wolf population in Scandinavia , 2015 .

[84]  D. Macdonald,et al.  Landscapes of Coexistence for terrestrial carnivores: the ecological consequences of being downgraded from ultimate to penultimate predator by humans , 2015 .

[85]  R. Briers,et al.  Eurasian lynx natal den site and maternal home‐range selection in multi‐use landscapes of Norway , 2015 .

[86]  J. Persson,et al.  Paying for an Endangered Predator Leads to Population Recovery , 2015 .

[87]  C. Darimont,et al.  The unique ecology of human predators , 2015, Science.

[88]  S. Redpath,et al.  When the hunter becomes the hunted , 2015, Science.

[89]  J. Swenson,et al.  Behavioural Differences between Single Scandinavian Brown Bears (Ursus arctos) and Females with Dependent Young When Experimentally Approached by Humans , 2015, PloS one.

[90]  C. Wilmers,et al.  Top carnivores increase their kill rates on prey as a response to human-induced fear , 2015, Proceedings of the Royal Society B: Biological Sciences.

[91]  B. D. Inouye,et al.  Bears benefit plants via a cascade with both antagonistic and mutualistic interactions. , 2015, Ecology letters.

[92]  J. Gilroy,et al.  Carnivore coexistence: value the wilderness. , 2015, Science.

[93]  O. Liberg,et al.  Recovery of large carnivores in Europe’s modern human-dominated landscapes , 2014, Science.

[94]  Douglas W. Smith,et al.  Trophic cascades in a multicausal world: Isle Royale and Yellowstone , 2014 .

[95]  J. Swenson,et al.  Brown bear circadian behavior reveals human environmental encroachment , 2014 .

[96]  M. Hebblewhite,et al.  Status and Ecological Effects of the World’s Largest Carnivores , 2014, Science.

[97]  J. Swenson,et al.  Saving large carnivores, but losing the apex predator? , 2013 .

[98]  Gregory J. McDermid,et al.  Relationships between grizzly bears and human disturbances in a rapidly changing multi-use forest landscape , 2013 .

[99]  O. Liberg,et al.  Sharing the bounty—Adjusting harvest to predator return in the Scandinavian human–wolf–bear–moose system , 2013 .

[100]  Shannon E Albeke,et al.  Linking anti-predator behaviour to prey demography reveals limited risk effects of an actively hunting large carnivore. , 2013, Ecology letters.

[101]  N. Nibbelink,et al.  Recolonizing wolves trigger a trophic cascade in Wisconsin (USA) , 2013 .

[102]  Mark Hebblewhite,et al.  Humans Strengthen Bottom-Up Effects and Weaken Trophic Cascades in a Terrestrial Food Web , 2013, PloS one.

[103]  Solve Sæbø,et al.  Lasting behavioural responses of brown bears to experimental encounters with humans , 2013 .

[104]  W. Ripple,et al.  Widespread mesopredator effects after wolf extirpation , 2013 .

[105]  W. Sutherland,et al.  Understanding and managing conservation conflicts. , 2013, Trends in ecology & evolution.

[106]  K. Eldegard,et al.  From overlooking to concealed: predator avoidance in an apex carnivore , 2012, European Journal of Wildlife Research.

[107]  Pieter A. Zuidema,et al.  Making conservation research more relevant for conservation practitioners , 2012 .

[108]  M. Delibes,et al.  Do bears know they are being hunted , 2012 .

[109]  D. Macdonald,et al.  Environmental determinants of habitat and kill site selection in a large carnivore: scale matters , 2012 .

[110]  M. Mangel,et al.  Deer, predators, and the emergence of Lyme disease , 2012, Proceedings of the National Academy of Sciences.

[111]  J. Swenson,et al.  Behaviour of Solitary Adult Scandinavian Brown Bears (Ursus arctos) when Approached by Humans on Foot , 2012, PloS one.

