Effects of risk assessment, predator behavior, and habitat on escape behavior in Columbian black-tailed deer

The relationship between preflight risk assessment by prey and the escape behaviors they perform while fleeing from predators is relatively unexplored. To examine this relationship, a human observer approached groups of Columbian black-tailed deer (Odocoileus hemionus columbianus), varying his behavior to simulate more or less threatening behavior. We measured the focal deer's angle of escape, distance moved during flight, duration of trotting and stotting behavior, and change in elevation during flight. Analyses revealed positive relationships between the distance moved during flight and the distance at which they fled. When flight was initiated when the approacher was close, deer fled relatively shorter distances and took flight paths at more acute angles, a property that would force a real predator to change direction suddenly. Our results indicate that deer do not compensate for allowing the observer to approach more closely by fleeing greater distances. Rather, distance moved and flight initiation distance are linked by level of reactivity and habituation: more reactive or less habituated deer both flee at a greater distance and move away to a greater distance during flight. More threatening behavior by the approacher led to longer durations of rapid flight behavior (e.g., trotting and stotting), and deer tended to flee uphill and into taller vegetation, using these landscape features as refuge from danger. Finally, we provide the first evidence for Pitcher's untested "antiambush" hypothesis for the function of stotting and discuss its significance. In general, both preflight predator behavior and habitat features influence both duration and direction of escape. Copyright 2007, Oxford University Press.

[1]  D. Kramer,et al.  Direction of predator approach and the decision to flee to a refuge , 1997, Animal Behaviour.

[2]  T. Caro,et al.  The functions of stotting in Thomson's gazelles: some tests of the predictions , 1986, Animal Behaviour.

[3]  F. J. Singer,et al.  Responses of Desert Bighorn Sheep to Increased Human Recreation , 2001 .

[4]  O. C. Wallmo Mule and black-tailed deer of North America. , 1982 .

[5]  R. Dasmann,et al.  Behavior of Columbian Black-Tailed Deer with Reference to Population Ecology , 1956 .

[6]  Richard G. Coss,et al.  Effects of predator behavior and proximity on risk assessment by Columbian black-tailed deer , 2006 .

[7]  Scott G. Miller,et al.  Wildlife responses to pedestrians and dogs. , 2001 .

[8]  J. Hamr Disturbance behaviour of chamois in an Alpine tourist area of Austria. , 1988 .

[9]  D. J. Freddy,et al.  RESPONSES OF MULE DEER TO DISTURBANCE BY PERSONS AFOOT AND SNOWMOBILES , 1986 .

[10]  F. James Rohlf,et al.  Biometry: The Principles and Practice of Statistics in Biological Research , 1969 .

[11]  W. Cooper EFFECT OF TEMPERATURE ON ESCAPE BEHAVIOUR BY AN ECTOTHERMIC VERTEBRATE, THE KEELED EARLESS LIZARD (HOLBROOKIA PROPINQUA) , 2000 .

[12]  S. Bratton Boat Disturbance of Ciconiiformes in Georgia Estuaries , 1990 .

[13]  Janice C. Daniel,et al.  Locomotor Ability and Wariness in Yellow‐Bellied Marmots , 2004 .

[14]  W. Cooper When and how do predator starting distances affect flight initiation distances , 2005 .

[15]  E. Curio,et al.  Proximate and Developmental Aspects of Antipredator Behavior , 1993 .

[16]  G. Helfman Threat-sensitive predator avoidance in damselfish-trumpetfish interactions , 2004, Behavioral Ecology and Sociobiology.

[17]  D. F. Behrend,et al.  SUMMER FLIGHT BEHAVIOR OF WHITE-TAILED DEER IN TWO ADIRONDACK FORESTS1 , 1968 .

[18]  William E. Cooper,et al.  ESCAPE BY A REFUGING PREY, THE BROAD-HEADED SKINK (EUMECES LATICEPS) , 1997 .

[19]  R. L. Knight,et al.  WILDLIFE RESPONSES TO RECREATION AND ASSOCIATED VISITOR PERCEPTIONS , 2003 .

[20]  Susan Lingle ESCAPE GAITS OF WHITE-TAILED DEER, MULE DEER AND THEIR HYBRIDS: GAITS OBSERVED AND PATTERNS OF LIMB COORDINATION , 1992 .

[21]  R. G. Coss,et al.  PERCEPTUAL ASPECTS OF LEOPARD RECOGNITION BY WILD BONNET MACAQUES (MACACA RADIATA) , 2000 .

[22]  A. Sih,et al.  A framework for determining the fitness consequences of antipredator behavior , 2007 .

[23]  K. S. Smallwood Mountain Lion Vocalizations and Hunting Behavior , 1993 .

[24]  T. Caro,et al.  The functions of stotting: a review of the hypotheses , 1986, Animal Behaviour.

[25]  E. F. Cassirer,et al.  ELK RESPONSES TO DISTURBANCE BY CROSS-COUNTRY SKIERS IN YELLOWSTONE NATIONAL PARK , 1992 .

[26]  D. Blumstein,et al.  Relationships of anti-predator escape and post-escape responses with body mass and morphology: a comparative avian study , 2006 .