[112]  H. Andrén,et al.  Influence of intraguild interactions on resource use by wolverines and Eurasian lynx , 2011 .

[113]  M. Hebblewhite,et al.  Human Activity Differentially Redistributes Large Mammals in the Canadian Rockies National Parks , 2011 .

[114]  A. Zedrosser,et al.  Brown bear conservation and the ghost of persecution past , 2011 .

[115]  E. K. Pikitch,et al.  Trophic Downgrading of Planet Earth , 2011, Science.

[116]  Alessandro Massolo,et al.  Human Activity Helps Prey Win the Predator-Prey Space Race , 2011, PloS one.

[117]  M. Delibes,et al.  Predators or prey? Spatio-temporal discrimination of human-derived risk by brown bears , 2011, Oecologia.

[118]  Dror Hawlena,et al.  Predator control of ecosystem nutrient dynamics. , 2010, Ecology letters.

[119]  Lori B. Shelby,et al.  Human Dimensions of Large Carnivore Conservation and Management: Introduction to the Special Issue , 2010 .

[120]  D. Allainé,et al.  Coping with human disturbance: spatial and temporal tactics of the brown bear (Ursus arctos) , 2010 .

[121]  S. Rushton,et al.  Top predators, mesopredators and their prey: interference ecosystems along bioclimatic productivity gradients. , 2010, The Journal of animal ecology.

[122]  M. Genovart,et al.  The Young, the Weak and the Sick: Evidence of Natural Selection by Predation , 2010, PloS one.

[123]  W. Ripple,et al.  The Landscape of Fear: Ecological Implications of Being Afraid~!2009-09-09~!2009-11-16~!2010-02-02~! , 2010 .

[124]  S. Vignieri,et al.  The Northern Rocky Mountain Gray Wolf is Not Yet Recovered , 2009 .

[125]  Arian D. Wallach,et al.  Threatened species indicate hot-spots of top-down regulation , 2009, Animal Biodiversity and Conservation.

[126]  A. Treves.  Hunting for large carnivore conservation , 2009 .

[127]  W. Ripple,et al.  Large predators and trophic cascades in terrestrial ecosystems of the western United States , 2009 .

[128]  A. Laliberte,et al.  The Rise of the Mesopredator , 2009 .

[129]  R. Peterson,et al.  Ungulate Carcasses Perforate Ecological Filters and Create Biogeochemical Hotspots in Forest Herbaceous Layers Allowing Trees a Competitive Advantage , 2009, Ecosystems.

[130]  E. Ritchie,et al.  More than Mere Numbers: The Impact of Lethal Control on the Social Stability of a Top-Order Predator , 2009, PloS one.

[131]  Christopher N. Johnson,et al.  Predator interactions, mesopredator release and biodiversity conservation. , 2009, Ecology letters.

[132]  J. Gaillard,et al.  What shapes Eurasian lynx distribution in human dominated landscapes: selecting prey or avoiding people? , 2009 .

[133]  Olivier Gimenez,et al.  The magnitude and selectivity of natural and multiple anthropogenic mortality causes in hunted brown bears. , 2009, The Journal of animal ecology.

[134]  Peter Segerström,et al.  Human caused mortality in the endangered Scandinavian wolverine population , 2009 .

[135]  A. Mysterud,et al.  Diet shift of a facultative scavenger, the wolverine, following recolonization of wolves. , 2008, The Journal of animal ecology.

[136]  R. May,et al.  Habitat differentiation within the large-carnivore community of Norway's multiple-use landscapes , 2008, The Journal of applied ecology.

[137]  T. Stankowich.  Ungulate flight responses to human disturbance: a review and meta-analysis. , 2008 .

[138]  Richard M Cowling,et al.  Knowing But Not Doing: Selecting Priority Conservation Areas and the Research–Implementation Gap , 2008, Conservation biology : the journal of the Society for Conservation Biology.

[139]  K. Kunkel,et al.  The potential for large carnivores to act as biodiversity surrogates in southern Africa , 2008, Biodiversity and Conservation.

[140]  S. Creel,et al.  Relationships between direct predation and risk effects. , 2008, Trends in ecology & evolution.

[141]  Carlos Nores,et al.  Wild boar Sus scrofa mortality by hunting and wolf Canis lupus predation: an example in northern Spain , 2008 .

[142]  Carrie V. Kappel,et al.  A Global Map of Human Impact on Marine Ecosystems , 2008, Science.

[143]  J. Berger.  Fear, human shields and the redistribution of prey and predators in protected areas , 2007, Biology Letters.

[144]  R. Hilborn,et al.  Predation by Bears Drives Senescence in Natural Populations of Salmon , 2007, PloS one.

[145]  J. Swenson,et al.  Terrain use by an expanding brown bear population in relation to age, recreational resorts and human settlements , 2007 .

[146]  H. Haberl,et al.  Quantifying and mapping the human appropriation of net primary production in earth's terrestrial ecosystems , 2007, Proceedings of the National Academy of Sciences.

[147]  J. Swenson,et al.  Importance of diel behaviour when studying habitat selection : examples from female Scandinavian brown bears (Ursus arctos) , 2007 .

[148]  J. Gill.  Approaches to measuring the effects of human disturbance on birds , 2007 .

[149]  S. Rushton,et al.  Trophic control of mesopredators in terrestrial ecosystems: top-down or bottom-up? , 2007, Ecology letters.

[150]  D. Blumstein,et al.  Evaluating methods to quantify anthropogenic stressors on wild animals , 2007 .

[151]  O. Liberg,et al.  Lynx (Lynx lynx) killing red foxes (Vulpes vulpes) in boreal Sweden – frequency and population effects , 2006 .

[152]  W. Ripple,et al.  Linking a cougar decline, trophic cascade, and catastrophic regime shift in Zion National Park , 2006 .

[153]  G. Vogel,et al.  The Carnivore Comeback , 2006, Science.

[154]  Arlyne Johnson,et al.  Effects of human–carnivore conflict on tiger (Panthera tigris) and prey populations in Lao PDR , 2006 .

[155]  K. Crooks,et al.  Recreation and large mammal activity in an urban nature reserve , 2006 .

[156]  R. May,et al.  Impact of infrastructure on habitat selection of wolverines Gulo gulo , 2006 .

[157]  O. Liberg,et al.  Cross-continental differences in patterns of predation: will naive moose in Scandinavia ever learn? , 2006, Proceedings of the Royal Society B: Biological Sciences.

[158]  K. Berger.  Carnivore‐Livestock Conflicts: Effects of Subsidized Predator Control and Economic Correlates on the Sheep Industry , 2006, Conservation biology : the journal of the Society for Conservation Biology.

[159]  R. Ostfeld,et al.  A call to ecologists: measuring, analyzing, and managing ecosystem services , 2005 .

[160]  M. Hebblewhite,et al.  HUMAN ACTIVITY MEDIATES A TROPHIC CASCADE CAUSED BY WOLVES , 2005, Ecology.

[161]  W. Ripple,et al.  Linking Wolves and Plants: Aldo Leopold on Trophic Cascades , 2005 .

[162]  M. Hebblewhite.  Predation by wolves interacts with the North Pacific Oscillation (NPO) on a western North American elk population , 2005 .

[163]  J. Estes,et al.  Strongly Interacting Species: Conservation Policy, Management, and Ethics , 2005 .

[164]  C. Beale,et al.  Behavioural responses to human disturbance: a matter of choice? , 2004, Animal Behaviour.

[165]  W. Ripple,et al.  Wolves and the Ecology of Fear: Can Predation Risk Structure Ecosystems? , 2004 .

[166]  Stephen Herrero,et al.  Distribution of subadult grizzly bears in relation to human development in the Bow River Watershed, Alberta , 2004 .