[27]  Theodore Stankowich,et al.  The re-emergence of felid camouflage with the decay of predator recognition in deer under relaxed selection , 2007, Proceedings of the Royal Society B: Biological Sciences.

[28]  D. Blumstein MULTIVARIATE ANALYSIS OF GOLDEN MARMOT MAXIMUM RUNNING SPEED: A NEW METHOD TO STUDY MRS IN THE FIELD' , 1992 .

[29]  T. Caro,et al.  Antipredator Defenses in Birds and Mammals , 2006 .

[30]  S. Blamires Factors influencing the escape response of an arboreal agamid lizard of tropical Australia (Lophognathus temporalis) in an urban environment , 1999 .

[31]  Christina M. Payne,et al.  Recognition systems and biological invasions , 2004 .

[32]  R. Swaisgood,et al.  Assessment of rattlesnake dangerousness by California ground squirrels: exploitation of cues from rattling sounds , 1999, Animal Behaviour.

[33]  R. Andersen,et al.  Short term behavioural and physiological response of moose Alces alces to military disturbance in Norway , 1996 .

[34]  J. Linsdale,et al.  A Herd of Mule Deer , 1954 .

[35]  J. Garner,et al.  Is fearfulness a trait that can be measured with behavioural tests? A validation of four fear tests for Japanese quail , 2006, Animal Behaviour.

[36]  M. Hildebrand Analysis of Asymmetrical Gaits , 1977 .

[37]  He Who Hesitates, Lives. Is Stotting Antiambush Behavior? , 1979, The American Naturalist.

[38]  Susan Lingle COYOTE PREDATION AND HABITAT SEGREGATION OF WHITE-TAILED DEER AND MULE DEER , 2002 .

[39]  D. Bauwens,et al.  Escape Tactics and Vulnerability to Predation Associated with Reproduction in the Lizard Lacerta vivipara , 1981 .

[40]  R. Swaisgood,et al.  Conflict and assessment in a predator–prey system: ground squirrels versus rattlesnakes , 1999, Animal Behaviour.

[41]  G. Kass-simon,et al.  Molt-related and size-dependent differences in the escape response and post-threat behavior of the American lobster, Homarus americanus. , 2000, The Biological bulletin.

[42]  Michael H. Kutner Applied Linear Statistical Models , 1974 .

[43]  T. Caro,et al.  Adaptive significance of antipredator behaviour in artiodactyls , 2004, Animal Behaviour.

[44]  S. Pellis,et al.  Fight or flight? Antipredator behavior and the escalation of coyote encounters with deer , 2002, Oecologia.

[45]  Johan Lind,et al.  Determining the fitness consequences of antipredation behavior , 2005 .

[46]  Susan Lingle,et al.  Detection and Avoidance of Predators in White-Tailed Deer (Odocoileus virginianus) and Mule Deer (O. hemionus) , 2001 .

[47]  José Martín,et al.  Changes in the Escape Responses of the Lizard Acanthodactylus erythrurus under Persistent Predatory Attacks , 2003, Copeia.

[48]  D. Blumstein Flight-initiation distance in birds is dependent on intruder starting distance , 2003 .

[49]  J. Losos Thermoregulatory correlates of escape behavior by a desert lizard, Ctenophorus isolepis , 1988 .

[50]  A. Sih,et al.  Trait compensation and cospecialization in a freshwater snail: size, shape and antipredator behaviour , 1999, Animal Behaviour.

[51]  T. Stankowich Marginal predation methodologies and the importance of predator preferences , 2003, Animal Behaviour.

[52]  Donald H. Owings,et al.  Different Functions of "Alarm" Calling for Different Time Scales: a Preliminary Report On Ground Squirrels , 1986 .

[53]  V. Barnett,et al.  Applied Linear Statistical Models , 1975 .

[54]  Sokal Rr,et al.  Biometry: the principles and practice of statistics in biological research 2nd edition. , 1981 .

[55]  C. Dwyer How has the risk of predation shaped the behavioural responses of sheep to fear and distress? , 2004, Animal Welfare.

[56]  Patrick A. Zollner,et al.  Inter‐specific variation in avian responses to human disturbance , 2005 .

[57]  S. L. Lima,et al.  Behavioral decisions made under the risk of predation: a review and prospectus , 1990 .

[58]  W. Cooper,et al.  Magnitude of food reward affects escape behavior and acceptable risk in Balearic lizards, Podarcis lilfordi , 2006 .

[59]  R. Shine,et al.  Costs of reproduction in lizards: escape tactics and susceptibility to predation , 1992, Behavioral Ecology and Sociobiology.

[60]  P. Aastrup,et al.  Responses of West Greenland caribou to the approach of humans on foot , 2000 .

[61]  Lawrence M. Dill,et al.  The Economics of Fleeing from Predators , 1986 .

[62]  K. D. Roeder,et al.  The detection and evasion of bats by moths , 1961 .

[63]  Daniel T Blumstein,et al.  Fear in animals: a meta-analysis and review of risk assessment , 2005, Proceedings of the Royal Society B: Biological Sciences.

[64]  Donald H. Owings,et al.  Snake Mobbing By California Ground Squirrels: Adaptive Variation and Ontogeny , 1977 .