[167]  R. Peterson,et al.  The influence of top–down, bottom–up and abiotic factors on the moose (Alces alces) population of Isle Royale , 2004, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[168]  M. Ogada,et al.  Limiting Depredation by African Carnivores: the Role of Livestock Husbandry , 2003 .

[169]  N. Schumaker,et al.  USE OF POPULATION VIABILITY ANALYSIS AND RESERVE SELECTION ALGORITHMS IN REGIONAL CONSERVATION PLANS , 2003 .

[170]  W. Getz,et al.  Trophic facilitation by introduced top predators: grey wolf subsidies to scavengers in Yellowstone National Park , 2003 .

[171]  T. Devault,et al.  Scavenging by vertebrates: behavioral, ecological, and evolutionary perspectives on an important energy transfer pathway in terrestrial ecosystems , 2003 .

[172]  Douglas W. Smith,et al.  Yellowstone after Wolves , 2003 .

[173]  Rolf O. Peterson,et al.  THE EFFECT OF PREY AND PREDATOR DENSITIES ON WOLF PREDATION , 2002 .

[174]  C. Krebs,et al.  Complex numerical responses to top-down and bottom-up processes in vertebrate populations. , 2002, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[175]  A. Frid,et al.  Synthesis Human-caused Disturbance Stimuli as a Form of Predation Risk , 2002 .

[176]  Jörn Theuerkauf,et al.  Daily movements and territory use by radio-collared wolves (Canis lupus) in Bialowieza Primeval Forest in Poland , 2001 .

[177]  J. L. Gittleman,et al.  Predicting extinction risk in declining species , 2000, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[178]  C. Krebs,et al.  Testing hypotheses of trophic level interactions: a boreal forest ecosystem , 2000 .

[179]  J. Swenson,et al.  Bears and ants: myrmecophagy by brown bears in central Scandinavia , 1999 .

[180]  M. Soulé,et al.  Mesopredator release and avifaunal extinctions in a fragmented system , 1999, Nature.

[181]  J. Ginsberg,et al.  Edge effects and the extinction of populations inside protected areas , 1998, Science.

[182]  H. Mooney,et al.  Human Domination of Earth’s Ecosystems , 1997, Renewable Energy.

[183]  W. Weber,et al.  A Global Perspective on Large Carnivore Conservation , 1996 .

[184]  F. Palomares,et al.  Positive Effects on Game Species of Top Predators by Controlling Smaller Predator Populations: An Example with Lynx, Mongooses, and Rabbits , 1995 .

[185]  R. Peterson,et al.  Wolves, Moose, and Tree Rings on Isle Royale , 1994, Science.

[186]  J. Andrew Royle,et al.  Living on the edge: Opportunities for Amur tiger recovery in China , 2018 .

[187]  Taal Levi,et al.  The primacy of bears as seed dispersers in salmon‐bearing ecosystems , 2018 .

[188]  C. Krebs,et al.  Using experimentation to understand the 10‐year snowshoe hare cycle in the boreal forest of North America , 2018, The Journal of animal ecology.

[189]  P. Quenette,et al.  Exploring the potential of brown bear (Ursus arctos arctos) as a long-distance seed disperser: a pilot study in South-Western Europe , 2016 .

[190]  W. Ripple,et al.  The ecological effects of providing resource subsidies to predators , 2015 .

[191]  O. Liberg,et al.  Monitoring of wolves in Scandinavia , 2012 .

[192]  Justina C. Ray,et al.  Large carnivores and the conservation of biodiversity , 2005 .

[193]  Stephen Herrero,et al.  Bears : status survey and conservation action plan , 1999 .

[194]  R. L. Knight,et al.  Understanding wildlife responses to humans , 1998 .

[195]  S. L. Lima.  Nonlethal Effects in the Ecology of Predator-Prey Interactions What are the ecological effects of anti-predator decision-making? , 1998 .